Artículos de revistas
Disentangling Host, Pathogen, And Environmental Determinants Of A Recently Emerged Wildlife Disease: Lessons From The First 15 Years Of Amphibian Chytridiomycosis Research
Registro en:
Ecology And Evolution. John Wiley And Sons Ltd, v. 5, n. 18, p. 4079 - 4097, 2015.
20457758
10.1002/ece3.1672
2-s2.0-84942367159
Institución
Resumen
The amphibian fungal disease chytridiomycosis, which affects species across all continents, recently emerged as one of the greatest threats to biodiversity. Yet, many aspects of the basic biology and epidemiology of the pathogen, Batrachochytrium dendrobatidis (Bd), are still unknown, such as when and from where did Bd emerge and what is its true ecological niche? Here, we review the ecology and evolution of Bd in the Americas and highlight controversies that make this disease so enigmatic. We explore factors associated with variance in severity of epizootics focusing on the disease triangle of host susceptibility, pathogen virulence, and environment. Reevaluating the causes of the panzootic is timely given the wealth of data on Bd prevalence across hosts and communities and the recent discoveries suggesting co-evolutionary potential of hosts and Bd. We generate a new species distribution model for Bd in the Americas based on over 30,000 records and suggest a novel future research agenda. Instead of focusing on pathogen "hot spots," we need to identify pathogen "cold spots" so that we can better understand what limits the pathogen's distribution. Finally, we introduce the concept of "the Ghost of Epizootics Past" to discuss expected patterns in postepizootic host communities. © 2015 John Wiley & Sons Ltd. 5 18 4079 4097 Adams, M.J., Miller, D.A.W., Muths, E., Corn, P.S., Grant, E.H.C., Bailey, L.L., Trends in amphibian occupancy in the United States (2013) PLoS ONE, 8, p. e64347 Allentoft, M.E., O'Brien, J., Global amphibian declines, loss of genetic diversity and fitness: a review (2010) Diversity, 2, pp. 47-71 Anagnostakis, S.L., Hillman, B., Evolution of the chestnut tree and its blight (1992) Arnoldia, 52, pp. 3-10 Baláž, V., Vörös, J., Civiš, P., Vojar, J., Hettyey, A., Sós, E., Assessing risk and guidance on monitoring of Batrachochytrium dendrobatidis in Europe through identification of taxonomic selectivity of infection (2014) Conserv. Biol., 28, pp. 213-223 Bataille, A., Fong, J.J., Cha, M., Wogan, G.O.U., Baek, H.J., Lee, H., Genetic evidence for a high diversity and wide distribution of endemic strains of the pathogenic chytrid fungus Batrachochytrium dendrobatidis in wild Asian amphibians (2013) Mol. Ecol., 22, pp. 4196-4209 Bataille, A., Cashins, S.D., Grogan, L., Skerratt, L.F., Hunter, D., McFadden, M., Susceptibility of amphibians to chytridiomycosis is associated with MHC class II conformation (2015) Proc. Biol. Sci., 282, p. 20143127 Becker, C.G., Zamudio, K.R., Tropical amphibian populations experience higher disease risk in natural habitats (2011) Proc. Natl. Acad. Sci. USA, 108, pp. 9893-9898 Becker, C.G., Rodriguez, D., Toledo, L.F., Longo, A.V., Lambertini, C., Correa, D.T., Partitioning the net effect of host diversity on an emerging amphibian pathogen (2014) Proc. Biol. Sci., 281, p. 20141796 Berger, L., Speare, R., Daszak, P., Green, D.E., Cunningham, A.A., Goggin, C.L., Chytridiomycosis causes amphibian mortality associated with population declines in the rain forests of Australia and Central America (1998) Proc. Natl. Acad. Sci. USA, 95, pp. 9031-9036 Berger, L., Marantelli, G., Skerratt, L.L., Speare, R., Virulence of the amphibian chytrid fungus Batrachochytrium dendrobatidis varies with the strain (2005) Dis. Aquat. Organ., 