Infection with Pseudogymnoascus destructans in bats (white-nose syndrome)
- Host Animals
- Hosts/Species Affected
- Distribution Table
- List of Symptoms/Signs
- Disease Course
- Impact: Economic
- Impact: Environmental
- Impact: Social
- Zoonoses and Food Safety
- Disease Treatment
- Prevention and Control
- Links to Websites
- Distribution Maps
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IdentityTop of page
Preferred Scientific Name
- Infection with Pseudogymnoascus destructans in bats
Preferred Common Name
- white-nose syndrome
International Common Names
- English: bat white-nose syndrome; Geomyces destructans infection; Pseudogymnoascus destructans infection
OverviewTop of page
White-nose syndrome (WNS) is an emerging disease of North American bats that has caused unprecedented population declines. It is caused by the psychrophilic (cold-loving) fungus Pseudogymnoascus destructans, which is believed to have been introduced to North America from Europe or Asia (where it is present but does not cause significant mortality), although the full extent of its native range is unknown. The route of introduction is also unknown. In North America, hibernating bats become infected with P. destructans when body temperature decreases during winter torpor into the range permissive for growth of this fungus. Infected bats may develop visible fungal growth on the nose or wings, awaken more frequently from torpor, and experience a cascade of physiologic changes that result in weight loss, dehydration, electrolyte imbalances, and death. P. destructans persists in the environments of underground bat hibernation sites (hibernacula) and is believed to spread primarily by natural movements of infected bats. The first evidence of WNS in North America is from a photograph of a hibernating bat taken during winter of 2005-2006 in a hibernaculum near Albany, New York. P. destructans subsequently spread rapidly from the northeastern United States throughout much of the eastern portions of the United States and Canada, and most recently (as of May 2017) was detected in Washington State. It has killed millions of bats, threatening some species with regional extirpation and putting at risk the valuable environmental services that bats provide by eating harmful insects.
Host AnimalsTop of page
|Animal name||Context||Life stage||System|
|Barbastella barbastellus||Wild host|
|Eptesicus fuscus||Wild host|
|Eptesicus nilssonii||Wild host|
|Miniopterus schreibersii||Wild host|
|Myotis austroriparius||Wild host|
|Myotis bechsteinii||Wild host|
|Myotis brandtii||Wild host|
|Myotis dasycneme||Wild host|
|Myotis daubentonii||Wild host|
|Myotis emarginatus||Wild host|
|Myotis grisescens||Wild host|
|Myotis leibii||Wild host|
|Myotis lucifugus||Wild host|
|Myotis myotis||Wild host|
|Myotis nattereri||Wild host|
|Myotis petax||Wild host|
|Myotis septentrionalis||Wild host|
|Myotis sodalis||Wild host|
|Myotis velifer||Wild host|
|Myotis volans||Wild host|
|Myotis yumanensis||Wild host|
|Perimyotis subflavus||Wild host|
|Plecotus auritus||Wild host|
|Rhinolophus euryale||Wild host|
|Rhinolophus hipposideros||Wild host|
Hosts/Species AffectedTop of page
White-nose syndrome (WNS) has been diagnosed in a diversity of bat species from North America, Europe, and Asia (Blehert et al., 2009; Turner et al., 2011; Pikula et al., 2012; Zukal et al., 2014; Hoyt et al., 2016; Zukal et al., 2016). In addition, P. destructans has been detected on a number of other bat species from these areas without documented signs of WNS (Martinkova et al., 2010; Wibbelt et al., 2010; Bernard et al., 2015; Hoyt et al., 2016). It is not known at this time why some North American species of bats experience more severe disease or higher mortality from WNS than others, nor what resistance mechanisms result in less severe disease in some species, although certain host traits and hibernation conditions are hypothesized to play a role (Flory et al., 2012; Hayman et al., 2016).
The Host Animals table lists species that, as of August 2018, were confirmed to demonstrate diagnostic evidence of white-nose syndrome associated with P. destructans infection. The fungus has been detected on a number of other bat species without histopathologic evidence of clinical disease; these species are not included in the table. An updated list of all bats that are either affected by white-nose syndrome or potentially carry the fungus is maintained at: https://www.whitenosesyndrome.org/about/bats-affected-wns.
Isolates of P. destructans from both North America and Europe have similar virulence in the North American little brown bat (Myotis lucifugus) (Warnecke et al., 2012). However, while skin infection with P. destructans has been detected in a number of European bat species, mass mortality attributable to infection has not been observed (Wibbelt et al., 2010; Puechmaille et al., 2011a; Puechmaille et al., 2011b; Pikula et al., 2012; Zukal et al., 2014).
DistributionTop of page
Pseudogymnoascus destructans has been detected on bats or in environments utilized by bats for hibernation in North America (United States and Canada), at least 18 European countries, and China. The native range of the fungus is unknown but is minimally presumed to include parts of Europe, where there is evidence of genetic diversification (Leopardi et al., 2015). The route of introduction to North America has not been determined. The disease has now been reported from 5 provinces in eastern Canada and 31 US states, mostly in the centre and east of the country; the fungus, but no clinical disease, has been found in a further two states. An interactive map of the spread and distribution of WNS in North America is available at: https://www.sciencebase.gov/gisviewer/wns/; the latest map of distribution, along with earlier versions, can be seen at https://www.whitenosesyndrome.org/resources/map.
Distribution TableTop of page
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.Last updated: 10 Jan 2020
PathologyTop of page
Gross signs of infection by Pseudogymnoascus destructans can include white, powdery growth of fungal hyphae and conidia on the muzzle and a white, tacky, opaque film on the wings. However, visible fungal growth is not always seen (Meteyer et al., 2009). Changes to the skin of affected bats can be variable, including: patches of rough skin on the face, ears, forearms, wing membrane, and feet; pinpoint white foci that resemble comedones on the muzzle; loss of sheen on glabrous skin; and irregular pigmentation with areas of contraction or tears in the wing membrane (Meteyer et al., 2009). Bats may be in poor body condition with inadequate fat stores but do not demonstrate evidence of major organ failure or other consistent non-cutaneous pathologies (Courtin et al., 2010).
Direct cutaneous penetration and proliferation by hyphae of P. destructans causes characteristic epidermal erosions and ulcers (Meteyer et al., 2009). Unlike most transmissible dermatophytes, P. destructans invades the underlying connective tissue (Meteyer et al., 2009; Reichard and Kunz, 2009; Cryan et al., 2010). Cup-like epidermal erosions filled with fungal hyphae, or ulceration and fungal invasion of underlying connective tissue, are often observed on histopathology (Meteyer et al., 2009). Fungal hyphae in tissue sections are branching and septate with curved conidia on short aerial hyphae, although the latter structures are not always observed (Meteyer et al., 2009).
