Invasive Species Compendium

Detailed coverage of invasive species threatening livelihoods and the environment worldwide

Datasheet

Batrachochytrium dendrobatidis
(Bd)

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Datasheet

Batrachochytrium dendrobatidis (Bd)

Summary

  • Last modified
  • 20 November 2018
  • Datasheet Type(s)
  • Invasive Species
  • Natural Enemy
  • Preferred Scientific Name
  • Batrachochytrium dendrobatidis
  • Preferred Common Name
  • Bd
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Fungi
  •     Phylum: Chytridiomycota
  •       Class: Chytridiomycetes
  •         Order: Rhizophydiales
  • Summary of Invasiveness
  • The zoosporic fungus Batrachochytrium dendrobatidis (Bd) is the causative agent of the infectious disease chytridiomycosis (

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Pictures

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PictureTitleCaptionCopyright
Chytridiomycosis; swabbing a North American bullfrog, farmed for the food trade in China, to test for Batrachochytrium dendrobatidis infection.
TitleSwabbing
CaptionChytridiomycosis; swabbing a North American bullfrog, farmed for the food trade in China, to test for Batrachochytrium dendrobatidis infection.
Copyright©Lisa M. Schloegel
Chytridiomycosis; swabbing a North American bullfrog, farmed for the food trade in China, to test for Batrachochytrium dendrobatidis infection.
SwabbingChytridiomycosis; swabbing a North American bullfrog, farmed for the food trade in China, to test for Batrachochytrium dendrobatidis infection.©Lisa M. Schloegel
Chytridiomycosis; swabbing the hindfoot webbing of a North American bullfrog. to test for infection by the fungus Batrachochytrium dendrobatidis.
TitleSwabbing
CaptionChytridiomycosis; swabbing the hindfoot webbing of a North American bullfrog. to test for infection by the fungus Batrachochytrium dendrobatidis.
Copyright©Lisa M. Schloegel
Chytridiomycosis; swabbing the hindfoot webbing of a North American bullfrog. to test for infection by the fungus Batrachochytrium dendrobatidis.
SwabbingChytridiomycosis; swabbing the hindfoot webbing of a North American bullfrog. to test for infection by the fungus Batrachochytrium dendrobatidis.©Lisa M. Schloegel
Chytridiomycosis; North American bullfrogs in Taiwan, farmed for the international food trade.
TitleNorth American bullfrogs in culture
CaptionChytridiomycosis; North American bullfrogs in Taiwan, farmed for the international food trade.
Copyright©Lisa M. Schloegel
Chytridiomycosis; North American bullfrogs in Taiwan, farmed for the international food trade.
North American bullfrogs in cultureChytridiomycosis; North American bullfrogs in Taiwan, farmed for the international food trade.©Lisa M. Schloegel
Chytridiomycosis; North American bullfrogs in Taiwan, farmed for the international food trade.
TitleNorth American bullfrogs in culture
CaptionChytridiomycosis; North American bullfrogs in Taiwan, farmed for the international food trade.
Copyright©Lisa M. Schloegel
Chytridiomycosis; North American bullfrogs in Taiwan, farmed for the international food trade.
North American bullfrogs in cultureChytridiomycosis; North American bullfrogs in Taiwan, farmed for the international food trade.©Lisa M. Schloegel
Chytridiomycosis; a chytrid-infected frog showing symptoms of Bd.
TitleSymptoms
CaptionChytridiomycosis; a chytrid-infected frog showing symptoms of Bd.
Copyright©Forrest Brem - This image was published in a Public Library of Science journal. PLoS journals are published under a CC BY 2.5 license
Chytridiomycosis; a chytrid-infected frog showing symptoms of Bd.
SymptomsChytridiomycosis; a chytrid-infected frog showing symptoms of Bd.©Forrest Brem - This image was published in a Public Library of Science journal. PLoS journals are published under a CC BY 2.5 license
Chytridiomycosis; close view of north American bullfrogs in Taiwan, farmed for the international food trade.
TitleNorth American bullfrogs in culture
CaptionChytridiomycosis; close view of north American bullfrogs in Taiwan, farmed for the international food trade.
Copyright©Lisa M. Schloegel
Chytridiomycosis; close view of north American bullfrogs in Taiwan, farmed for the international food trade.
North American bullfrogs in cultureChytridiomycosis; close view of north American bullfrogs in Taiwan, farmed for the international food trade.©Lisa M. Schloegel

Identity

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Preferred Scientific Name

  • Batrachochytrium dendrobatidis

Preferred Common Name

  • Bd

International Common Names

  • English: amphibian chytrid fungus; chytrid frog fungus; chytridiomycosis

Local Common Names

  • Germany: Chytridpilz

Summary of Invasiveness

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The zoosporic fungus Batrachochytrium dendrobatidis (Bd) is the causative agent of the infectious disease chytridiomycosis (Berger et al., 1998; Daszak et al., 2000). Chytridiomycosis results from a sustained cutaneous infection by B. dendrobatidis; it is an emerging infectious disease of amphibians causing mass mortality and population declines worldwide (Berger et al., 1998; Daszak et al., 2003).