68, pp. 47-50 Bielby, J., Cooper, N., Cunningham, A.A., Garner, T.W.J., Purvis, A., Predicting susceptibility to future declines in the world's frogs (2008) Conserv. Lett., 1, pp. 82-90 Blehert, D.S., Hicks, A.C., Behr, M., Meteyer, C.U., Berlowski-Zier, B.M., Buckles, E.L., Bat white-nose syndrome: An emerging fungal pathogen? (2009) Science, 323, pp. 227-227 Bletz, M.C., Loudon, A.H., Becker, M.H., Bell, S.C., Woodhams, D.C., Minbiole, K.P.C., Mitigating amphibian chytridiomycosis with bioaugmentation: characteristics of effective probiotics and strategies for their selection and use (2013) Ecol. Lett., 16, pp. 807-820 Briggs, C.J., Knapp, R.A., Vredenburg, V.T., Enzootic and epizootic dynamics of the chytrid fungal pathogen of amphibians (2010) Proc. Natl. Acad. Sci. USA, 107, pp. 9695-9700 Cashins, S.D., Grogan, L.F., McFadden, M., Hunter, D., Harlow, P.S., Berger, L., Prior infection does not improve survival against the amphibian disease chytridiomycosis (2013) PLoS ONE, 8, p. e56747 Catenazzi, A., Lehr, E., Rodriguez, L.O., Vredenburg, V.T., Batrachochytrium dendrobatidis and the collapse of anuran species richness and abundance in the Upper Manu National Park, southeastern Peru (2011) Conserv. Biol., 25, pp. 382-391 Cavender-Bares, J., Kozak, K.H., Fine, P.V.A., Kembel, S.W., The merging of community ecology and phylogenetic biology (2009) Ecol. Lett., 12, pp. 693-715 Cheng, T.L., Rovito, S.M., Wake, D.B., Vredenburg, V.T., Coincident mass extirpation of neotropical amphibians with the emergence of the infectious fungal pathogen Batrachochytrium dendrobatidis (2011) Proc. Natl. Acad. Sci. USA, 108, pp. 9502-9507 Chestnut, T., Anderson, C., Popa, R., Blaustein, A.R., Voytek, M., Olson, D.H., Heterogeneous occupancy and density estimates of the pathogenic fungus Batrachochytrium dendrobatidis in waters of North America (2014) PLoS ONE, 9, p. e106790 Collins, J.P., Crump, M.L., (2009) Extinction in our times. Global amphibian decline, , Oxford University Press, Oxford Crawford, A.J., Lips, K.R., Bermingham, E., Epidemic disease decimates amphibian abundance, species diversity, and evolutionary history in the highlands of central Panama (2010) Proc. Natl. Acad. Sci. USA, 107, pp. 13777-13782 Daszak, P., Cunningham, A.A., Hyatt, A.D., Anthropogenic environmental change and the emergence of infectious diseases in wildlife (2001) Acta Trop., 78, pp. 103-116 DiRenzo, G.V., Langhammer, P.F., Zamudio, K.R., Lips, K.R., Fungal infection intensity and zoospore output of Atelopus zeteki, a potential acute chytrid supershedder (2014) PLoS ONE, 9, p. e93356 Ellison, A.R., Savage, A.E., DiRenzo, G.V., Langhammer, P., Lips, K.R., Zamudio, K.R., Fighting a losing battle: vigorous immune response countered by pathogen suppression of host defenses in the chytridiomycosis-susceptible frog Atelopus zeteki (2014) G3 (Bethesda), 4, pp. 1275-1289 Ellison, A.R., Tunstall, T., DiRenzo, G.V., Hughey, M.C., Rebollar, E.A., Belden, L.K., More than skin deep: Functional genomic basis for resistance to amphibian chytridiomycosis (2015) Genome Biol. Evol., 7, pp. 286-298 Eterovick, P.C., Carnaval, A., Borges-Nojosa, D.M., Silvano, D.L., Segalla, M.V., Sazima, I., Amphibian declines in Brazil: an overview (2005) Biotropica, 37, pp. 166-179 Ewald, P.W., (1994) Evolution of infectious disease, , Oxford University Press, Oxford Farrer, R.A., Weinert, L.A., Bielby, J., Garner, T.W., Balloux, F., Clare, F., Multiple emergences of genetically diverse amphibian-infecting chytrids include a globalized hypervirulent recombinant lineage (2011) Proc. Natl. Acad. Sci. USA, 108, pp. 18732-18736 