A notable observation on histopathology is the absence of inflammation in skin from hibernating bats, even when there is evidence for extensive invasion by P. destructans (Meteyer et al., 2009). Analysis of gene expression during infection reveals that infected hibernating bats initiate a partial immune response to fungal invasion, including activation of acute inflammation and wound healing pathways, but recruitment of white blood cells to affected sites for clearing infection is inadequate during torpor (Rapin et al., 2014; Field et al., 2015). Paradoxically, damage to the wing membranes is most notable in bats examined just after spring emergence from hibernation, which may be attributable to immune-mediated damage to invaded tissues on return to euthermia (Meteyer et al., 2012).
DiagnosisTop of page
Pseudogymnoascus destructans can be detected in environmental substrates from bat hibernation environments even when bats are absent. The fungus can additionally be detected on bats in the absence of clinical signs of disease. Thus the case definition for white-nose syndrome (WNS) distinguishes between detection of the pathogen by culture or molecular methods and confirmation of WNS through a combination of pathogen detection and observation of histologic signs of skin infection (USGS National Wildlife Health Center, 2015). Detection of P. destructans in the absence of histopathologic support or other field observations can only confirm that the fungus is present in a population or location.
P. destructans can be grown from a variety of sample types (e.g., soil, skin biopsies, and skin swabs), by culture on Sabouraud dextrose agar at 7°C (Gargas et al., 2009; Lorch et al., 2013a), or by polymerase chain reaction (PCR) targeting the intergenic spacer region of the rRNA gene complex (Muller et al., 2013). Confirmation of skin infection with fungal invasion characteristic of clinical WNS requires microscopic examination of skin cross-sections (Meteyer et al., 2009). When screening for presence of P. destructans, PCR is preferred over culture due to greater sensitivity (USGS National Wildlife Health Center, 2015).
Field signs that a hibernating bat population may be affected by WNS include: excessive or unexplained mortality at or near a hibernaculum; visible fungal growth on the muzzle or wings of live or freshly dead bats; abnormal daytime activity or movement towards hibernaculum openings; and moderate to severe wing damage in non-torpid bats in thin body condition (although the latter two signs are both nonspecific when observed in isolation) (USGS National Wildlife Health Center, 2015). In the field, ultraviolet light can also be used to screen hibernating bats for potential infection. Long-wave ultraviolet light elicits a distinct orange-yellow fluorescence when the fungus has invaded skin tissues (Turner et al., 2014). Observation of this characteristic fluorescence can be a valuable screening tool for guiding sample collection for laboratory testing (Turner et al., 2014). Field observations compatible with WNS, or detection of P. destructans by culture or molecular methods, can be paired with observation of UV fluorescence to support a diagnosis of suspected WNS.
List of Symptoms/SignsTop of page
|General Signs / Dehydration||Other:All Stages||Sign|
|General Signs / Underweight, poor condition, thin, emaciated, unthriftiness, ill thrift||Other:All Stages||Sign|
|General Signs / Weight loss||Other:All Stages||Sign|
|Nervous Signs / Abnormal behavior, aggression, changing habits||Other:All Stages||Sign|
|Nervous Signs / Excessive or decreased sleeping||Other:All Stages||Sign|
|Skin / Integumentary Signs / Scarred skin||Other:All Stages||Sign|
|Skin / Integumentary Signs / Skin crusts, scabs||Other:All Stages||Sign|
|Skin / Integumentary Signs / Skin ulcer, erosion, excoriation||Other:All Stages||Diagnosis|
Disease CourseTop of page
Pathogenesis of white-nose syndrome (WNS) has emerged as a unique, multi-stage model whereby a skin infection progresses to causing abnormal behaviors and systemic illness due to physiological changes experienced by infected bats during winter torpor. Infection leading to disease is limited to hibernating bats whose body temperature is low enough to allow fungal growth and invasion (Cryan et al., 2010; Verant et al., 2012). Infected bats may develop visible fungal growth around the nose or on the wings as the fungus invades cutaneous tissues (although this sign is not always observed). This visible white growth on hibernating bats tends to be noticeable after midwinter, but phenology of hyphal and conidiophore growth above skin surfaces is not well characterized. Infected bats may also demonstrate abnormal hibernation behaviors, such as more frequent arousal from torpor, moving among roosting sites or congregating near hibernaculum entrances, and flying by day outside of hibernation sites (Castle and Cryan, 2010; Cryan et al., 2010; Frick et al., 2010; Blehert, 2012; Reeder et al., 2012;). Subsequently, infected bats experience a cascade of physiological changes that result in weight loss, dehydration, and electrolyte imbalances, potentially culminating in death (Willis et al., 2011; Cryan et al., 2013a; Verant et al., 2014).
Bats can recover from WNS in a captive setting following removal from torpor and provision of supportive care. Rehabilitation of individual animals, however, is a labour intensive process that is not broadly applicable to management of free-ranging bat populations (Fuller et al., 2011; Meteyer et al., 2011). Upon emergence from hibernation in spring, infected bats can naturally recover from WNS without human intervention, but individuals with extensive infections are less likely to survive the healing process (Dobony et al., 2011; Fuller et al., 2011). Following arousal from hibernation, the immune system response against Pseudogymnoascus destructans, which is not observed in torpid animals, can be so vigorous as to fatally impair the host (Meteyer et al., 2012).
EpidemiologyTop of page
White-nose syndrome (WNS) is primarily observed during winter months when bat skin temperatures are conducive to growth of Pseudogymnoascus destructans (Blehert, 2012; Verant et al., 2012; Langwig et al., 2015a). Infected bats are hypothesized to be a primary mechanism for moving the fungus among hibernation sites (Lorch et al., 2011; Lorch et al., 2013b; Hoyt et al., 2015b). Prevalence of P. destructans in bat populations peaks during late winter, but the fungus can be detected on bats at lower prevalence through early spring emergence; pathogen prevalence declines during summer months (Langwig et al., 2015a). The fungus can also persist outside the host for extended periods in substrates from hibernation sites, meaning that once infested, hibernacula serve as persistent reservoirs of the pathogen (Lorch et al., 2013b; Reynolds et al., 2015).