Although the origin of the fungus is still disputed (see 'History of Introduction and Spread' section), the nature of Bd-associated amphibian mortalities and molecular evidence suggests its emergence and spread is a recent event.

Bd is among the world’s 100 worst invasive alien species (Lowe et al., 2004). It has been recognised by the Aquatic Animal Health Standards Comission (OIE) ad hoc group on Amphibian Diseases as one of the two pathogens of particular importance in amphibians (the other one being viruses belonging to the familiy Iridoviridae and genus Ranavirus) (Schloegel et al., 2010). In 2008, chytridiomycosis was added to the OIE’s list of notifiable diseases due to increasing evidence of the spread of the pathogen through the live amphibian trade.

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Fungi
  •         Phylum: Chytridiomycota
  •             Class: Chytridiomycetes
  •                 Order: Rhizophydiales
  •                     Genus: Batrachochytrium
  •                         Species: Batrachochytrium dendrobatidis

Diseases Table

Top of page Chytridiomycosis

Distribution

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At the time of its discovery in 1998, Bd had already achieved a global distribution and its presence has since been confirmed on every major continent except Antarctica (where amphibian fauna are not present) (Fisher et al., 2009b). It is now known to infect more than 350 of the more than 5000 recognized amphibian species (Fisher et al., 2009b). The number of infected species is likely an underestimate of the true number infected by the pathogen (owing to a lack of surveys in many regions of the world). The distribution of the fungus in Asia, for instance, is still largely un-assessed. New studies, however, are beginning to reveal the presence of Bd in this region of the world (e.g. Japan, Indonesia, Taiwan), both past and present (Kusrini et al., 2008; Goka et al., 2009; L. Schloegel et al., Wildlife Trust, New York, USA, unpublished data). Goka et al. (2009), for instance, reported the presence of Bd in preserved specimens of Andrias japonicus (Japanese giant salamander) from Japan in 1902, the earliest record known to date.

The origin of Bd is still largely disputed. (For more on this subject, including evidence from its distribution, see the 'History of Introduction and Spread' section). Histological analysis of preserved museum specimens has aided researchers in assessing the historical presence of the pathogen in some areas. Until reports by Goka et al. (2009), the oldest known record of Bd was found in a Xenopus laevis (African clawed frog) specimen from South Africa preserved in 1938 (Weldon et al., 2004), leading many to speculate that Africa was the pathogen’s place of origin. X. laevis was heavily traded for use in human pregnancy testing in the 1930s and 1940s, and is still the most widely used species for research today. Invasive populations of this amphibian have been documented in many areas. Further support for the hypothesis of African Xenopus spp. being the original source of Bd is provided by Soto-Azat et al. (2009), who studied museum specimens of frogs from Africa and South America and found Bd only in Xenopus from Africa. The distribution of X. laevis populations, however, appears to be inconsistent with the nature of Bd outbreaks, indicating that additional hosts must play a role in the spread of this disease to new locales (Rachowicz et al., 2005).

The current distribution of B. dendrobatidis is mapped on the website http://www.spatialepidemiology.net/bd/.

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

ArmeniaDisease not reportedOuellet et al., 2005
ChinaPresentBai et al., 2012
-Hong KongDisease not reported2006Rowley et al., 2007
-SichuanPresentZeng et al., 2011
IndiaPresentPresent based on regional distribution.
-KeralaLocalisedNair et al., 2011
IndonesiaPresentPresent based on regional distribution.
-JavaPresent2007Kusrini et al., 2008Mount Gede Pangrango National Park
IranDisease not reportedOuellet et al., 2005
JapanWidespreadIntroducedGoka et al., 2009
-HonshuPresent only in captivity/cultivation2007IntroducedUne et al., 2008Tokyo
Korea, Republic ofPresentYang et al., 2009
MalaysiaPresentPresent based on regional distribution.
-Peninsular MalaysiaPresentSavage et al., 2011
ThailandDisease not reportedMcLeod et al., 2008

Africa

CameroonAbsent, unreliable recordDoherty-Bone et al., 2008; Soto-Azat et al., 2009
Congo Democratic RepublicPresentGreenbaum et al., 2008Kahuzi Beiga National Park, South Kivu Province
GabonPresentBell et al., 2011
KenyaWidespreadOuellet et al., 2005; Kielgast et al., 2010
MadagascarDisease not reportedWeldon et al., 2008
MalawiAbsent, unreliable recordSoto-Azat et al., 2009
MoroccoLocalisedEl-Mouden et al., 2011
South AfricaWidespreadNativeWeldon et al., 2004
SwazilandPresentWeldon et al., 2004
UgandaPresent2006Goldberg et al., 2007Kibale National Park, Western Uganda