Fisher, M.C., Bosch, J., Yin, Z., Stead, D.A., Walker, J., Selway, L., Proteomic and phenotypic profiling of the amphibian pathogen Batrachochytrium dendrobatidis shows that genotype is linked to virulence (2009) Mol. Ecol., 18, pp. 415-429 Fisher, M.C., Walker, S.F., Garner, T.W.J., The global emergence of Batrachochytrium dendrobatidis in space, time, and host (2009) Annu. Rev. Microbiol., 63, pp. 291-310 Fisher, M.C., Henk, D.A., Briggs, C.J., Brownstein, J.S., Madoff, L.C., McCraw, S.L., Emerging fungal threats to animal, plant and ecosystem health (2012) Nature, 484, pp. 186-194 Gahl, M.K., Longcore, J.E., Houlahan, J.E., Varying responses of northeastern North American amphibians to the chytrid pathogen Batrachochytrium dendrobatidis (2012) Conserv. Biol., 26, pp. 135-141 Garner, T.W.J., Perkins, M.W., Govindarajulu, P., Seglie, D., Walker, S., Cunningham, A.A., The emerging amphibian pathogen Batrachochytrium dendrobatidis globally infects introduced populations of the North American bullfrog, Rana catesbeiana (2006) Biol. Lett., 2, pp. 455-459 Greenspan, S.E., Longcore, J.E., Calhoun, A.J.K., Host invasion by Batrachochytrium dendrobatidis: fungal and epidermal ultrastructure in model anurans (2012) Dis. Aquat. Organ., 100, pp. 201-210 Gründler, M.C., Toledo, L.F., Parra-Olea, G., Haddad, C.F.B., Giasson, L.O.M., Sawaya, R.J., Interaction between breeding habitat and elevation affects prevalence but not infection intensity of Batrachochytrium dendrobatidis in Brazilian anuran assemblages (2012) Dis. Aquat. Organ., 97, pp. 173-184 Goka, K., Yokoyama, J., Une, Y., Kuroki, T., Suzuki, K., Nakahara, M., Kobayashi, A., Hyatt, A.D., Amphibian chytridiomycosis in Japan: distribution, haplotypes and possible route of entry into Japan (2009) Mol. Ecol., 18, pp. 4757-4774 Gurr, S., Samalova, M., Fisher, M., The rise and rise of emerging infectious fungi challenges food security and ecosystem health (2011) Fungal Biol. Rev., 25, pp. 181-188 Hanlon, S.M., Parris, M.J., The interactive effects of chytrid fungus, pesticides, and exposure timing on gray treefrog (Hyla versicolor) larvae (2014) Environ. Toxicol. Chem., 33, pp. 216-222 Hine, R.L., Les, B.L., Hellmich, B.F., (1981) Leopard frog populations and mortality in Wisconsin, 1974-1976, , Technical Bulletin No. 122, Department of Natural Resources, Madison, Wisconsin Houlahan, J.E., Findlay, C.S., Schmidt, B.R., Meyer, A.H., Kuzmin, S.L., Quantitative evidence for global amphibian population declines (2000) Nature, 404, pp. 752-755 James, T.Y., Litvintseva, A.P., Vilgalys, R., Morgan, J.A.T., Taylor, J.W., Fisher, M.C., Rapid expansion of an emerging fungal disease into declining and healthy amphibian populations (2009) PLoS Pathogens, 5, p. e1000458 Johnson, M.L., Speare, R., Survival of Batrachochytrium dendrobatidis in water: Quarantine and disease control implications (2003) Emerg. Infect. Dis., 9, pp. 922-925 Jones, K.E., Patel, N.G., Levy, M.A., Storeygard, A., Balk, D., Gittleman, J.L., Global trends in emerging infectious diseases (2008) Nature, 451. , 990-9U4 Keesing, F., Holt, R.D., Ostfeld, R.S., Effects of species diversity on disease risk (2006) Ecol. Lett., 9, pp. 485-498 Kilpatrick, A.M., Briggs, C.J., Daszak, P., The ecology and impact of chytridiomycosis: an emerging disease of amphibians (2010) Trends Ecol. Evol., 25, pp. 109-118 Knapp, R.A., Briggs, C.J., Smith, T.C., Maurer, J.R., Nowhere to hide: impact of a temperature-sensitive amphibian pathogen along an elevation gradient in the temperate zone (2011) Ecosphere, 2 (8) Kriger, K.M., Hero, J.M., The chytrid fungus Batrachochytrium dendrobatidis is non-randomly distributed across amphibian breeding habitats (2007) Divers. Distrib., 13, pp. 781-788 Kriger, K.M., Pereoglou, F., Hero, J.M., Latitudinal variation in the prevalence and intensity of chytrid (Batrachochytrium dendrobatidis) infection in Eastern Australia (2007) Conserv. Biol., 21, pp. 1280-1290 Langhammer, P.F., Lips, K.R., Burrowes, P.A., Tunstall, T., Palmer, C.M., Collins, J.P., A fungal pathogen of amphibians, Batrachochytrium dendrobatidis, attenuates in pathogenicity with in vitro passages (2013) PLoS ONE, 8, p. e77630 Langwig, K.E., Frick, W.F., Bried, J.T., Hicks, A.C., Kunz, T.H., Kilpatrick, A.M., Sociality, density-dependence and microclimates determine the persistence of populations suffering from a novel fungal disease, white-nose syndrome (2012) Ecol. Lett., 15, pp. 1050-1057 Lees, A.C., Pimm, S.L., Species, extinct before we know them? (2015) Curr. Biol., 25, pp. R177-R180 Lips, K.R., Decline of a tropical amphibian fauna (1998) Conserv. Biol., 12, pp. 106-117 Lips, K.R., Brem, F., Brenes, R., Reeve, J.D., Alford, R.A., Voyles, J., Emerging infectious disease and the loss of biodiversity in a Neotropical amphibian community (2006) Proc. Natl. Acad. Sci. USA, 103, pp. 3165-3170 Liu, X., Rohr, J.R., Li, Y.M., Climate, vegetation, introduced hosts and trade shape a global wildlife pandemic (2013) Proc. Biol. Sci., 280 Longcore, J.E., Pessier, A.P., Nichols, D.K., Batrachochytrium dendrobatidis gen et sp nov, a chytrid pathogenic to amphibians (1999) Mycologia, 91, pp. 219-227 Longo, A.V., Burrowes, P.A., Joglar, R.L., Seasonality of Batrachochytrium dendrobatidis infection in direct-developing frogs suggests a mechanism for persistence (2010) Dis. Aquat. Organ., 92, pp. 253-260 Longo, A.V., Rodriguez, D., Leite, D.D., Toledo, L.F., Almeralla, C.M., Burrowes, P.A., ITS1 copy number varies among Batrachochytrium dendrobatidis strains: Implications for qPCR estimates of infection intensity from field-collected amphibian skin swabs (2013) PLoS ONE, 8, p. e59499 Martel, A., Spitzen-van der Sluijs, A., Blooi, M., Bert, W., Ducatelle, R., Fisher, M.C., Batrachochytrium salamandrivorans sp nov causes lethal chytridiomycosis in amphibians (2013) Proc. Natl. Acad. Sci. USA, 110, pp. 15325-15329 Martel, A., Blooi, M., Adriaensen, C., Van Rooij, P., Beukema, W., Fisher, M.C., Recent introduction of a chytrid fungus endangers Western Palearctic salamanders (2014) Science, 346, pp. 630-631 McCracken, S., Gaertner, J.P., Forstner, M.R.J., Hahn, D., Detection of Batrachochytrium dendrobatidis in amphibians from the forest floor to the upper canopy of an Ecuadorian Amazon lowland rainforest (2009) Herpetol. Rev., 40, p. 190 McMahon, T.A., Brannelly, L.A., Chatfield, M.W.H., Johnson, P.T.J., Joseph, M.B., McKenzie, V.J., Chytrid fungus Batrachochytrium dendrobatidis has nonamphibian hosts and releases chemicals that cause pathology in the absence of infection (2012) Proc. Natl. Acad. Sci. USA, 110, pp. 210-215 McMahon, T.A., Sears, B.F., Venesky, M.D., Bessler, S.M., Brown, J.M., Deutsch, K., Amphibians acquire resistance to live and dead fungus overcoming fungal immunosuppression (2014) Nature, 511, pp. 224-227 Meller, L., Cabeza, M., Pironon, S., Barbet-Massin, M., Maiorano, L., Georges, D., Ensemble distribution models in conservation prioritization: from consensus predictions to consensus reserve networks (2014) Divers. Distrib., 20, pp. 309-321 Mitchell, K.M., Churcher, T.S., Garner, T.W.J., Fisher, M.C., Persistence of the emerging pathogen Batrachochytrium dendrobatidis outside the amphibian host greatly increases the probability of host extinction (2008) Proc. Biol. Sci., 275, pp. 329-334 Monastersky, R., Biodiversity: Life-a status report (2014) Nature, 516, pp. 158-161 Morgan, J.A.T., Vredenburg, V.T., Rachowicz, L.J., Knapp, R.A., Stice, M.J., Tunstall, T., Population genetics of the frog-killing fungus Batrachochytrium dendrobatidis (2007) Proc. Natl. Acad. Sci. USA, 104, pp. 13845-13850 Olson, D.H., Aanensen, D.M., Ronnenberg, K.L., Powell, C.I., Walker, S.F., Bielby, J., Mapping the global emergence of Batrachochytrium dendrobatidis, the amphibian chytrid fungus (2013) PLoS ONE, 8, p. e56802 Peterson, A.C., McKenzie, V.J., Investigating differences across host species and scales to explain the distribution of the amphibian pathogen Batrachochytrium dendrobatidis (2014) PLoS ONE, 9, p. e107441 Piotrowski, J.S., Annis, S.L., Longcore, J.E., Physiology of Batrachochytrium dendrobatidis, a chytrid pathogen of amphibians (2004) Mycologia, 96, pp. 9-15 Puschendorf, R., Carnaval, A.C., VanDerWal, J., Zumbado-Ulate, H., Chaves, G., Bolanos, F., Distribution models for the amphibian chytrid Batrachochytrium dendrobatidis in Costa Rica: proposing climatic refuges as a conservation tool (2009) Divers. Distrib., 15, pp. 401-408 Rachowicz, L.J., Hero, J.M., Alford, R.A., Taylor, J.W., Morgan, J.A.T., Vredenburg, V.T., The novel and endemic pathogen hypotheses: Competing explanations for the origin of emerging infectious diseases of wildlife (2005) Conserv. Biol., 19, pp. 1441-1448 Ren, P., Haman, K.H., Last, L.A., Rajkumar, S.S., Keel, M.K., Chaturvedi, V., Clonal spread of Geomyces destructans among bats, midwestern and southern United States (2012) Emerg. Infect. Dis., 18, pp. 883-885 Rödder, D., Kielgast, J., Bielby, J., Schmidtlein, S., Bosch, J., Garner, T.W.J., Global amphibian extinction risk assessment for the panzootic chytrid fungus (2009) Diversity, 1, pp. 52-66 Rodriguez, D., Longo, A.V., Zamudio, K.R., Magnetic capture hybridization of Batrachochytrium dendrobatidis genomic DNA (2012) J. Microbiol. Methods, 90, pp. 156-159 Rodriguez, D., Becker, C.G., Pupin, N.C., Haddad, C.F.B., Zamudio, K.R., Long-term endemism of two highly divergent lineages of the amphibian-killing fungus in the Atlantic Forest of Brazil (2014) Mol. Ecol., 23, pp. 774-787 Rosenblum, E.B., Poorten, T.J., Settles, M., Murdoch, G.K., Only skin deep: shared genetic response to the deadly chytrid fungus in susceptible frog species (2012) Mol. Ecol., 21, pp. 3110-3120 Rosenblum, E.B., James, T.Y., Zamudio, K.R., Poorten, T.J., Ilut, D., Rodriguez, D., Complex history of the amphibian-killing chytrid fungus revealed with genome resequencing data (2013) Proc. Natl. Acad. Sci. USA, 110, pp. 9385-9390 Rowley, J.J.L., Alford, R.A., Hot bodies protect amphibians against chytrid infection in nature (2013) Sci. Rep., 3, p. 1515 Sanderson, E.W., Jaiteh, M., Levy, M.A., Redford, K.H., Wannebo, A.V., Woolmer, G., The human footprint and the last of the wild (2002) Bioscience, 52, pp. 891-904 Savage, A.E., Zamudio, K.R., MHC genotypes associate with resistance to a frog-killing fungus (2011) Proc. Natl. Acad. Sci. USA, 108, pp. 16705-16710 Schierup, M.H., Mikkelsen, A.M., Hein, J., Recombination, balancing selection and phylogenies in MHC and self-incompatibility genes (2001) Genetics, 159, pp. 1833-1844 Schloegel, L.M., Toledo, L.F., Longcore, J.E., Greenspan, S.E., Viera, C.A., Lee, M., Novel, panzootic, and hybrid genotypes of amphibian chytridiomycosis associated with the bullfrog trade (2012) Mol. Ecol., 21, pp. 5162-5177 Scholthof, K.B.G., The disease triangle: pathogens, the environment and society (2007) Nat. Rev. Microbiol., 5, pp. 152-156 Searle, C.L., Biga, L.M., Spatafora, J.W., Blaustein, A.R., A dilution effect in the emerging amphibian pathogen Batrachochytrium