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Kiesecker 2010 Global stressors and the global decline of amphibians:tipping the stress immunocompetency axis

Kiesecker 2010 Global stressors and the global decline of amphibians:tipping the stress immunocompetency axis

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Published by joekiesecker
Abstract There is a widespread consensus that the earth
is experiencing a mass extinction event and at the
forefront are amphibians, the most threatened of all
vertebrate taxa. A recent assessment found that nearly
one-third (32%, 1,856 species) of the world’s amphibian
species are threatened. Amphibians have existed on the
earth for over 300 million years, yet in just the last two
decades there have been an alarming number of extinctions,
nearly 168 species are believed to have gone extinct
and at least 2,469 (43%) more have populations
that are declining. Infectious diseases have been recognized
as one major cause of worldwide amphibian
population declines. This could be the result of the
appearance of novel pathogens, or it could be that
exposure to environmental stressors is increasing the
susceptibility of amphibians to opportunistic pathogens.
Here I review the potential effects of stressors on disease
susceptibility in amphibians and relate this to disease
emergence in human and other wildlife populations. I
will present a series of case studies that illustrate the role
of stress in disease outbreaks that have resulted in
amphibian declines. First, I will examine how elevated
sea-surface temperatures in the tropical Pacific since the
mid-1970s have affected climate over much of the world
and could be setting the stage for pathogen-mediated
amphibian declines in many regions. Finally, I will discuss
how the apparently rapid increase in the prevalence
of amphibian limb deformities is linked to the synergistic
effects of trematode infection and exposure to chemical
contaminants.
Abstract There is a widespread consensus that the earth
is experiencing a mass extinction event and at the
forefront are amphibians, the most threatened of all
vertebrate taxa. A recent assessment found that nearly
one-third (32%, 1,856 species) of the world’s amphibian
species are threatened. Amphibians have existed on the
earth for over 300 million years, yet in just the last two
decades there have been an alarming number of extinctions,
nearly 168 species are believed to have gone extinct
and at least 2,469 (43%) more have populations
that are declining. Infectious diseases have been recognized
as one major cause of worldwide amphibian
population declines. This could be the result of the
appearance of novel pathogens, or it could be that
exposure to environmental stressors is increasing the
susceptibility of amphibians to opportunistic pathogens.
Here I review the potential effects of stressors on disease
susceptibility in amphibians and relate this to disease
emergence in human and other wildlife populations. I
will present a series of case studies that illustrate the role
of stress in disease outbreaks that have resulted in
amphibian declines. First, I will examine how elevated
sea-surface temperatures in the tropical Pacific since the
mid-1970s have affected climate over much of the world
and could be setting the stage for pathogen-mediated
amphibian declines in many regions. Finally, I will discuss
how the apparently rapid increase in the prevalence
of amphibian limb deformities is linked to the synergistic
effects of trematode infection and exposure to chemical
contaminants.

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Published by: joekiesecker on Jan 09, 2011
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04/27/2012

