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The fungal pathogen Batrachochytrium dendrobatidis (Bd) poses a major threat to amphibian populations. To assist efforts to address such threats, we examined differences in Bd host infection prevalence among amphibian species and its relations to both local environmental factors in breeding habitats and landscape variables measured at three scales (500, 2000 and 5000 m radii) around breeding sites in southernmost Sweden. We sampled 947 anurans of six species in 31 ponds and assessed their infection status. We then examined correlations of infection prevalence with canopy cover, pond perimeter and pH (treated as local-scale pond characteristics), and the number of ponds, area of arable land, area of mature forest, number of resident people and presence of sea within the three radii (treated as landscape variables). The Bd infection prevalence was very low, 0.5-1.0%, in two of the six anuran species (Bufo bufo and Rana temporaria), and substantially higher (13-64%) in the other four (Bombina bombina, Bufotes variabilis, Epidalea calamita, Rana arvalis). In the latter four species Bd infection prevalence was positively associated with ponds' pH (site range: 5.3-8.1), and negatively associated with areas of mature forest and/or wetlands in the surroundings. Our results show that the infection dynamics of Bd are complex and associated with host species, local pond characteristics and several landscape variables at larger spatial scales. Knowledge of environmental factors associated with Bd infections and differences in species' susceptibility may help to counter further spread of the disease and guide conservation action plans, especially for the most threatened species.

Batrachochytrium dendrobatidis ( Bd ) is a fungal pathogen that has decimated amphibian populations worldwide for several decades. We examined the changes in gene expression in response to Bd infection in two populations of the common toad, Bufo bufo , in a laboratory experiment. We collected B. bufo eggs in southern and northern Sweden, and infected the laboratory-raised metamorphs with two strains of the global panzoonotic lineage Bd -GPL. Differential expression analysis showed significant differences between infected and control individuals in both liver and skin. The skin samples showed no discernible differences in gene expression between the two strains used, while liver samples were differentiated by strain, with one of the strains eliciting no immune response from infected toads. Immune system genes were overexpressed in skin samples from surviving infected individuals, while in liver samples the pattern was more diffuse. Splitting samples by population revealed a stronger immune response in northern individuals. Differences in transcriptional regulation between populations are particularly relevant to study in Swedish amphibians, which may have experienced varying exposure to Bd . Earlier exposure to this pathogen and subsequent adaptation or selection pressure may contribute to the survival of some populations over others, while standing genetic diversity in different populations may also affect the infection outcome.

Species in the fungal genus Batrachochytrium are responsible for severe declines in the populations of amphibians globally. The sources of these pathogens have been uncertain. O'Hanlon et al. used genomics on a panel of more than 200 isolates to trace the source of the frog pathogen B. dendrobatidis to a hyperdiverse hotspot in the Korean peninsula (see the Perspective by Lips). Over the past century, the trade in amphibian species has accelerated, and now all lineages of B. dendrobatidis occur in traded amphibians; the fungus has become ubiquitous and is diversifying rapidly. Science , this issue p. 621 ; see also p. 604 The chytrid fungus responsible for global amphibian declines originated in the Korean peninsula and spread during the past century by human trade. Globalized infectious diseases are causing species declines worldwide, but their source often remains elusive. We used whole-genome sequencing to solve the spatiotemporal origins of the most devastating panzootic to date, caused by the fungus Batrachochytrium dendrobatidis , a proximate driver of global amphibian declines. We traced the source of B. dendrobatidis to the Korean peninsula, where one lineage, Bd ASIA-1, exhibits the genetic hallmarks of an ancestral population that seeded the panzootic. We date the emergence of this pathogen to the early 20th century, coinciding with the global expansion of commercial trade in amphibians, and we show that intercontinental transmission is ongoing. Our findings point to East Asia as a geographic hotspot for B. dendrobatidis biodiversity and the original source of these lineages that now parasitize amphibians worldwide.