P. destructans can also likely be transferred among locations by humans on clothing, footwear, or equipment (Sleeman, 2011; Shelley et al., 2013; Ballmann et al., 2017). Restricting movement of these items from areas affected by WNS to unaffected areas and attention to decontamination is strongly advised as a precautionary principle for persons who visit potential bat hibernation habitats (e.g. recreational cavers, biologists, or miners). Guidance for decontaminating and restricting movement of equipment used in bat hibernation sites has been developed to reduce risks of anthropogenic contributions to the spread of WNS: https://www.whitenosesyndrome.org/resources/cavers.
Impact: EconomicTop of page
Millions of North American bats have died from white-nose syndrome (WNS), resulting in dramatic regional bat population declines (Frick et al., 2010; Frick et al., 2015; Reynolds et al., 2016). Insectivorous bats provide valuable pest control services. For example, bats eat insects that damage crops and forests, and that carry diseases. In the USA alone, insectivorous bats are estimated to save farmers over $3 billion annually in pest suppression services (Boyles et al., 2011). Recent experiments indicate that bat suppression of insect pests in corn (maize) fields could scale globally to one billion dollars in savings per year for that crop alone (Maine and Boyles, 2015). Many bat species in tropical and subtropical regions are also important pollinators and dispersers of seeds (Boyles et al., 2011; Kunz et al., 2011). Mortality due to WNS and other population stressors is expected to reduce the levels at which bat populations provide these valuable and irreplaceable ecosystem services.
Impact: EnvironmentalTop of page
White-nose syndrome affects all life stages of hibernating bats, and mortality at newly-affected hibernacula can be very high, resulting in substantial and rapid decreases in bat abundance (Frick et al., 2010). Millions of North American bats have died from WNS, and population declines for heavily impacted species could result in regional extirpation of some previously common species such as the little brown bat (Myotis lucifugus) and northern long-eared bat (M. septentrionalis) (Frick et al., 2010; Frick et al., 2015; Reynolds et al., 2016; Brooks, 2011; Thogmartin et al., 2013; Erickson et al., 2016).
Bat populations affected by WNS are expected to be slow to recover due to low annual fecundity, and some populations may struggle to achieve the numbers seen prior to emergence of this disease (Russell et al., 2015). Cumulative effects of other population stressors, such as mortality from wind-turbine collisions, could exacerbate disease impacts (Erickson et al., 2016). Long-term monitoring of bat populations will be vital for understanding recovery trajectories and guiding evidence-based management decisions (Russell et al., 2015).
Additionally, shifts in bat community composition and abundance have been documented in areas of the United States and Canada that have been affected by WNS (Francl et al., 2012; Frick et al., 2015). Although the implications of these declines and populations shifts are not fully understood, they are likely to exert ecological impacts (Brooks, 2011; Frick et al., 2015).
Large-scale impacts of WNS are confined to North America. Isolates of P. destructans from both North America and Europe have similar virulence in the North American little brown bat (Myotis lucifugus) (Warnecke et al., 2012). However, while skin infection with P. destructans has been detected in a number of European bat species, mass mortality attributable to infection has not been observed (Wibbelt et al., 2010; Puechmaille et al., 2011a; Puechmaille et al., 2011b; Pikula et al., 2012; Zukal et al., 2014).
Impact: SocialTop of page
Some bat species, such as the northern long-eared bat (M. septentrionalis), may consume large quantities of mosquitoes (Reiskind and Wund, 2009), and can therefore potentially benefit human health by reducing risks for transmission of vector-borne disease. Large-scale mortality due to WNS is expected to reduce these benefits.
Zoonoses and Food SafetyTop of page
Direct human health implications of Pseudogymnoascus destructans are unknown. However, there have been no reports to date of P. destructans infection occurring in human or animal species other than bats. Furthermore, human skin and core temperatures exceed the range permissive to growth of P. destructans.
Disease TreatmentTop of page
Captive experiments have demonstrated that individual animals removed from torpor can recover from infection through provision of appropriate supportive care (Meteyer et al., 2011). The fungus is also susceptible to a range of common antifungal medications that are used to treat other fungal infections through topical or systemic administration (Chaturvedi et al., 2011). However, appropriate dosing and tolerance of these medications is not established in bats. While treatment and rehabilitation of individual animals is possible (Meteyer et al., 2011) and could be deployed as a targeted intervention for critically endangered bat species that tolerate captivity, individualized disease treatment can make only small contributions to mitigating the impacts of white-nose syndrome on a population level.
Prevention and ControlTop of page
The U.S. Fish and Wildlife Service and the Canadian Wildlife Health Cooperative coordinate national and international partnerships to investigate and respond to white-nose syndrome (WNS) (U.S. Fish and Wildlife Service, 2011; Canadian Wildlife Health Cooperative, 2015). These interagency efforts have fostered considerable collaboration that led to rapid progress in monitoring and understanding the spread of WNS, characterizing Pseudogymnoascus destructans as a pathogen, developing effective diagnostic approaches, organizing multi-agency working groups to encourage collaboration, and developing federal-state surveillance partnerships to track pathogen spread (Voyles et al., 2015; Lankau and Moede Rogall, 2016).
Management of hibernacula, including restriction of human access to some sites, and development of procedures and requirements to disinfect caving equipment, was instituted in partnership with recreational caving communities to limit anthropogenic spread of P. destructans (Sleeman, 2011; Shelley et al., 2013; Ballmann et al., 2017). Preventing spread by natural movement of bats is not feasible given the mobility of free-ranging wildlife. Additionally, culling of infected individuals within a hibernaculum does not reduce morbidity and mortality from WNS because of the rapid spread of the fungus within dense populations and because of the role of environmental substrates as a persistent reservoir of P. destructans (Foley et al., 2011; Hallam and McCracken, 2011; Crozier and Schulte-Hostedde, 2014; Meyer et al., 2016).
Current research efforts to reduce bat population losses and support their recovery are increasingly focused on developing mitigation strategies to limit the expansion of WNS. Understanding of the pathobiology of P. destructans infection suggests avenues that could reduce prevalence or severity of clinical disease in hibernating bats. For example, a topic of active investigation is the potential to reduce burdens of P. destructans in hibernation environments to lessen morbidity and mortality of bats at a population level through application of biological- or chemical-control strategies (Cornelison et al., 2014a; Cornelison et al., 2014b; Frank et al., 2014; Hoyt et al., 2015a; Zhang et al., 2015; Boire et al., 2016; Cheng et al., 2016; Frank et al., 2016). Additionally, development of an oral vaccine that could be used to immunize bat populations against infection by P. destructans is underway; proof-of-concept testing of a candidate recombinant raccoonpox virus platform for oral immunization of bats was recently completed (Stading et al., 2016).