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SPECIAL FEATURE
Environmental change, pathogens and human linkages
Joseph M. Kiesecker
Global stressors and the global decline of amphibians: tipping the stress immunocompetency axis
Received: 2 February 2009/ Accepted: 1 February 2010
Ó
The Ecological Society of Japan 2010
Abstract
There is a widespread consensus that the earthis experiencing a mass extinction event and at theforefront are amphibians, the most threatened of allvertebrate taxa. A recent assessment found that nearlyone-third (32%, 1,856 species) of the world’s amphibianspecies are threatened. Amphibians have existed on theearth for over 300 million years, yet in just the last twodecades there have been an alarming number of extinc-tions, nearly 168 species are believed to have gone ex-tinct and at least 2,469 (43%) more have populationsthat are declining. Infectious diseases have been recog-nized as one major cause of worldwide amphibianpopulation declines. This could be the result of theappearance of novel pathogens, or it could be thatexposure to environmental stressors is increasing thesusceptibility of amphibians to opportunistic pathogens.Here I review the potential effects of stressors on diseasesusceptibility in amphibians and relate this to diseaseemergence in human and other wildlife populations. Iwill present a series of case studies that illustrate the roleof stress in disease outbreaks that have resulted inamphibian declines. First, I will examine how elevatedsea-surface temperatures in the tropical Pacific since themid-1970s have affected climate over much of the worldand could be setting the stage for pathogen-mediatedamphibian declines in many regions. Finally, I will dis-cuss how the apparently rapid increase in the prevalenceof amphibian limb deformities is linked to the synergisticeffects of trematode infection and exposure to chemicalcontaminants.
Keywords
Global amphibian declines
Æ
Stress-mediatedinfection
Æ
Climate change
Æ
Pesticide exposure
Introduction
What does the global decline in amphibians have to dowith disease outbreaks in human populations and otherspecies of wildlife? In the past two decades at least 20major human diseases have reemerged in novel, morevirulent forms (Epstein1999). Worldwide, scientistshave discovered at least 30 previously unknown humandiseases, such as Ebola, AIDS, and SARS (Fauci2001;Weinhold2004). Moreover, the last decade has seen anumber of epidemics that have caused large-scaledeclines in several wildlife species (Dobson and Carper1993; Daszak et al.2000). These changes have occurred in synchrony with the global decline of amphibian spe-cies (Blaustein and Kiesecker2002; Collins and Storfer2003; Semlitsch2003). The global loss of amphibian populations was first recognized in 1989 as a phenome-non that deserved world-wide attention (Blaustein andWake1990; Blaustein et al.1994). To date, approxi- mately 168 species of frogs and salamanders are severelythreatened or have gone extinct (GAA2004). Includedwith many of these declines are reports of massivemortality associated with pathogenic infection and anincreased incidence of developmental malformations(Daszak et al.2001; Blaustein and Kiesecker2002). In some populations, more than 80% of the individualsexhibit severe deformities, including extra or missinglimbs (Ankley et al.2002; Johnson et al.2001,2002; Johnson and Sutherland2003; see also Hayes et al.2002a,2002b,2003). The widespread nature of the de- clines and the apparent increase in the prevalence of deformities has lead to substantial interest from scien-tists and the general public.How is the global decline of amphibians related toincreased disease prevalence in human and wildlifepopulations? As human populations have grown, theworld has dramatically changed. Between one-third andone-half of Earth’s land surface has been altered byhuman action (Vitousek et al.1997). The concentrationof carbon dioxide in the atmosphere has increased by
J. M. Kiesecker (
&
)North America Conservation Region, The Nature Conservancy,117 E Mountain, Suite 201, Fort Collins, CO 80524, USAE-mail: jkiesecker@tnc.orgTel.: +1-970-4849598Ecol ResDOI 10.1007/s11284-010-0702-6
 