Factors behind intraspecific variation in sensitivity to pathogens remain poorly understood. We investigated how geographical origin in two North European amphibians affects tolerance to infection by the chytrid fungus Batrachochytrium dendrobatidis (Bd), a generalist pathogen which has caused amphibian population declines worldwide. We exposed newly metamorphosed individuals of moor frog Rana arvalis and common toad Bufo bufo from two latitudinal regions to two different BdGPL strains. We measured survival and growth as infections may cause sub-lethal effects in fitness components even in the absence of mortality. Infection loads were higher in B. bufo than in R. arvalis, and smaller individuals had generally higher infection loads. B. bufo had high mortality in response to Bd infection, whereas there was little mortality in R. arvalis. Bd-mediated mortality was size-dependent and high-latitude individuals were smaller leading to high mortality in the northern B. bufo. Bd exposure led to sub-lethal effects in terms of reduced growth suggesting that individuals surviving the infection may have reduced fitness mediated by smaller body size. In both host species, the Swedish Bd strain caused stronger sublethal effects than the British strain. We suggest that high-latitude populations can be more vulnerable to chytrids than those from lower latitudes and discuss the possible mechanisms how body size and host geographical origin contribute to the present results.

Genetic variation is often lower at high latitudes, which may compromise the adaptability and hence survival of organisms. Here we show that genetic variability is negatively correlated with northern latitude in European green toads (Bufotes viridis). The result holds true for both putatively neutral microsatellite variation and supposedly adaptive MHC Class IIB variation. In particular, our findings have bearing on the conservation status of this species in Sweden, on the northern limit of its distribution where local populations are small and fragmented. These genetically impoverished populations are closely related to other populations found around the Baltic Sea basin. The low neutral and adaptive variation in these fringe populations compared to population at central ranges confirms a pattern shared across all other amphibians so far studied. In Sweden, the situation of green toads is of concern as the remaining populations may not have the evolutionary potential to cope with present and future environmental challenges.

Identifying the routes and rates of introductions is fundamental for the understanding of marine invasions. Recurring introductions over the last 50 yr have led to the establishment of feral Pacific oyster Crassostrea gigas populations throughout Europe. In the northern countries, Sweden and Norway, the species first occurred in large numbers in 2006. Here, we investigated the relative importance of introduction via re-laying of cultured oysters imported for consumption from France, Ireland or the Netherlands, and dispersal of oyster larvae by ocean currents from wild oyster populations in Denmark. Using microsatellite DNA markers, we estimated genetic differentiation among Pacific oysters collected at 4 Swedish locations, 3 Norwegian locations and 9 potential source locations in Denmark, Ireland, the Netherlands and France. All Swedish samples and 1 Norwegian sample (Tromlingene) were genetically similar to each other and the Danish samples and showed significant genetic differentiation from all other populations. Consequently, it appears that the Pacific oyster populations in Sweden, Denmark and Tromlingene are closely connected and/or share a recent origin. The 2 remaining Norwegian samples (Hui and Espevik) differed from each other and all other populations, but showed similarities to wild oyster samples from Scandinavia and Ireland, respectively. Overall, the results underline a complex origin of Norwegian oysters, with gene flow from Swedish/Danish populations, as well as other unidentified sources. The apparent connectivity among most of the Scandinavian populations has implications for regional management of this invasive species, and highlights possible scenarios for other marine invasive species with a similar life history.

Emerging infectious diseases are causing mortality and declines in wildlife populations globally. My thesis aims to get as clear a picture as possible of the effect the invasive chytrid fungus Batrachochytrium dendrobatidis has on the Swedish amphibian community.In Paper I I performed a large-scale survey testing for the presence of Bd in three regions in Sweden (Southern, Central and Northern). I sampled 1917 amphibians from 101 localities and found that Bd was widespread in southern and central Sweden, occurring in all nine investigated species and in 45.5 % of the sampled sites with an overall prevalence of 13.8%. I found a positive correlation between the temperature at spawning for each species and species prevalence. Species that require higher temperatures for egg-laying are distributed in the southern parts of the country, which led to a higher prevalence in the southern region.In Paper II, I investigated which local environmental factors in breeding habitats, landscape structure and amphibian community affect the occurrence and prevalence of Bd among breeding sites in southern Sweden. Bd prevalence in the four species with the highest prevalence (Bombina bombina, Bufotes variabilis, Epidalea calamita and Rana arvalis) was higher in ponds surrounded by less mature forest, few wetlands, and higher pH.In Papers III and IV, I looked at species and population differences in responses to Bd infection. I performed an infection experiment described in Paper III, where I exposed individuals from two common Swedish species (moor frog R. arvalis and common toad Bufo bufo) originating from two regions (north and south) with two different strains of Bd (from Sweden and the UK). I found that infection led to lower survival and growth in both species, more so in B. bufo than in R. arvalis. Small size proved to be a strong determinant of survival. As individuals from the northern population were significantly smaller than the southern ones, this may have led to the northern populations being more affected by Bd infection. In Paper IV, I studied variation in MHC Class IIB loci in B. bufo along a latitudinal gradient across Sweden. Variation in MCH genes decreased from south to north. Also, differences in survival from the experiment in Paper III could be explained by MHC haplotypes. I found that survival in the southern region was dependent on both Bd-strain and MHC haplotype.