dendrobatidis (2011) Proc. Natl. Acad. Sci. USA, 108, pp. 16322-16326 Searle, C.L., Gervasi, S.S., Hua, J., Hammond, J.I., Relyea, R.A., Olson, D.H., Differential host susceptibility to Batrachochytrium dendrobatidis, an emerging amphibian pathogen (2011) Conserv. Biol., 25, pp. 965-974 Soto-Azat, C., Valenzuela-Sanchez, A., Collen, B., Rowcliffe, J.M., Veloso, A., Cunningham, A.A., The population decline and extinction of Darwin's frogs (2013) PLoS ONE, 8, p. e66957 Talley, B.L., Muletz, C.R., Vredenburg, V.T., Fleischer, R.C., Lips, K.R., A century of Batrachochytrium dendrobatidis in Illinois amphibians (1888-1989) (2015) Biol. Conserv., 182, pp. 254-261 Thuiller, W., Georges, D., Engler, R., (2014) biomod2: Ensemble platform for species distribution modeling, pp. 1-64. , http://CRAN.R-project.org/package=biomod2, R package version 3 Tinsley, R.C., Coxhead, P.G., Stott, L.C., Tinsley, M.C., Piccinni, M.Z., Guille, M.J., Chytrid fungus infections in laboratory and introduced Xenopus laevis populations: assessing the risks for U.K. native amphibians (2015) Biol. Conserv., 184, pp. 380-388 Trabucco, A., Zomer, R.J., Global Aridity Index (Global-Aridity) and Global Potential Evapo-Transpiration (Global-PET) Geospatial Database (2009), http://www.csi.cgiar.org/, Published online, available from the CGIAR-CSI GeoPortal at:CGIAR Consortium for Spatial InformationVelo-Anton, G., Rodriguez, D., Savage, A.E., Parra-Olea, G., Lips, K.R., Zamudio, K.R., Amphibian-killing fungus loses genetic diversity as it spreads across the New World (2012) Biol. Conserv., 146, pp. 213-218 Venesky, M.D., Kerby, J.L., Storfer, A., Parris, M.J., Can differences in host behavior drive patterns of disease prevalence in tadpoles? (2011) PLoS ONE, 6, p. e24991 Venesky, M.D., Liu, X., Sauer, E.L., Rohr, J.R., Linking manipulative experiments to field data to test the dilution effect (2014) J. Anim. Ecol., 83, pp. 557-565 Voyles, J., Johnson, L.R., Briggs, C.J., Cashins, S.D., Alford, R.A., Berger, L., Experimental evolution alters the rate and temporal pattern of population growth in Batrachochytrium dendrobatidis, a lethal fungal pathogen of amphibians (2014) Ecol. Evol., 4, pp. 3633-3641 Voyles, J., Kilpatrick, A.M., Collins, J.P., Fisher, M.C., Frick, W.F., McCallum, H., Moving beyond too little, too late: Managing emerging infectious diseases in wild populations requires international policy and partnerships (2014) EcoHealth, 2014, pp. 1-4 Vrålstad, T., Strand, D.A., Grandjean, F., Kvellestad, A., Håstein, T., Knutsen, A.K., Molecular detection and genotyping of Aphanomyces astaci directly from preserved crayfish samples uncovers the Norwegian crayfish plague disease history (2014) Vet. Microbiol., 173, pp. 66-75 Vredenburg, V.T., Knapp, R.A., Tunstall, T.S., Briggs, C.J., Dynamics of an emerging disease drive large-scale amphibian population extinctions (2010) Proc. Natl. Acad. Sci. USA, 107, pp. 9689-9694 Vredenburg, V.T., Felt, S.A., Morgan, E.C., McNally, S.V.G., Wilson, S., Green, S.L., Prevalence of Batrachochytrium dendrobatidis in Xenopus collected in Africa (1871-2000) and in California (2001-2010) (2013) PLoS ONE, 8, p. e63791 Walker, S.F., Bosch, J., James, T.Y., Litvintseva, A.P., Valls, J.A.O., Piña, S., Introduced pathogens threaten species recovery programs (2008) Curr. Biol., 18, pp. R853-R854 Weldon, C., du Preez, L.H., Hyatt, A.D., Muller, R., Speare, R., Origin of the amphibian chytrid fungus (2004) Emerg. Infect. Dis., 10, pp. 2100-2105 Wilcoxen, T.E., Boughton, R.K., Schoech, S.J., Selection on innate immunity and body condition in Florida scrub-jays throughout an epidemic (2010) Biol. Lett., 6, pp. 552-554