Many scientists argue that we are either entering or in the midst of the sixth great mass extinction. Intense human pressure, both direct and indirect, is having profound effects on natural environments. The amphibians-frogs, salamanders, and caecilians-may be the only major group currently at risk globally. A detailed worldwide assessment and subsequent updates show that onethird or more of the 6,300 species are threatened with extinction. This trend is likely to accelerate because most amphibians occur in the tropics and have small geographic ranges that make them susceptible to extinction. The increasing pressure from habitat destruction and climate change is likely to have major impacts on narrowly adapted and distributed species. We show that salamanders on tropical mountains are particularly at risk. A new and significant threat to amphibians is a virulent, emerging infectious disease, chytridiomycosis, which appears to be globally distributed, and its effects may be exacerbated by global warming. This disease, which is caused by a fungal pathogen and implicated in serious declines and extinctions of >200 species of amphibians, poses the greatest threat to biodiversity of any known disease. Our data for frogs in the Sierra Nevada of California show that the fungus is having a devastating impact on native species, already weakened by the effects of pollution and introduced predators. A general message from amphibians is that we may have little time to stave off a potential mass extinction.chytridiomycosis ͉ climate change ͉ population declines ͉ Batrachochytrium dendrobatidis ͉ emerging disease

Epidemiological theory generally suggests that pathogens will not cause host extinctions because the pathogen should fade out when the host population is driven below some threshold density. An emerging infectious disease, chytridiomycosis, caused by the fungal pathogen Batrachochytrium dendrobatidis (Bd) is directly linked to the recent extinction or serious decline of hundreds of amphibian species. Despite continued spread of this pathogen into uninfected areas, the dynamics of the host-pathogen interaction remain unknown. We use fine-scale spatiotemporal data to describe (i) the invasion and spread of Bd through three lake basins, each containing multiple populations of the mountain yellow-legged frog, and (ii) the accompanying host-pathogen dynamics. Despite intensive sampling, Bd was not detected on frogs in study basins until just before epidemics began. Following Bd arrival in a basin, the disease spread to neighboring populations at ≈700 m/yr in a wave-like pattern until all populations were infected. Within a population, infection prevalence rapidly reached 100% and infection intensity on individual frogs increased in parallel. Frog mass mortality began only when infection intensity reached a critical threshold and repeatedly led to extinction of populations. Our results indicate that the high growth rate and virulence of Bd allow the nearsimultaneous infection and buildup of high infection intensities in all host individuals; subsequent host population crashes therefore occur before Bd is limited by density-dependent factors. Preventing infection intensities in host populations from reaching this threshold could provide an effective strategy to avoid the extinction of susceptible amphibian species in the wild.amphibian declines | Batrachochytrium dendrobatidis | chytridiomycosis | emerging infectious disease | Rana muscosa E arth's biodiversity is increasingly threatened with extinction.

Chytridiomycosis, the disease caused by the chytrid fungus, Batrachochytrium dendrobatidis (Bd), has contributed to amphibian population declines and extinctions worldwide. The impact of this pathogen, however, varies markedly among amphibian species and populations. Following invasion into some areas of California's Sierra Nevada, Bd leads to rapid declines and local extinctions of frog populations ( Rana muscosa , R. sierrae ). In other areas, infected populations of the same frog species have declined but persisted at low host densities for many years. We present results of a 5-year study showing that infected adult frogs in persistent populations have low fungal loads, are surviving between years, and frequently lose and regain the infection. Here we put forward the hypothesis that fungal load dynamics can explain the different population-level outcomes of Bd observed in different areas of the Sierra Nevada and possibly throughout the world. We develop a model that incorporates the biological details of the Bd-host interaction. Importantly, model results suggest that host persistence versus extinction does not require differences in host susceptibility, pathogen virulence, or environmental conditions, and may be just epidemic and endemic population dynamics of the same host–pathogen system. The different disease outcomes seen in natural populations may result solely from density-dependent host–pathogen dynamics. The model also shows that persistence of Bd is enhanced by the long-lived tadpole stage that characterize these two frog species, and by nonhost Bd reservoirs.

Emerging infectious diseases threaten human and wildlife populations. Altered ecological interactions between mutualistic microbes and hosts can result in disease, but an understanding of interactions between host, microbes and disease-causing organisms may lead to management strategies to affect disease outcomes. Many amphibian species in relatively pristine habitats are experiencing dramatic population declines and extinctions due to the skin disease chytridiomycosis, which is caused by the chytrid fungus Batrachochytrium dendrobatidis. Using a randomized, replicated experiment, we show that adding an antifungal bacterial species, Janthinobacterium lividum, found on several species of amphibians to the skins of the frog Rana muscosa prevented morbidity and mortality caused by the pathogen. The bacterial species produces the anti-chytrid metabolite violacein, which was found in much higher concentrations on frog skins in the treatments where J. lividum was added. Our results show that cutaneous microbes are a part of amphibians' innate immune system, the microbial community structure on frog skins is a determinant of disease outcome and altering microbial interactions on frog skins can prevent a lethal disease outcome. A bioaugmentation strategy may be an effective management tool to control chytridiomycosis in amphibian survival assurance colonies and in nature.