approximately 30% since the late 1800s. Human actionsresult in the fixation of more nitrogen than all naturalterrestrial sources and over half of accessible surfacefreshwater is utilized by human actions (Vitousek et al.1997). I suggest that these changes have created anenvironment that fosters emergence of infectious dis-eases for both wildlife and human populations (Fig.1).Emerging diseases are those that have increased inincidence, pathogenicity, geographic range, have shiftedhosts or recently evolved new strains (Daszak et al.2000). Emerging infectious diseases fall into two over-arching categories. The first category includes parasiticorganisms that have invaded into new populations.These diseases are characterized by an explosive spreadthrough the naive host population after the introductionof the novel pathogen. The second category includesoutbreaks where the parasite is native to a particularhost and geographic range but is spreading becausesome external factor either promotes transmission be-tween hosts or decreases a host’s response to infection.
 Amphibians as bioindicators of environmental health
Concern about amphibians is in large part due to theirvalue as indicators of environmental health (Blausteinand Kiesecker2002; Storfer2003; Kiesecker et al.2004). Relative to other vertebrates, amphibians have unpro-tected permeable skin, lacking scales or hair (Blausteinand Bancroft2007; Han et al.2008). Their embryos develop without the benefit of shells and thus are directlyexposed to environmental conditions. Their complexlifecycles expose them to both aquatic and terrestrialhazards. These characteristics make them particularlysensitive to changes in environmental conditions. Thewidespread nature of the disease-mediated declines andthe increase in deformities could be a warning of envi-ronmental degradation (Lannoo2008). Understandingthe factors that drive increased incidence of infectiousdisease in amphibian populations is central to under-standing increased disease prevalence in human andwildlife populations because the majority of emergingdiseases are linked to human-induced environmentalchange (Daszak et al.2001; Kiesecker et al.2004). Moreover, wildlife populations have long been consid-ered an important link to pathogen emergence, as theyserve as reservoirs for zoonotic diseases (pathogenstransmissible between humans and animals). Many of the emerging infectious agents of humans are zoonoticdiseases (Daszak et al.2000). Thus, increased pathogenicinfection in wildlife populations could translate into in-creased risk of human infection. Here I consider a fewcases of amphibian mortality and deformity that havebeen associated with increased pathogenic infection todemonstrate that the global decline in amphibians isrelated to a larger phenomena associated with increaseddisease prevalence in human and wildlife populations. Iwill examine cases of decline that fall under the twobroad classes of emerging infectious disease, environ-mental stress and introduced pathogens, to illustrate thesimilarity of mechanisms that have promoted diseaseoutbreaks.
Important amphibian pathogens
Although little is known about the effects of pathogenson wild amphibians, there are a number of studiesshowing that they may be significant contributors todeclining amphibian populations. A variety of patho-gens may affect wild amphibian populations. These in-clude viruses, bacteria, trematode parasites, protozoans,oomycetes, and fungi. These pathogens can be theproximate causes of mortality or they can cause suble-thal damage such as severe developmental and physio-logical deformities.Three pathogens have received a great deal of atten-tion recently with regard to amphibian population de-clines and deformities; a chytridomycete fungus,
Batrachochytrium dendrobatidis
, hereafter referred to asBD (Berger et al.1998), a fungal-like oomycete,
Sapro-legnia ferax,
hereafter SF (Kiesecker and Blaustein1995;Kiesecker and Blaustein1997a; Kiesecker et al.2001a) and a digenetic trematode,
Ribeiroia ondatrae,
hereafterRO (Johnson et al.2002). I recognize that numerouspathogens are associated with the amphibian decline/deformities phenomena (see for example Di Rosa et al.2007; Greer and Collins2007; Duffus et al.2008) but here I focus on these three pathogens because of thewealth of studies examining the role of environmentalchange in their emergence. BD is found in several areaswhere population declines have occurred and is fatal tofrogs under certain experimental conditions (Bergeret al.1998; Lips et al.2003; Ron et al.2003).
Saprolegnia
has long been known to affect amphibian embryos, andlarvae and SF has been associated with massive embryomortality in the Cascade Mountains of Oregon, USA.Recent experimental evidence suggests that SF may
Fig. 1
Relationship of habitat alteration often associated withincreased human development that may alter disease dynamics
 