ReferencesTop of page
Alves DMCC, Terribile LC, Brito D, 2014. The potential impact of white-nose syndrome on the conservation status of North American bats. PLoS ONE, 9(9):e107395. http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0107395
Ballmann, A. E., Torkelson, M. R., Bohuski, E. A., Russell, R. E., Blehert, D. S., 2017. Dispersal hazards of Pseudogymnoascus destructans by bats and human activity at hibernacula in summer. Journal of Wildlife Diseases, 53(4), 725-735. http://www.bioone.org/loi/jwdi doi: 10.7589/2016-09-206
Barlow AM, Worledge L, Miller H, Drees KP, Wright P, Foster JT, Sobek C, Borman AM, Fraser M, 2015. First confirmation of Pseudogymnoascus destructans in British bats and hibernacula. Veterinary Record, 177(3):73. http://veterinaryrecord.bvapublications.com/archive/
Bernard RF, Foster JT, Willcox EV, Parise KL, McCracken GF, 2015. Molecular detection of the causative agent of white-nose syndrome on Rafinesque's big-eared bats (Corynorhinus rafinesquii) and two species of migratory bats in the southeastern USA. Journal of Wildlife Diseases, 51(2):519-522. http://www.jwildlifedis.org/doi/full/10.7589/2014-08-202
Blehert DS, Hicks AC, Behr M, Meteyer CU, Berlowski-Zier BM, Buckles EL, Coleman JTH, Darling SR, Gargas A, Niver R, Okoniewski JC, Rudd RJ, Stone WB, 2009. Bat white-nose syndrome: An emerging fungal pathogen? Science, 323(5911):227
Boire N, Zhang S, Khuvis J, Lee R, Rivers J, Crandall P, Keel MK, Parrish N, 2016. Potent inhibition of Pseudogymnoascus destructans, the causative agent of White-nose Syndrome in bats, by cold-pressed, terpeneless, Valencia orange oil. PLoS ONE, 11(2):e0148473. http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0148473
Brack V Jr, 2007. Temperatures and locations used by hibernating bats, including Myotis sodalis (Indiana bat), in a limestone mine: implications for conservation and management. Environmental Management, 40(5):739-746. http://www.springerlink.com/content/p7545307343m2645/?p=a3d536471fa14277ad1d157fd4c89996&pi=1
Brooks RT, 2011. Declines in summer bat activity in central New England 4 years following the initial detection of white-nose syndrome. Biodiversity and Conservation, 20(11):2537-2541. http://www.springerlink.com/content/9716632784163261/
Canadian Wildlife Health Cooperative, 2015. A national Plan to manage white nose syndrome in bats in Canada. Saskatoon, Saskatchewan, Canada: Canadian Wildlife Health Cooperative, 14 pp. http://www.cwhc-rcsf.ca/docs/BatWhiteNoseSyndrome-NationalPlan.pdf
Carpenter GM, Willcox EV, Bernard RF, Stiver WH, 2016. Detection of Pseudogymnoascus destructans on free-flying male bats captured during summer in the Southeastern USA. Journal of Wildlife Diseases, 52(4):922-926. http://www.bioone.org/loi/jwdi
Castle KT, Cryan PM, 2010. White-nose syndrome in bats: a primer for resource managers. Park Science, 27(1):20-25
Chaturvedi S, Rajkumar SS, Li XJ, Hurteau GJ, Shtutman M, Chaturvedi V, 2011. Antifungal testing and high-throughput screening of compound library against Geomyces destructans, the etiologic agent of geomycosis (WNS) in bats. PLoS ONE, No.March:e17032. http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0017032
Chaturvedi V, Springer DJ, Behr MJ, Ramani R, Li XJ, Peck MK, Ren P, Bopp DJ, Wood B, Samsonoff WA, Butchkoski CM, Hicks AC, Stone WB, Rudd RJ, Chaturvedi S, 2010. Morphological and molecular characterizations of psychrophilic fungus Geomyces destructans from New York bats with white nose syndrome (WNS). PLoS ONE, No.May:e10783. http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0010783
Cheng TL, Mayberry H, McGuire LP, Hoyt JR, Langwig KE, Nguyen H, Parise KL, Foster JT, Willis CKR, Kilpatrick AM, Frick WF, 2016. Efficacy of a probiotic bacterium to treat bats affected by the disease white-nose syndrome. Journal of Applied Ecology. http://dx.doi.org/10.1111/1365-2664.12757
Cornelison CT, Gabriel KT, Barlament C, Crow SA Jr, 2014. Inhibition of Pseudogymnoascus destructans growth from conidia and mycelial extension by bacterially produced volatile organic compounds. Mycopathologia, 177(1/2):1-10. http://rd.springer.com/journal/11046
Cornelison CT, Keel MK, Gabriel KT, Barlament CK, Tucker TA, Pierce GE, Crow SA, 2014. A preliminary report on the contact-independent antagonism of Pseudogymnoascus destructans by Rhodococcus rhodochrous strain DAP96253. BMC Microbiology, 14:26
Courtin F, Stone WB, Risatti G, Gilbert K, Kruiningen HJ van, 2010. Pathologic findings and liver elements in hibernating bats with white-nose syndrome. Veterinary Pathology, 47(2):214-219. http://vet.sagepub.com/content/by/year
Crozier GKD, Schulte-Hostedde AI, 2014. The ethical dimensions of wildlife disease management in an evolutionary context. Evolutionary Applications, 7(7):788-798. http://onlinelibrary.wiley.com/journal/10.1111/(ISSN)1752-4571
Cryan PM, Meteyer CU, Blehert DS, Lorch JM, Reeder DM, Turner GG, Webb J, Behr M, Verant M, Russell RE, Castle KT, 2013. Electrolyte depletion in white-nose syndrome bats. Journal of Wildlife Diseases, 49(2):398-402. http://www.jwildlifedis.org/content/49/2/398.full
Cryan PM, Meteyer CU, Boyles JG, Blehert DS, 2010. Wing pathology of white-nose syndrome in bats suggests life-threatening disruption of physiology. BMC Biology, 8(135):(11 November 2010). http://www.biomedcentral.com/1741-7007/8/135
Cryan PM, Meteyer CU, Boyles JG, Blehert DS, 2013. White-nose syndrome in bats: illuminating the darkness. BMC Biology, 11:47