North America

CanadaPresentPresent based on regional distribution.
-British ColumbiaWidespreadGarner et al., 2006
-ManitobaDisease not reportedOuellet et al., 2005
-Northwest TerritoriesPresentSchock et al., 2010
-OntarioWidespreadGarner et al., 2006
MexicoLocalised2006Frías-Alvarez et al., 2008
USAPresentPresent based on regional distribution.
-AlabamaPresent2006Byrne et al., 2008
-AlaskaPresentReeves and Green, 2006
-ArizonaWidespread2006Schlaepfer et al., 2007; Picco and Collins, 2008
-ArkansasWidespread2006Rothermel et al., 2008
-CaliforniaWidespreadIntroduced Invasive Padgett-Flohr and Hopkins, 2009
-ColoradoWidespread2004Muths et al., 2008
-FloridaDisease not reported2006Rothermel et al., 2008
-GeorgiaWidespread2006Rothermel et al., 2008
-HawaiiWidespread2004IntroducedBeard and O'Neill, 2005
-IdahoWidespread2004Muths et al., 2008
-IndianaPresent1970sOuellet et al., 2005
-KansasDisease not reportedOuellet et al., 2005
-LouisianaWidespread2006Rothermel et al., 2008
-MaineWidespreadLongcore et al., 2007National wildlife refuges and federal lands
-MarylandPresentGrant et al., 2008Chesapeake and Ohio Canal National Historic Park
-MassachusettsWidespreadLongcore et al., 2007Federal refuges
-MichiganDisease not reportedOuellet et al., 2005
-MinnesotaPresent1980sOuellet et al., 2005
-MississippiDisease not reportedRothermel et al., 2008
-MissouriPresent1990-2001Ouellet et al., 2005
-MontanaWidespread2004Muths et al., 2008
-New HampshireWidespreadLongcore et al., 2007Federal refuges
-New YorkWidespreadLongcore et al., 2007
-North CarolinaWidespread2006Rothermel et al., 2008
-North DakotaDisease not reportedOuellet et al., 2005
-OregonWidespread2006Pearl et al., 2007
-PennsylvaniaPresentRaffel et al., 2010
-South CarolinaWidespread2006Rothermel et al., 2008
-TennesseeDisease not reportedRothermel et al., 2008
-TexasPresentGaertner et al., 2009Central Texas
-VermontWidespreadLongcore et al., 2007Federal refuges
-VirginiaWidespread2006Rothermel et al., 2008
-WashingtonWidespread2006Pearl et al., 2007
-WisconsinPresent1980sOuellet et al., 2005
-WyomingWidespread2006Murphy et al., 2009

Central America and Caribbean

BarbadosDisease not reportedOuellet et al., 2005
Costa RicaWidespreadPuschendorf et al., 2009
CubaPresentDíaz et al., 2007Central Cuba
DominicaPresentMalhotra et al., 2007
El SalvadorPresentLawson et al., 2011
GuatemalaPresentMendelson et al., 2004
HondurasPresentPuschendorf et al., 2006
PanamaWidespread2007Introduced Invasive Woodhams et al., 2008b; Brem and Lips, 2008
Puerto RicoPresentBurrowes et al., 2004
Trinidad and TobagoWidespreadAlemu et al., 2008Tobago

South America

ArgentinaWidespreadHerrera et al., 2005; Barrionuevo and Ponssa, 2008
BoliviaPresentBarrionuevo and Ponssa, 2008Huayramayu River, Carrasco National Park, Cochabamba
BrazilPresentPresent based on regional distribution.
-GoiasPresentGarner et al., 2006Brasilia
-Minas GeraisPresent2005Garner et al., 2006
-ParaPresent only in captivity/cultivation2008Schloegel et al., 2009
-PernambucoPresentCarnaval et al., 2006
-Rio de JaneiroPresentCarnaval et al., 2006
-Sao PauloPresent2006Schloegel et al., 2009
ChileLocalisedSolís et al., 2010
ColombiaPresent2005Ruiz and Rueda-Almonacid, 2008
EcuadorPresent1998Santiago and Merino, 2000
PeruPresent2003Seimon et al., 2007
UruguayPresent, few occurrencesMazzoni et al., 2003; Borteiro et al., 2009
VenezuelaWidespread2006Sánchez et al., 2008

Europe

AustriaPresentSztatecsny, 2008; Sztatecsny and Glaser, 2011
BelgiumPresent, few occurrencesMutschmann et al., 2000; Pasmans et al., 2010
DenmarkPresentScalera et al., 2008
FranceWidespreadGarner et al., 2006
GermanyPresentMutschmann et al., 2000; Ohst et al., 2011
ItalyWidespreadSimoncelli et al., 2005
JerseyAbsent, confirmed by surveyCunningham and Minting, 2008
LatviaDisease not reportedOuellet et al., 2005
LuxembourgLocalisedWood et al., 2009
NetherlandsPresent only in captivity/cultivationSpitzen-van der Sluijs et al., 2011
PortugalWidespread2004Garner et al., 2005
Russian FederationDisease not reportedOuellet et al., 2005
SpainWidespread2004Garner et al., 2005
SwitzerlandWidespread2004Garner et al., 2005
UKPresentGarner et al., 2006; Cunningham and Minting, 2008

Oceania

AustraliaPresentPresent based on regional distribution.
-New South WalesWidespread2005Kriger et al., 2007
-QueenslandWidespread2008Kriger and Hero, 2008; North and Alford, 2008
FijiDisease not reportedOuellet et al., 2005
New ZealandPresent2001Bell et al., 2004
Papua New GuineaDisease not reportedOuellet et al., 2005