Summary Life‐history theory predicts a trade‐off between early‐life fitness and life span. While the focus traditionally has been on the fecundity‐life span trade‐off, there are strong reasons to expect trade‐offs with growth rate and/or development time. We investigated the roles of growth rate and development time in the evolution of life span in two independent selection experiments in the outcrossing nematode Caenorhabditis remanei. First, we found that selection under heat‐shock leads to the evolution of increased life span without fecundity costs, but at the cost of slower development. Thereafter, the putative evolutionary links between development time, growth rate, fecundity, heat‐shock resistance and life span were independently assessed in the second experiment by directly selecting for fast or slow development. This experiment confirmed our initial findings, since selection for slow development resulted in the evolution of long life span and increased heat‐shock resistance. Because there were no consistent trade‐offs with growth rate or fecundity, our results highlight the key role of development rate – differentiation of the somatic cells per unit of time – in the evolution of life span. Since development time is under strong selection in nature, reduced somatic maintenance resulting in shorter life span may be a widespread cost of rapid development. A lay summary is available for this article. Lay Summary

Parasitic chytrid fungi have emerged as a significant threat to amphibian species worldwide, necessitating the development of techniques to isolate these pathogens into culture for research purposes. However, early methods of isolating chytrids from their hosts relied on killing amphibians. We modified a pre-existing protocol for isolating chytrids from infected animals to use toe clips and biopsies from toe webbing rather than euthanizing hosts, and distributed the protocol to researchers as part of the BiodivERsA project RACE; here called the RML protocol. In tandem, we developed a lethal procedure for isolating chytrids from tadpole mouthparts. Reviewing a database of use a decade after their inception, we find that these methods have been applied across 5 continents, 23 countries and in 62 amphibian species. Isolation of chytrids by the non-lethal RML protocol occured in 18% of attempts with 207 fungal isolates and three species of chytrid being recovered. Isolation of chytrids from tadpoles occured in 43% of attempts with 334 fungal isolates of one species (Batrachochytrium dendrobatidis) being recovered. Together, these methods have resulted in a significant reduction and refinement of our use of threatened amphibian species and have improved our ability to work with this group of emerging pathogens.

Populations of the same species may differ in their sensitivity to pathogens but the factors behind this variation are poorly understood. Moreover, infections may cause sub-lethal fitness effects even in species resistant or tolerant to disease. The chytrid fungus Batrachochytrium dendrobatidis (Bd), is a generalist pathogen which has caused amphibian population declines worldwide. In many species, Bd infection causes the disease chytridiomycosis, often leading to high mortality. We investigated how geographical origin affects tolerance to Bd by exposing newly metamorphosed individuals of two North European amphibians (moor frog Rana arvalis, common toad Bufo bufo) from two latitudinal regions to two different BdGPL strains. Bd exposure strongly lowered survival in B. bufo, and in both species survival was lower in the northern region, this difference being much stronger in B. bufo. Northern individuals were smaller in both species, and the survival difference between the regions was size-mediated with smaller individuals being more sensitive to Bd. In both species, Bd exposure led to sub-lethal effects in terms of reduced growth suggesting that even individuals surviving the infection may have reduced fitness mediated by smaller body size. Bd strain affected size-dependent mortality differently in the two regions. We discuss the possible mechanisms how body size and geographical origin can contribute to the present results.