Dobony CA, Hicks AC, Langwing KE, Linden RI von, Okoniewski JC, Rainbolt RE, 2011. Little brown myotis persist despite exposure to white-nose syndrome. Journal of Fish and Wildlife Management, 2(2):190-195
Drees KP, Palmer JM, Sebra R, Lorch JM, Chen C, Wu ChengCang, Bok JinWoo, Keller NP, Blehert DS, Cuomo CA, Lindner DL, Foster JT, 2016. Use of multiple sequencing technologies to produce a high-quality genome of the fungus Pseudogymnoascus destructans, the causative agent of bat white-nose syndrome. Genome Announcements, 4(3):e00445-16. http://genomea.asm.org/content/4/3/e00445-16.abstract
Erickson RA, Thogmartin WE, Diffendorfer JE, Russell RE, Szymanski JA, 2016. Effects of wind energy generation and white-nose syndrome on the viability of the Indiana bat. Peerj, 4:e2830
Escobar LE, Lira-Noriega A, Medina-Vogel G, Peterson AT, 2014. Potential for spread of the white-nose fungus (Pseudogymnoascus destructans) in the Americas: use of maxent and nichea to assure strict model transference. Geospatial Health, 9(1):221-229. http://www.geospatialhealth.net/index.php/gh/article/view/19/19
Field KA, Johnson JS, Lilley TM, Reeder SM, Rogers EJ, Behr MJ, Reeder DM, 2015. The white-nose syndrome transcriptome: activation of anti-fungal host responses in wing tissue of hibernating little brown myotis. PLoS Pathogens, 11(10):e1005168. http://journals.plos.org/plospathogens/article?id=10.1371/journal.ppat.1005168
Fisher MC, Henk DA, Briggs CJ, Brownstein JS, Madoff LC, McCraw SL, Gurr SJ, 2012. Emerging fungal threats to animal, plant and ecosystem health. Nature (London), 484(7393):186-194. http://www.nature.com/nature
Flieger M, Bandouchova H, Cerny J, Chudickova M, Kolarik M, Kovacova V, Martinkova N, Novak P, Sebesta O, Stodulkova E, Pikula J, 2016. Vitamin B-2 as a virulence factor in Pseudogymnoascus destructans skin infection. Scientific Reports, 6:33200
Flory AR, Kumar S, Stohlgren TJ, Cryan PM, 2012. Environmental conditions associated with bat white-nose syndrome mortality in the north-eastern United States. Journal of Applied Ecology, 49(3):680-689. http://onlinelibrary.wiley.com/journal/10.1111/(ISSN)1365-2664
Foley J, Clifford D, Castle K, Cryan P, Ostfeld RS, 2011. Investigating and managing the rapid emergence of white-nose syndrome, a novel, fatal, infectious disease of hibernating bats. Conservation Biology, 25(2):223-231. http://onlinelibrary.wiley.com/journal/10.1111/(ISSN)1523-1739
Francl KE, Ford WM, Sparks DW, Brack V, 2012. Capture and reproductive trends in summer bat communities in West Virginia: Assessing the impact of white-nose syndrome. Journal of Fish and Wildlife Management, 3(1):33-42
Frank CL, Ingala MR, Ravenelle RE, Dougherty-Howard K, Wicks SO, Herzog C, Rudd RJ, 2016. The effects of cutaneous fatty acids on the growth of Pseudogymnoascus destructans, the etiological agent of White-Nose Syndrome (WNS). PLoS ONE, 11(4):e0153535. http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0153535
Frank CL, Michalski A, McDonough AA, Rahimian M, Rudd RJ, Herzog C, 2014. The resistance of a North American bat species (Eptesicus fuscus) to white-nose syndrome (WNS). PLoS One, 9(12):e113958
Frick WF, Pollock JF, Hicks AC, Langwig KE, Reynolds DS, Turner GG, Butchkoski CM, Kunz TH, 2010. An emerging disease causes regional population collapse of a common North American bat species. Science (Washington), 329(5992):679-682. http://www.sciencemag.org
Frick WF, Puechmaille SJ, Hoyt JR, Nickel BA, Langwig KE, Foster JT, Barlow KE, Bartonicka T, Feller D, Haarsma AJ, Herzog C, Horácek I, Kooij J van der, Mulkens B, Petrov B, Reynolds R, Rodrigues L, Stihler CW, Turner GG, Kilpatrick AM, 2015. Disease alters macroecological patterns of North American bats. Global Ecology and Biogeography, 24(7):741-749. http://onlinelibrary.wiley.com/journal/10.1111/(ISSN)1365-2486
Froschauer A, Coleman J, 2012. News release: North American bat death toll exceeds 5.5 million from white-nose syndrome. Arlington, Virginia, USA: U.S. Fish and Wildlife Service, 2 pp. https://www.whitenosesyndrome.org/sites/default/files/files/wns_mortality_2012_nr_final_0.pdf
Fuller NW, Reichard JD, Nabhan ML, Fellows SR, Pepin LC, Kunz TH, 2011. Free-ranging little brown myotis (Myotis lucifugus) heal from wing damage associated with white-nose syndrome. EcoHealth, 8(2):154-162. http://www.springerlink.com/content/r11rlp2u53268426/
Hallam TG, Federico P, 2012. The panzootic white-nose syndrome: an environmentally constrained disease? Transboundary and Emerging Diseases, 59(3):269-278. http://onlinelibrary.wiley.com/doi/10.1111/j.1865-1682.2011.01268.x/full
Hallam TG, McCracken GF, 2011. Management of the panzootic white-nose syndrome through culling of bats. Conservation Biology, 25(1):189-194. http://onlinelibrary.wiley.com/journal/10.1111/(ISSN)1523-1739
Hayman DTS, Pulliam JCR, Marshall JC, Cryan PM, Webb CT, 2016. Environment, host and fungal traits predict continental scale white-nose syndrome in bats. Science Advances, 2:e1500831
Hoyt JR, Cheng TL, Langwig KE, Hee MM, Frick WF, Kilpatrick AM, 2015. Bacteria isolated from bats inhibit the growth of Pseudogymnoascus destructans, the causative agent of white-nose syndrome. PLoS ONE, 10(4):e0121329. http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0121329
Hoyt JR, Langwig KE, Okoniewski J, Frick WF, Stone WB, Kilpatrick AM, 2015. Long-term persistence of Pseudogymnoascus destructans, the causative agent of white-nose syndrome, in the absence of bats. EcoHealth, 12(2):330-333. http://link.springer.com/article/10.1007%2Fs10393-014-0981-4