History of Introduction and Spread

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There are two circulating theories regarding the origin of Bd (Rachowicz et al., 2005). The first, the Novel Pathogen Hypothesis (NPH), states that Bd has recently spread into new geographic regions. The second, the Endemic Pathogen Hypothesis (EPH), suggests that the fungus has been present in the environment, but that environmental change has recently altered the relationship with its amphibian hosts, leading to emergence of the disease. Molecular evidence appears to suggest that the global spread of Bd was a recent event (Morehouse et al., 2003; James et al., 2009). There is increasing data to support the notion that the anthropogenic trade in, or the introduction of, amphibians is responsible for the recent spread of Bd, with papers reporting the presence of the fungus in the pet trade, zoological collections, introduced species and the laboratory animal trade, among others (Nichols et al., 2001; Daszak et al., 2003; Une et al., 2008; Schloegel et al., 2009; Gratwicke et al., 2010; Spitzen-van der Sluijs et al., 2011).

The North American bullfrog (Rana catesbeiana, or Lithobates catesbeianus) is thought to be a carrier of Bd, exhibiting no clinical signs when experimentally or naturally infected with the fungus (Daszak et al., 2004); it is a globally traded commodity, and is sold live in markets throughout the world (Schlaepfer et al., 2005; Schloegel et al., 2009).  Sampling of native, feral and captive populations of R.catesbeiana has so far established the presence of Bd in this species on the North American, South American, European and Asian continents (Hanselmann et al., 2004; Garner et al., 2006; Goka et al., 2009; Schloegel et al., 2010). It is thought that the international movement of R.catesbeiana has served as a pathway of introduction in many regions of the world. Sequencing analyses of 59 strains of Bd from around the world found the highest degree of allelic diversity in an isolate obtained from this species, suggesting a single clonal lineage and recent spread (James et al., 2009). Further studies are currently under way to test this hypothesis.

Research in Central America indicates a North to South pattern of spread for Bd, suggesting that the pathogen is invasive in this region (Lips et al., 2006). Bd-associated anuran declines were observed in Monteverde, Costa Rica in 1987-88 and then spread at a mean rate of 26km/year before reaching El Cope, Panama in 2004 (Lips et al., 2006; Lips et al., 2008). Statistical analyses of amphibian declines in the Andean regions of South America also support the theory that Bd is an invasive pathogen, which was most likely introduced to the region in the late 1970s/early 1980s (Lips et al., 2008). Lips et al. (2008) propose that there were three separate introduction events occurring in Ecuador, Venezuela and Brazil. The mode of introduction and timing of introductions are still unknown in many regions, however; it is thought that sequencing data of Bd isolates will help to unravel this mystery.

Morgan et al. (2007) found evidence supporting both the novel and endemic pathogen hypotheses through the genotyping of isolates from the Sierra Nevada, California, USA. Additional sequencing of a more globalized dataset, however, appears to support the novel pathogen hypothesis (James et al., 2009). It is becoming evident that there is still much that is not known about the current and historical distribution of Bd. It is hoped that a catalogue of global isolates for comparative analyses will shed light on its origin and subsequent spread.

A recent genetic study by Farrer et al. (2011) identified three clades of Bd, indicated that the most common and most virulent one had emerged across five continents in the 20th century, and suggested that it was likely to have arisen as a result of contact between previously isolated populations.

Further discussion of the NPH and EPH is included in a review about B. dendrobatidis by Spitzen-van der Sluijs and Zollinger (2010).

Introductions

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Introduced toIntroduced fromYearReasonIntroduced byEstablished in wild throughReferencesNotes
Natural reproductionContinuous restocking
Balearic Islands Channel Islands 1991 Hitchhiker (pathway cause) Yes No Walker et al. (2008) Captive-bred Alytes muletenis reintroduced to Mallorca by Jersey Wildlife Preservation Trust.

Risk of Introduction

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The scale of the international trade in amphibians is in the millions of animals per year, with frogs being sold for human consumption, for pets, as bait, for experimental use in laboratories, as exhibits in zoological institutions and for bio-control purposes. (Parker, 2004; Warkentin et al., 2009). Each of these trade routes is capable of spreading Bd to new locales. While it appears that Bd is already widespread, recent studies show that different strains of the pathogen can vary in virulence (Berger et al., 2005b).  The introduction of a more virulent strain could therefore have a greater impact on the survival of amphibians in an area where a less virulent strain may already be present.

The live amphibian trade has been implicated in the spread of Bd by numerous authors (Fisher and Garner et al., 2007; Schloegel et al. 2009). Recent studies are beginning to illustrate the extent of the presence of the pathogen in trade routes. A survey of North American bullfrogs (Rana catesbeiana) farmed abroad, imported into the U.S. and sold at live markets in three major U.S. cities found that they harboured Bd infections (Schloegel et al., 2009). Bd has also been detected on bullfrog farms in Uruguay and Brazil (Mazzoni et al., 2003; Schloegel et al., 2010). Should infected individuals escape into the wild and interact with free-living species, native amphibians could be at risk of infection. The emergence of chytridiomycosis in the United Kingdom, for instance, was associated with infected, feral R. catesbeiana (Cunningham et al., 2005). A recent comparison of genetic sequences found that Bd isolates from Brazilian bullfrog farms were similar to those from wild anurans, indicating either that there has been transmisison between wild and captive hosts, or that the infections have a common source (Schloegel et al., 2010); as the farms are known not to be secure, it is likely that they at least act as reservoirs for the persistence and spread of the fungus, and possible that they may have introduced it to the country.