Amphibians highlight the global biodiversity crisis because ∼40% of all amphibian species are currently in decline. Species have disappeared even in protected habitats (e.g., the enigmatic extinction of the golden toad, Bufo periglenes, from Costa Rica). The emergence of a fungal pathogen, Batrachochytrium dendrobatidis (Bd), has been implicated in a number of declines that have occurred in the last decade, but few studies have been able to test retroactively whether Bd emergence was linked to earlier declines and extinctions. We describe a noninvasive PCR sampling technique that detects Bd in formalin-preserved museum specimens. We detected Bd by PCR in 83-90% (n = 38) of samples that were identified as positive by histology. We examined specimens collected before, during, and after major amphibian decline events at established study sites in southern Mexico, Guatemala, and Costa Rica. A pattern of Bd emergence coincident with decline at these localities is revealed-the absence of Bd over multiple years at all localities followed by the concurrent emergence of Bd in various species at each locality during a period of population decline. The geographical and chronological emergence of Bd at these localities also indicates a southbound spread from southern Mexico in the early 1970s to western Guatemala in the 1980s/1990s and to Monteverde, Costa Rica by 1987. We find evidence of a historical "Bd epidemic wave" that began in Mexico and subsequently spread to Central America. We describe a technique that can be used to screen museum specimens from other amphibian decline sites around the world.chytridiomycosis | emerging infectious disease | epizootic

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Amphibian population declines and extinctions are occurring even in the world's least impacted areas. The introduction and spread of nonnative predators is one of many proposed causes of amphibian declines. Correlational studies have shown a negative relationship between introduced fishes and declining amphibians, but little direct experimental evidence is available. This study experimentally manipulated the presence and absence of widely introduced salmonids rainbow trout (Oncorhynchus mykiss) and brook trout (Salvelinus fontinalis) to test the hypothesis that their introduction has contributed to the decline of the mountain yellow-legged frog (Rana muscosa). From 1996 to 2003, the introduced trout were removed from 5 lakes in a remote protected area of the Sierra Nevada, and 16 nearby lakes were used as controls, 8 with introduced trout and 8 without. To determine the vulnerable life stage, rainbow trout were placed in cages in three lakes containing amphibians. Removal of introduced trout resulted in rapid recovery of frog populations, and, in the caging experiment, tadpoles were found to be vulnerable to trout predation. Together, these experiments illustrate that introduced trout are effective predators on R. muscosa tadpoles and suggest (i) that the introduction of trout is the most likely mechanism responsible for the decline of this mountain frog and (ii) that these negative effects can be reversed.

Chytridiomycosis, caused by the fungal pathogen Batrachochytrium dendrobatidis, is an emerging infectious disease implicated in declines of amphibian populations around the globe. An emerging infectious disease is one that has recently been discovered; has recently increased in incidence, geography, or host range; or is newly evolved. For any given outbreak of an emerging disease, it is therefore possible to state two hypotheses regarding its origin. The novel pathogen hypothesis states that the disease has recently spread into new geographic areas, whereas the endemic pathogen hypothesis suggests that it has been present in the environment but recently has increased in host range or pathogenicity. Distinguishing between these hypotheses is important, because the conservation measures needed to slow or stop the spread of a novel pathogen are likely to differ from those needed to prevent outbreaks of an endemic pathogen. Population genetics may help discriminate among the possible origins of an emerging disease. Current evidence suggests chytridiomycosis may be a novel pathogen being spread worldwide by carriers; until we know how much genetic variation to expect in an endemic strain, however, we cannot yet conclude that B. dendrobatidis is a novel pathogen. La Hipótesis del Patógeno Incipiente y Endémico: Explicaciones Opuestas del Origen de Enfermedades Infecciosas Emergentes en la Vida Silvestre Resumen: La quitridiomicosis, causada por el hongo patógeno Batrachochytrium dendrobatidis, es una enfermedad infecciosa emergente implicada en las declinaciones de poblaciones de anfibios en el mundo. Una enfermedad infecciosa emergente es una que ha sido descubierta recientemente; que ha incrementado en su rango de incidencia, geográfico o de huéspedes recientemente; o que ha evolucionado recientemente. Para cualquier brote de una enfermedad emergente es posible enunciar dos hipótesis en relación con su origen. La hipótesis del patógeno incipiente establece que el la enfermedad se ha extendido recientemente haciaáreas geográficas nuevas, mientras que la hipótesis del patógeno endémico sugiere que ha estado presente en el ambiente pero que ha incrementado en el rango de huéspedes o de patogenicidad. Es importante distinguir entre estas dos hipótesis, porque es probable que las medidas de conservación que se requieren para reducir o detener la dispersión de un patógeno incipiente sean diferentes a las requeridas para prevenir brotes de ‡ ‡Current address: Resources Management and Sciences, 1442 Novel and Endemic Pathogen Hypotheses Rachowicz et al. un patógeno endémico. La genética de poblaciones puede ayudar a distinguir entre los posibles orígenes de una enfermedad emergente. La evidencia actual sugiere que la quitridiomicosis puede ser un patógeno incipiente que esta siendo dispersado por transportistas mundialmente; sin embargo, no podemos concluir que B. dendrobatidis es un patógeno incipiente hasta que no se conozca la variación genética esperada en una cepa endémica.Palabras Clave: Batrachochytrium dendrobatidis,...