Hoyt JR, Sun KePing, Parise KL, Lu GuanJun, Langwig KE, Jiang TingLei, Yang ShuBao, Frick WF, Kilpatrick AM, Foster JT, Feng Jiang, 2016. Widespread bat white-nose syndrome fungus, northeastern China. Emerging Infectious Diseases, 12(1):140-142. http://wwwnc.cdc.gov/eid/article/22/1/pdfs/15-1314.pdf
IUCN, 2016. The IUCN (the International Union for Conservation of Nature) Red List of Threatened Species. http://www.iucnredlist.org/
Khankhet J, Vanderwolf KJ, McAlpine DF, McBurney S, Overy DP, Slavic D, Xu J, 2014. Clonal expansion of the Pseudogymnoascus destructans genotype in North America is accompanied by significant variation in phenotypic expression. PLoS One, 9(8):e104684
Knudsen GR, Dixon RD, Amelon SK, 2013. Potential spread of white-nose syndrome of bats to the northwest: epidemiological considerations. Northwest Science, 87(4):292-306. http://www.bioone.org/doi/abs/10.3955/046.087.0401
Kunz TH, Torrez EB de, Bauer D, Lobova T, Fleming TH, 2011. Ecosystem services provided by bats. Annals of the New York Academy of Sciences, 1223:1-38
Langwig KE, Frick WF, Reynolds R, Parise KL, Drees KP, Hoyt JR, Cheng TL, Kunz TH, Foster JT, Kilpatrick AM, 2015. Host and pathogen ecology drive the seasonal dynamics of a fungal disease, white-nose syndrome. Proceedings of the Royal Society of London. Series B, Biological Sciences, 282(1799):20142335. http://rspb.royalsocietypublishing.org/content/282/1799/20142335
Langwig KE, Voyles J, Wilber MQ, Frick WF, Murray KA, Bolker BM, Collins JP, Cheng TL, Fisher MC, Hoyt JR, Lindner DL, McCallum HI, Puschendorf R, Rosenblum EB, Toothman M, Willis CK, Briggs CJ, Kilpatrick AM, 2015. Context-dependent conservation responses to emerging wildlife diseases. Frontiers in Ecology and the Environment, 13(4):195-202
Lankau EW, Moede Rogall G, 2016. White-nose syndrome in North American bats - U.S. Geological Survey updates. Madison, Wisconsin, USA: USGS National Wildlife Health Center, 4 pp. [Fact Sheet 2016-3084.] https://pubs.usgs.gov/fs/2016/3084/fs20163084.pdf
Leopardi S, Blake D, Puechmaille SJ, 2015. White-nose syndrome fungus introduced from Europe to North America. Current Biology, 25(6):R217-R219
Lorch JM, Lindner DL, Gargas A, Muller LK, Minnis AM, Blehert DS, 2013. A culture-based survey of fungi in soil from bat hibernacula in the eastern United States and its implications for detection of Geomyces destructans, the causal agent of bat white-nose syndrome. Mycologia, 105(2):237-252
Lorch JM, Meteyer CU, Behr MJ, Boyles JG, Cryan PM, Hicks AC, Ballmann AE, Coleman JTH, Redell DN, Reeder DM, Blehert DS, 2011. Experimental infection of bats with Geomyces destructans causes white-nose syndrome. Nature (London), 480(7377):376-378. http://www.nature.com/nature
Lorch JM, Muller LK, Russell RE, O'Connor M, Lindner DL, Blehert DS, 2013. Distribution and environmental persistence of the causative agent of white-nose syndrome, geomyces destructans, in bat hibernacula of the eastern United States. Applied and Environmental Microbiology, 79(4):1293-1301. http://aem.asm.org/content/79/4/1293.abstract
Lorch JM, Palmer JM, Lindner DL, Ballmann AE, George KG, Griffin K, Knowles S, Huckabee JR, Haman KH, Anderson CD, Becker PA, Buchanan JB, Foster JT, Blehert DS, 2016. First detection of bat white-nose syndrome in western North America. mSphere, 1(4):00148-16. http://msphere.asm.org/content/1/4/e00148-16
Maher SP, Kramer AM, Pulliam JT, Zokan MA, Bowden SE, Barton HD, Magori K, Drake JM, 2012. Spread of white-nose syndrome on a network regulated by geography and climate. Nature Communications, 3:1306
Maine JJ, Boyles JG, 2015. Bats initiate vital agroecological interactions in corn. Proceedings of the National Academy of Sciences of the United States of America, 112(40):12438-12443. http://www.pnas.org/content/112/40/12438.full
Martínková N, Backor P, Bartonicka T, Blazková P, Cervený J, Falteisek L, Gaisler J, Hanzal V, Horácek D, Hubálek Z, Jahelková H, Kolarík M, Korytár L, Kubátová A, Lehotská B, Lehotský R, Lucan RK, Májek O, Mateju J, Rehák Z, Safár J, Tájek P, Tkadlec E, Uhrin M, Wagner J, Weinfurtová D (et al), 2010. Increasing incidence of Geomyces destructans fungus in bats from the Czech Republic and Slovakia. PLoS ONE, No.November:e13853. http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0013853
Mascuch SJ, Moree WJ, Hsu CC, Turner GG, Cheng TL, Blehert DS, Kilpatrick AM, Frick WF, Meehan MJ, Dorrestein PC, Gerwick L, 2015. Direct detection of fungal siderophores on bats with white-nose syndrome via fluorescence microscopy-guided ambient ionization mass spectrometry. PLoS ONE, 10(3):e0119668. http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0119668
Meteyer CU, Barber D, Mandl JN, 2012. Pathology in euthermic bats with white nose syndrome suggests a natural manifestation of immune reconstitution inflammatory syndrome. Virulence, 3(7):583-588. http://www.landesbioscience.com/journals/virulence/article/22330/?show_full_text=true
Meteyer CU, Buckles EL, Blehert DS, Hicks AC, Green DE, Shearn-Bochsler V, Thomas NJ, Gargas A, Behr MJ, 2009. Histopathologic criteria to confirm white-nose syndrome in bats. Journal of Veterinary Diagnostic Investigation, 21(4):411-414. http://jvdi.org/archive/
Meteyer CU, Valent M, Kashmer J, Buckles EL, Lorch JM, Blehert DS, Lollar A, Berndt D, Wheeler E, White CL, Ballmann AE, 2011. Recovery of little brown bats (Myotis lucifugus) from natural infection with Geomyces destructans, white-nose syndrome. Journal of Wildlife Diseases, 47(3):618-626. http://www.jwildlifedis.org/cgi/content/full/47/3/618