Bd has been detected in numerous other captive populations around the world (Berger et al., 1998; Longcore et al., 1999; Mazzoni et al., 2003; Une et al., 2008; Marantelli et al., 2004; Schloegel et al., 2009; Spitzen-van der Sluijs et al., 2011). When captive-reared animals are released intentionally (e.g. reintroduction programs), or unintentionally (e.g. the release of unwanted pets), native amphibian fauna could be at risk. Reintroduction efforts of the Mallorcan midwife toad Alytes muletensis, for instance, appear to have introduced Bd to Mallorca through the release of infected individuals (Walker et al., 2008). Ongoing studies continue to highlight the potential for the national and international movement of live amphibians to spread pathogens that could negatively impact the health and persistence of amphibian populations worldwide.

Pathogen Characteristics

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Batrachochytrium dendrobatidis (Bd) is an asexual, spherical, eukaryotic, fungal pathogen that develops in the keratinized skin cells of amphibians. As a dispersal stage, it produces swimming zoospores characteristic of the members of the Chytridiomycota. Zoospores are typically 3-5 µm in diameter, lack prominent lipid globules and have a posterior flagellum (Longcore et al., 1999). A distinguishing feature of Bd is the presence of internal septa (forming colonial thalli). Zoosporangia have thread-like rhizoids arising from single or multiple areas on the developing zoosporangium and a microtubule root beginning at kinetosome triplets 9-1 and extending parallel to the kinetosome into the aggregation of ribosomes. The kinetosome is connected to a nonflagellated centriole with overlapping fibers. The mature zoosporangium has numerous lipid globules, lacks a rumposome and transition zone plug and has one or more operculate discharge papillae (in instances where septa are present, each thallus formed by the septa has its own discharge papilla). A resting spore has not been observed for this species (Longcore et al., 1999).

B. dendrobatidis is well adapted to living in the dynamic tissue of the stratified epidermis. Sporangia live inside epidermal cells, initially parasitizing cells a few layers deep, and have a rate of development that coincides with the maturing of the cell as it moves outwards and keratinizes. They grow initially in living cells but complete their development in dead superficial keratinized cells that lack organelles. Discharge tubes have the ability to merge with and dissolve the epidermal cell membrane and open on to the surface of the cell, usually the surface distal from the body. These specialized adaptations suggest that B. dendrobatidis has long evolved to live in skin. The duration of the life cycle in vitro is 4 to 5 days at 22°C and is assumed to be the same in amphibian skin, although this has not been tested (Berger et al., 2005a, in Spitzen-van der Sluijs and Zollinger, 2010). 

B. dendrobatidis presents a wide environmental tolerance and is able to persist under a variety of temperature and precipitation regimes. The optimal condition for its growth in the laboratory is between 17 and 25°C; at 28°C growth ceases, and the organism dies after a week at above 29°C (Spitzen-van der Sluijs and Zollinger, 2010).

Habitat List

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CategorySub-CategoryHabitatPresenceStatus
Multiple

Host Animals

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Animal nameContextLife stageSystem
AnuraDomesticated host, Wild hostOther: All Stages
CaudataDomesticated host, Wild hostOther: All Stages

Climate

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ClimateStatusDescriptionRemark
A - Tropical/Megathermal climate Tolerated Average temp. of coolest month > 18°C, > 1500mm precipitation annually
Af - Tropical rainforest climate Preferred > 60mm precipitation per month
Am - Tropical monsoon climate Tolerated Tropical monsoon climate ( < 60mm precipitation driest month but > (100 - [total annual precipitation(mm}/25]))
As - Tropical savanna climate with dry summer Tolerated < 60mm precipitation driest month (in summer) and < (100 - [total annual precipitation{mm}/25])
Aw - Tropical wet and dry savanna climate Tolerated < 60mm precipitation driest month (in winter) and < (100 - [total annual precipitation{mm}/25])
BS - Steppe climate Tolerated > 430mm and < 860mm annual precipitation
Cf - Warm temperate climate, wet all year Preferred Warm average temp. > 10°C, Cold average temp. > 0°C, wet all year
Cs - Warm temperate climate with dry summer Tolerated Warm average temp. > 10°C, Cold average temp. > 0°C, dry summers
Cw - Warm temperate climate with dry winter Tolerated Warm temperate climate with dry winter (Warm average temp. > 10°C, Cold average temp. > 0°C, dry winters)
Df - Continental climate, wet all year Tolerated Continental climate, wet all year (Warm average temp. > 10°C, coldest month < 0°C, wet all year)
Ds - Continental climate with dry summer Tolerated Continental climate with dry summer (Warm average temp. > 10°C, coldest month < 0°C, dry summers)
Dw - Continental climate with dry winter Tolerated Continental climate with dry winter (Warm average temp. > 10°C, coldest month < 0°C, dry winters)
ET - Tundra climate Tolerated Tundra climate (Average temp. of warmest month < 10°C and > 0°C)