affect amphibians in several ways and its infection rateand virulence depend upon a number of factors(Kiesecker and Blaustein1995; Kiesecker et al.2001a, 2001b). The presence of trematode parasite, RO hasbeen associated with the occurrence of limb deformitiesat natural breeding sites. Experimental studies haveshown that tadpoles exposed to larval RO develop limbdeformities similar to those seen at natural breeding sites(Blaustein and Johnson2003)
.
Below I examine thecontext-dependent nature of these host–pathogen inter-actions and compare these with representative diseaseoutbreaks in other wildlife species and human popula-tions.
Environmental stress and increased disease prevalence
It has long been known that environmental stressors canincrease host susceptibility to disease (Daszak et al.2001;Lafferty and Holt2003). A number of environmentalstressors (i.e., habitat modification, UV-B radiation, cli-mate change, pesticides) have been correlated with diseaseoutbreak in amphibian populations as well as otherwildlife and human populations. In this section, I willreview some examples of environmental changes that af-fectinfection riskfor amphibians and relate thisto similaroutbreaks in other wildlife and human populations.Global climate change and increased disease prevalenceAnthropogenic global climate change is likely to causechanges in the geographic range and incidence of manyinfectious diseases (Nicholls1993; Dobson and Carper1993; Reeves et al.1994; Patz et al.1996). Of particular concern is the rapid warming that has occurred of thelatter half of the last century (Houghton et al.2001). Inthe last century, surface temperatures have risen anaverage of 0.5
°
C, with concurrent changes in precipita-tionpatternsandanincreaseinthefrequencyandseverityof extreme weather events. In particular is the El Nin ˜o-Southern Oscillation (ENSO) phenomenon that resultsfrom air–sea interactions in the Tropical Pacific that im-pacts weather patterns over the entire globe (Timmer-mannetal.1999).ElNin ˜oeventsrefertothereplacementof normally cold nutrient-rich water over most of theequatorial Pacific Ocean with warm nutrient-deficientsurfacewaterevery2–7 years.LaNin ˜a,theoppositeofElNin ˜o, refers extreme cold water over the equatorialPacific Ocean. In 1976, a large change in sea-surfacetemperatures in the South Pacific resulted in an increasein the frequency, duration, and intensity of ENSO events(Timmermann et al.1999). Because ENSO events affecttemperatureandmoisturepatternsoverland,amphibiansand other organisms are likely to be effected.In the Pacific Northwest of the United States, massiveamphibian embryo mortality is associated with thepresence of the oomycete pathogen, SF (Kiesecker andBlaustein1997a). SF outbreaks have been linked to in-creased UV-B radiation (Kiesecker and Blaustein1995).Attempts to link these disease outbreaks with environ-mental change have often postulated that these patternsare connected to ozone depletion. However, in somecases, climate change can be more effective thanstratospheric ozone depletion in increasing the exposureof aquatic organisms to biologically effective UV-Bradiation (Yan et al.1996; Kiesecker et al.2001a). Amphibians, particularly those that breed in shallowmontane lakes and ponds, may be quite susceptible toclimate-induced changes in UV-B exposure (Blausteinet al.1998; Blaustein and Belden2003; Han et al.2008). Amphibian embryos developing in montane lakes andponds are often exposed to direct sunlight, yet theoverlying water column may attenuate UV-B radiation.Where precipitation is reduced, however, associatedreduction in water depth at oviposition sites may en-hance UV-B exposure (Yan et al.1996). In the PacificNorthwest of the United States, where SF outbreakshave occurred, precipitation patterns are closely tied toENSO cycles (Redmond and Koch.1991). For the Pa-cific Northwest El Nin ˜o events result in a decrease inwinter precipitation. Hence, the increase in frequencyand magnitude of El Nin ˜o events following the step-likewarming of the tropical Pacific may have raised theincidence and severity of SF outbreaks by increasing theextent to which embryos are exposed to sunlight inshallow water (Kiesecker et al.2001a).To test this hypothesis, we (Kiesecker et al.2001a)examined the relationship between pathogen-mediatedembryo mortality and climate by quantifying mortalityin relation to water depth at natural oviposition sites inthe context of interannual variation in precipitation andthe Southern Oscillation Index (SOI). We used experi-mentation and observation to examine how mortalityassociated with SF infections at natural oviposition sitesis related to the water depth in which embryos develop,how water depth at an oviposition sites is a function of variability in precipitation associated with ENSO cycles,and how outbreaks of SF infection observed in shallowwaters is mediated by exposure to UV-B radiation.Observations of embryonic mortality patterns wereconsistent with the hypothesis that climate-induced wa-ter depth fluctuations influence the exposure of devel-oping embryos to UV-B radiation. The percentage of mortality associated with SF infection was dependent onthe water depth in which embryos developed. More than50% of the western toad embryos that developed inrelatively shallow water (<20 cm) consistently devel-oped SF infections (Fig.2). However, when eggs devel-oped in water >45 cm, SF associated mortality wasnever more than 19%. Water depth at oviposition siteswas in turn related to the amount of winter precipita-tion. The amount of winter precipitation in Oregon’sNorth Cascades Mountains during 1990–1999 was itself a function of the ENSO. Results from field experimentssuggest that variation in water level influenced SF-associated mortality patterns through exposure to UV-Bradiation (Fig.2).

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