Meyer AD, Stevens DF, Blackwood JC, 2016. Predicting bat colony survival under controls targeting multiple transmission routes of white-nose syndrome. Journal of Theoretical Biology, 409:60-69. http://www.sciencedirect.com/science/article/pii/S0022519316302752
Minnis AM, Lindner DL, 2013. Phylogenetic evaluation of Geomyces and allies reveals no close relatives of Pseudogymnoascus destructans, comb. nov., in bat hibernacula of eastern North America. Fungal Biology, 117(9):638-649. http://www.sciencedirect.com/science/article/pii/S1878614613001025
Mulec J, Covington E, Walochnik J, 2013. Is bat guano a reservoir of Geomyces destructans? Open Journal of Veterinary Medicine, 3(2):161-167. http://www.scirp.org/journal/PaperInformation.aspx?PaperID=32699
Muller LK, Lorch JM, Lindner DL, O'Connor M, Gargas A, Blehert DS, 2013. Bat white-nose syndrome: a real-time TaqMan polymerase chain reaction test targeting the intergenic spacer region of Geomyces destructans. Mycologia, 105(2):253-259
O'Donoghue AJ, Knudsen GM, Beekman C, Perry JA, Johnson AD, DeRisi JL, Craik CS, Bennet RJ, 2015. Destructin-1 is a collagen-degrading endopeptidase secreted by Pseudogymnoascus destructans, the causative agent of white-nose syndrome. Proceedings of the National Academy of Sciences of the USA, 112(24):7478-7483
O'Regan SM, Magori K, Pulliam JT, Zokan MA, Kaul RB, Barton HD, Drake JM, 2015. Multi-scale model of epidemic fade-out: will local extirpation events inhibit the spread of white-nose syndrome? Ecological Applications, 25(3):621-633. http://www.esajournals.org/doi/full/10.1890/14-0417.1
Paiva-Cardoso M das N, Morinha F, Barros P, Vale-Gonçalves H, Coelho AC, Fernandes L, Travassos P, Faria AS, Bastos E, Santos M, Cabral JA, 2014. First isolation of Pseudogymnoascus destructans in bats from Portugal. European Journal of Wildlife Research, 60(4):645-649. http://link.springer.com/article/10.1007%2Fs10344-014-0831-2
Pannkuk EL, Risch TS, Savary BJ, 2015. Isolation and Identification of an extracellular subtilisin-like serine protease secreted by the bat pathogen Pseudogymnoascus destructans. PLoS One, 10(3):e0120508
Pavlinic I, Ðakovic M, Lojkic I, 2015. Pseudogymnoascus destructans in Croatia confirmed. European Journal of Wildlife Research, 61(2):325-328. http://link.springer.com/article/10.1007%2Fs10344-014-0885-1
Pikula J, Bandouchova H, Novotný L, Meteyer CU, Zukal J, Irwin NR, Zima J, Martínková N, 2012. Histopathology confirms white-nose syndrome in bats in Europe. Journal of Wildlife Diseases, 48(1):207-211. http://www.jwildlifedis.org/content/48/1/207.full
Puechmaille SJ, Frick WF, Kunz TH, Racey PA, Voigt CC, Wibbelt G, Teeling EC, 2011. White-nose syndrome: is this emerging disease a threat to European bats? Trends in Ecology & Evolution, 26(11):570-576. http://www.sciencedirect.com/science/journal/01695347
Puechmaille SJ, Verdeyroux P, Fuller H, Gouilh MA, Bekaert M, Teeling EC, 2010. White-nose syndrome fungus (Geomyces destructans) in bat, France. Emerging Infectious Diseases, 16(2):290-293. http://www.cdc.gov/eid/content/16/2/290.htm
Puechmaille SJ, Wibbelt G, Korn V, Fuller H, Forget F, Mühldorfer K, Kurth A, Bogdanowicz W, Borel C, Bosch T, Cherezy T, Drebet M, Görföl T, Haarsma AJ, Herhaus F, Hallart G, Hammer M, Jungmann C, Bris Yle, Lutsar L, Masing M, Mulkens B, Passior K, Starrach M, Wojtaszewski A, Zöphel U (et al), 2011. Pan-European distribution of white-nose syndrome fungus (Geomyces destructans) not associated with mass mortality. PLoS ONE, No.April:e19167. http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0019167
Rajkumar SS, Li XJ, Rudd RJ, Okoniewski JC, Xu JP, Chaturvedi S, Chaturvedi V, 2011. Clonal genotype of Geomyces destructans among bats with white nose syndrome, New York, USA. Emerging Infectious Diseases, 17(7):1273-1276. http://www.cdc.gov/eid/content/17/7/1273.htm
Rapin N, Johns K, Martin L, Warnecke L, Turner JM, Bollinger TK, Willis CKR, Voyles J, Misra V, 2014. Activation of innate immune-response genes in little brown bats (Myotis lucifugus) infected with the fungus Pseudogymnoascus destructans. PLoS ONE, 9(11):e112285. http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0112285
Raudabaugh DB, Miller AN, 2013. Nutritional capability of and substrate suitability for Pseudogymnoascus destructans, the causal agent of bat white-nose syndrome. PLoS ONE, 8(10):e78300. http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0078300
Reeder DM, Frank CL, Turner GG, Meteyer CU, Kurta A, Britzke ER, Vodzak ME, Darling SR, Stihler CW, Hicks AC, Jacob R, Grieneisen LE, Brownlee SA, Muller LK, Blehert DS, 2012. Frequent arousal from hibernation linked to severity of infection and mortality in bats with white-nose syndrome. PLoS ONE, 7(6):e38920. http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0038920
Reichard JD, Kunz TH, 2009. White-nose syndrome inflicts lasting injuries to the wings of little brown myotis (Myotis lucifugus). Acta Chiropterologica, 11(2):457-464. http://www.bioone.org/doi/abs/10.3161/150811009X485684
Reiskind MH, Wund MA, 2009. Experimental assessment of the impacts of northern long-eared bats on ovipositing Culex (Diptera: Culicidae) mosquitoes. Journal of Medical Entomology, 46(5):1037-1044. http://docserver.ingentaconnect.com/deliver/connect/esa/00222585/v46n5/s10.pdf?expires=1257376399&id=0000&titleid=10266&checksum=30387CAF864617F53701316548939DF1