Latitude/Altitude Ranges

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Latitude North (°N)Latitude South (°S)Altitude Lower (m)Altitude Upper (m)
70 50

Air Temperature

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Parameter Lower limit Upper limit
Absolute minimum temperature (ºC) -19.6
Mean annual temperature (ºC) -2.1 26.1
Mean maximum temperature of hottest month (ºC) 10 39.8
Mean minimum temperature of coldest month (ºC) -17.6 20.9

Rainfall

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ParameterLower limitUpper limitDescription
Mean annual rainfall2904383mm; lower/upper limits

Rainfall Regime

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Summer
Uniform
Winter

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Daphnia magna Predator Spores not specific Buck et al., 2011

Pathway Vectors

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Economic Impact

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Amphibians make up a large proportion of the biomass in many tropical regions of the world. With their vibrant colours and unique life history traits, they can be a draw for many tourists. The loss of so many of the world’s anurans, and the effects of that loss on the surrounding environment, could have implications for the tourism industry upon which many people base their livelihoods (although Wollenberg et al. (2010) found that ecotourists in Madacascar, with little prior knowledge of the disease, were happy to follow preventive measures and were correctly not worried about it as a threat to their own health).

The economic impact of Bd on trade routes through the development of national and international policy is also impending. In May, 2008, the World Organization for Animal Health (OIE) listed Bd as a notifiable disease. The guidelines set forth by the OIE serve to increase awareness but may also be used as a basis for the implementation of policy in countries trading in amphibians. Efforts are already underway to incorporate OIE guidelines for Bd into legislation in the USA. Should legislation pass, it could mean that all live anuran imports would need to undergo routine quarantine and testing, among other things. With millions of amphibians imported into the USA each year (Schlaepfer et al., 2005; Gratwicke et al., 2009; Schloegel et al., 2009), the costs incurred by the implementation of quarantine standards could have a negative impact on the trade.

The economic costs of restoring disrupted ecosystems are likely to be higher than the costs of preventive or curative measures. Amphibians are important in the food chain, and their loss is detrimental to all ecosystems. For example they eat insects in large numbers, so their disappearance can cause significant problems in agriculture or human health. The costs of such losses are almost impossible to calculate, but should be taken into account (A. Spitzen, RAVON, Netherlands, personal communication, 2011).

Environmental Impact

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Impact on habitats

The presence of Bd is likely to have a significant impact on natural habitats through the loss of amphibian biodiversity. Amphibians are both aquatic and terrestrial, and any changes in their abundance will likely affect both systems. Tadpoles typically feed on algae, detritus and other animals (Whiles et al., 2006), and changes in abundance or species composition are known to alter algal community structure (Kupferberg, 1997). The presence of amphibians can impact nutrient cycling and leaf litter decomposition, plant communities and arthropod biomass (Beard et al., 2002). It is also thought that amphibians contribute to ecosystem recovery and resilience following events such as a hurricane (Beard et al., 2002). Amphibians are also an important prey item. A study in the Sierra Nevada, California, found that the presence of the Garter snake (Thamnophis elegans) is highly dependent on the availability of its primary prey item, amphibians (Jennings et al., 1992).

Impact on biodiversity

Current estimates indicate that 32% of amphibian species (1856 out of 5743) are threatened with extinction (Stuart et al., 2004). Not all the declines are Bd-related, but enigmatic declines (including disease) appear to be on the rise and are of major concern (Stuart et al., 2004). Smith et al. (2009) analyzed the patterns of amphibian declines in tropical American amphibians (an area known to have Bd-associated declines). The data indicate that those species most likely to be extirpated were those with low occupancy and a high degree of endemism. The resulting assemblage of amphibian fauna post-decline is a homogenization of species across a range of sites, thereby reducing biodiversity at the regional and global levels. There have been 9 amphibian species extinctions since 1980, an additional 113 are thought to be extinct and many more are considered to be threatened with extinction (Ron et al., 2005). These numbers are on the rise as Bd spreads into new regions and more data come to light.

Vredenburg et al. (2010) studied the dynamics of invasion of previously unexposed populations and found that it was the high growth rate and virulence of Bd that enabled it to reach high densities in all host individuals and cause population crashes before its spread was limited by reduced density of hosts.

The threatened species table contains those species for which: (1) there is a documented link between infection with Bd and declines and (2) the species is listed as threatened by the IUCN Red List of Endangered Species (i.e. Near Threatened, Vulnerable, Endangered, Critically Endangered, Extinct in the Wild or Extinct). Those species that filled the criteria for number 1, but for which an IUCN listing could not be found, were also included. 

There are many more species for which infection and/or dead and dying frogs have been documented, or which are in decline presumably due to chytridiomycosis, but detailed data linking disease with declines are lacking. Skerratt et al. (2007), for instance, state that Bd may be implicated in the decline or extinction of up to 200 species of amphibians (the most spectacular loss of vertebrate biodiversity due to disease in recorded history).

Further discussion of the importance of the impact of Bd on amphibian populations can be found in articles by Heard et al. (2011) and Duffus (2009).

Threatened Species

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Threatened SpeciesConservation StatusWhere ThreatenedMechanismReferencesNotes
Anaxyrus boreas (western toad)NT (IUCN red list: Near threatened) NT (IUCN red list: Near threatened)USAPathogenicMuths et al., 2003
Atelopus chiriquiensisCR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered)Costa RicaPathogenicBerger et al., 1998; Lips, 1998
Atelopus variusCR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered)Costa Rica; PanamaPathogenicBerger et al., 1998; Lips, 1999
Atelopus zeteki (Panamanian Golden Frog)CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered)PanamaPathogenicGewin, 2008
Bufo haematiticusLC (IUCN red list: Least concern) LC (IUCN red list: Least concern)PanamaPathogenicBerger et al., 1998; Lips et al., 2006
Craugastor azueroensisEN (IUCN red list: Endangered) EN (IUCN red list: Endangered)PanamaPathogenicLips et al., 2006
Craugastor bransfordiiLC (IUCN red list: Least concern) LC (IUCN red list: Least concern)PanamaPathogenicLips et al., 2006
Craugastor emcelaeCR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered)PathogenicBerger et al., 1998; Lips, 1999
Craugastor podiciferusNT (IUCN red list: Near threatened) NT (IUCN red list: Near threatened)PanamaPathogenicLips et al., 2006
Craugastor punctariolusEN (IUCN red list: Endangered) EN (IUCN red list: Endangered)PanamaPathogenicLips et al., 2006; Ryan et al., 2008
Craugastor rugulosusLC (IUCN red list: Least concern) LC (IUCN red list: Least concern)PathogenicBerger et al., 1998; Lips, 1999
Craugastor tabasaraeCR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered)PanamaPathogenicLips et al., 2006
Euproctus platycephalusEN (IUCN red list: Endangered) EN (IUCN red list: Endangered)ItalyPathogenicBovero et al., 2008
Gastrotheca cornutaEN (IUCN red list: Endangered) EN (IUCN red list: Endangered)Costa Rica; Panama; Colombia; EcuadorPathogenicLips et al., 2006
Hylomantis lemurCR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered)PanamaPathogenicBerger et al., 1998; Lips, 1999; Lips et al., 2006
Hyloscirtus colymbaCR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered)PanamaPathogenicBerger et al., 1998; Lips, 1999; Lips et al., 2006
Isthmohyla debilisCR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered)PathogenicBerger et al., 1998; Lips, 1999
Leiopelma archeyiCR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered)New ZealandPathogenicBell et al., 2004
Pristimantis caryophyllaceusNT (IUCN red list: Near threatened) NT (IUCN red list: Near threatened)PanamaPathogenicLips et al., 2006
Pristimantis museosusEN (IUCN red list: Endangered) EN (IUCN red list: Endangered)PanamaPathogenicLips et al., 2006
Rana muscosa (mountain yellow-legged frog)EN (IUCN red list: Endangered) EN (IUCN red list: Endangered); USA ESA listing as endangered species USA ESA listing as endangered speciesUSA; CaliforniaPathogenicUS Fish and Wildlife Service, 1999; Rachowicz et al., 2006
Rheobatrachus silusEX (IUCN red list: Extinct) EX (IUCN red list: Extinct)PathogenicRetallick et al., 2004
Rheobatrachus vitellinusEX (IUCN red list: Extinct) EX (IUCN red list: Extinct)PathogenicRetallick et al., 2004
Silverstoneia nubicolaNT (IUCN red list: Near threatened) NT (IUCN red list: Near threatened)Costa Rica; Panama; ColombiaPathogenicLips et al., 2006
Taudactylus acutirostrisCR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered)PathogenicBerger et al., 1998; Schloegel et al., 2006
Ambystoma cingulatum (frosted flatwoods salamander)VU (IUCN red list: Vulnerable) VU (IUCN red list: Vulnerable); USA ESA listing as threatened species USA ESA listing as threatened speciesFlorida; Georgia; South CarolinaPathogenicUS Fish and Wildlife Service, 2009
Eleutherodactylus cooki (guajon)VU (IUCN red list: Vulnerable) VU (IUCN red list: Vulnerable); USA ESA listing as threatened species USA ESA listing as threatened speciesPuerto RicoPest and disease transmissionUS Fish and Wildlife Service, 2004
Eleutherodactylus jasperi (golden coqui)CR (IUCN red list: Critically endangered) CR (IUCN red list: Critically endangered); USA ESA listing as threatened species USA ESA listing as threatened speciesPuerto RicoPest and disease transmissionUS Fish and Wildlife Service, 2013

Social Impact

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The various antimicrobial peptide (AMP) secretions from amphibians are thought to have numerous applications to human medicine. Many species are in decline due to infection with Bd; along with the disappearances of these species go any hopes of using their unique abilities that could greatly benefit human health. 

For example, epibatidine, extracted from the Ecuadorian poison frog (Epipedobates anthonyi) is thought to be an effective non-narcotic pain reliever (Garraffo et al., 2009) and a peptide extracted from the Chinese brown frog (Rana chensinensis) could have uses as an antimicrobial agent (Jin et al., 2009). Perhaps one of the most significant advances so far is the discovery of a series of peptides found to be effective at inhibiting HIV infection of T cells, the implications of which could be ground-breaking (Scott et al., 2005). 

Researchers believe that the Australian southern and northern gastric brooding frogs (Rheobatrachus silus and R. vitellinus) could have held a cure for the common ulcer. The fertilized eggs of these species were swallowed by the adult frogs. The gastric juices of the frog’s stomach could be halted during development, allowing the larval stage to grow without harm. These species are currently listed as extinct by the IUCN Red List of Endangered Species (Meyer et al., 2004; Hero et al., 2004).

Apart from any practical use of amphibian species, many people are likely to be distressed by their decline or disappearance (A. Spitzen, RAVON, Netherlands, personal communication, 2011).

Risk and Impact Factors

Top of page Invasiveness
  • Proved invasive outside its native range
  • Highly adaptable to different environments
  • Is a habitat generalist
  • Tolerant of shade
  • Highly mobile locally
  • Fast growing
  • Has high reproductive potential
  • Reproduces asexually
Impact outcomes
  • Altered trophic level
  • Changed gene pool/ selective loss of genotypes
  • Damaged ecosystem services
  • Ecosystem change/ habitat alteration
  • Host damage
  • Increases vulnerability to invasions
  • Infrastructure damage
  • Loss of medicinal resources
  • Modification of natural benthic communities
  • Modification of nutrient regime
  • Modification of successional patterns
  • Negatively impacts animal health
  • Negatively impacts tourism
  • Reduced native biodiversity
  • Threat to/ loss of endangered species
  • Threat to/ loss of native species
  • Negatively impacts animal/plant collections
  • Negatively impacts trade/international relations
Impact mechanisms
  • Pest and disease transmission
  • Interaction with other invasive species
  • Parasitism (incl. parasitoid)
  • Pathogenic
  • Rapid growth
Likelihood of entry/control
  • Highly likely to be transported internationally accidentally
  • Highly likely to be transported internationally illegally
  • Difficult to identify/detect as a commodity contaminant
  • Difficult to identify/detect in the field
  • Difficult/costly to control

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Woodhams DC, Ardipradja K, Alford RA, Marantelli G, Reinert LK, Rollins-Smith LA, 2007. Resistance to chytridiomycosis varies among amphibian species and is correlated with skin peptide defenses. Animal Conservation, 10(4):409-417. http://www.blackwell-synergy.com/doi/pdf/10.1111/j.1469-1795.2007.00130.x

Woodhams DC, Kilburn VL, Reinert LK, Voyles J, Medina D, Ibáñez R, Hyatt AD, Boyle DG, Pask JD, Green DM, Rollins-Smith LA, 2008. Chytridiomycosis and amphibian population declines continue to spread eastward in Panama. EcoHealth, 5(3):268-274. http://www.springerlink.com/content/u6l4775128852478/?p=49bbc10b47854c998770743b9d60a9ea&pi=5

Woodhams DC, Rollins-Smith LA, Alford RA, Simon MA, Harris RN, 2007. Innate immune defenses of amphibian skin: antimicrobial peptides and more. Animal Conservation, 10:425-428

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Zeng ZhaoHui, Bai ShiZhuo, Zhu YunQi, Wang XiaoLong, 2011. Genetic differentiation of the pathogen of Batrachochytrium dendrobatidis in toads. Journal of Economic Animal, 15(3):160-163. http://jdxb.jlau.edu.cn

Links to Websites

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WebsiteURLComment
AmphibiaWebhttp://amphibiaweb.org/
Bd-Mapshttp://www.spatialepidemiology.net/bd/
Global register of Introduced and Invasive species (GRIIS)http://griis.org/Data source for updated system data added to species habitat list.
OIE Aquatic Animal Health Codehttp://www.oie.int/fileadmin/Home/eng/Health_standards/aahc/2010/en_sommaire.htmThe aim of the Aquatic Animal Health Code is to assure the sanitary safety of international trade in aquatic animals (fish, molluscs and crustaceans) and their products.
Pet Industry Joint Advisory Council Bd-Free 'Phibs Campaignhttp://www.pijac.org/projects/project.asp?p=26

Organizations

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World: OIE (World Organisation for Animal Health), 12, rue de Prony, 75017 Paris, France, http://www.oie.int/

USA: EcoHealth Alliance, 460 West 34th Street, 17th Floor, New York NY 10001-2320, www.ecohealthalliance.org

USA: PARC - Partners in Amphibian and Reptile Conservation, www.parcplace.org

Contributors

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25/05/10 Original text by:

Lisa Schloegel, EcoHealth Alliance 460 West 34th Street, 17th floor, New York, NY 10001, USA

Peter Daszak, Wildlife Trust, USA

Distribution Maps

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