Ren P, Haman KH, Last LA, Rajkumar SS, Keel MK, Chaturvedi V, 2012. Clonal spread of Geomyces destructans among bats, Midwestern and Southern United States. Emerging Infectious Diseases, 18(5):883-885. http://wwwnc.cdc.gov/eid/article/18/5/pdfs/11-1711.pdf
Reynolds HT, Barton HA, 2014. Comparison of the white-nose syndrome agent Pseudogymnoascus destructans to cave-dwelling relatives suggests reduced saprotrophic enzyme activity. PLoS ONE, 9(1):e86437. http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0086437
Reynolds HT, Ingersoll T, Barton HA, 2015. Modeling the environmental growth of Pseudogymnoascus destructans and its impact on the white-nose syndrome epidemic. Journal of Wildlife Diseases, 51(2):318-331. http://www.jwildlifedis.org/doi/full/10.7589/2014-06-157
Reynolds RJ, Powers KE, Orndorff W, Ford WM, Hobson CS, 2016. Changes in rates of capture and demographics of Myotis septentrionalis (Northern Long-eared Bat) in western Virginia before and after onset of white-nose syndrome. Northeastern Naturalist, 23(2):195-204. http://www.bioone.org/loi/nena
Russell RE, Thogmartin WE, Erickson RA, Szymanski J, Tinsley K, 2015. Estimating the short-term recovery potential of little brown bats in the eastern United States in the face of White-nose syndrome. Ecological Modelling, 314:111-117. http://www.sciencedirect.com/science/article/pii/S0304380015003221
Sachanowicz K, Stepien A, Ciechanowski M, 2014. Prevalence and phenology of white-nose syndrome fungus Pseudogymnoascus destructans in bats from Poland. Central European Journal of Biology, 9(4):437-443. http://rd.springer.com/article/10.2478/s11535-013-0280-z
Shelley V, Kaiser S, Shelley E, Williams T, Kramer M, Haman K, Keel K, Barton HA, 2013. Evaluation of strategies for the decontamination of equipment for Geomyces destructans, the causative agent of white-nose syndrome (WNS). Journal of Cave and Karst Studies, 75(1):1-10
Simonovicov A, Pangallo D, Chovanová K, Lehotská B, 2011. Geomyces destructans associated with bat disease WNS detected in Slovakia. Biologia (Bratislava), 66(3):562-564. http://www.springerlink.com/content/16h34421556mg307/
Sleeman JM, 2011. Universal precautions for the management of bat white-nose syndrome (WNS):2 pp. [Wildlife Health Bulletin 2011-05.] https://www.nwhc.usgs.gov/publications/wildlife_health_bulletins/WHB_2011-05_UniversalPrecautions.pdf
Smyth C, Schlesinger S, Overton BE, Butchkoski C, 2013. Alternative host hypothesis and potential virulence genes in Geomyces destructans. Bat Research News, 54(2):17-24
Stading BR, Osorio JE, Velasco-Villa A, Smotherman M, Kingstad-Bakke B, Rocke TE, 2016. Infectivity of attenuated poxvirus vaccine vectors and immunogenicity of a raccoonpox vectored rabies vaccine in the Brazilian Free-tailed bat (Tadarida brasiliensis). Vaccine, 34(44):5352-5358. http://www.sciencedirect.com/science/journal/0264410X
Thapa V, Turner GG, Hafenstein S, Overton BE, Vanderwolf KJ, Roossinck MJ, 2016. Using a novel partitivirus in Pseudogymnoascus destructans to understand the epidemiology of White-nose syndrome. PLoS Pathogens, 12(12):e1006076. http://journals.plos.org/plospathogens/article?id=10.1371/journal.ppat.1006076
Thogmartin WE, Sanders-Reed CA, Szymanski JA, McKann PC, Pruitt L, King RA, Runge MC, Russell RE, 2013. White-nose syndrome is likely to extirpate the endangered Indiana bat over large parts of its range. Biological Conservation, 160:162-172
Turner GG, Meteyer CU, Barton H, Gumbs JF, Reeder DM, Overton B, Bandouchova H, Bartonicka T, Martínková N, Pikula J, Zukal J, Blehert DS, 2014. Nonlethal screening of bat-wing skin with the use of ultraviolet fluorescence to detect lesions indicative of white-nose syndrome. Journal of Wildlife Diseases, 50(3):566-573. http://www.jwildlifedis.org/doi/full/10.7589/2014-03-058
Turner GG, Reeder DM, Coleman JTH, 2011. Estimating the short-term recovery potential of little brown bats in the eastern United States in the face of white-nose syndrome. Bat Research News, 52(2):13-27
US Fish and Wildlife Service, 2011. A national plan for assisting states, federal agencies and tribes in managing whitenose syndrome in bats. Hadley, Massachusetts, USA: U.S. Fish and Wildlife Service, 17 pp. https://www.whitenosesyndrome.org/sites/default/files/white-nose_syndrome_national_plan_may_2011_0.pdf
US Fish and Wildlife Service, 2016. White-nose syndrome.org: A coordinated response to the devastating bat disease. Where is it now? Hadley, Massachusetts, USA: U.S. Fish and Wildlife Service. https://www.whitenosesyndrome.org/about/where-is-it-now
USGS National Wildlife Health Center, 2015. Diagnostic categories for reporting cases of bat white-nose syndrome (WNS) including a summary of revisions to WNS case definitions for the 2014/2015 season. Madison, Wisconsin, USA: USGS National Wildlife Health Center, 3 pp. https://www.nwhc.usgs.gov/disease_information/white-nose_syndrome/Case%20Defintions%20for%20WNS.pdf
Verant ML, Boyles JG, Waldrep W, Wibbelt G, Blehert DS, 2012. Temperature-dependent growth of Geomyces destructans, the fungus that causes bat white-nose syndrome. PLoS One, 7(9):e46280
Verant ML, Meteyer CU, Speakman JR, Cryan PM, Lorch JM, Blehert DS, 2014. White-nose syndrome initiates a cascade of physiologic disturbances in the hibernating bat host. BMC Physiology, 14(10):(9 December 2014). http://www.biomedcentral.com/content/pdf/s12899-014-0010-4.pdf
Voyles J, Kilpatrick AM, Collins JP, Fisher MC, Frick WF, McCallum H, Willis CKR, Blehert DS, Murray KA, Puschendorf R, Rosenblum EB, Bolker BM, Cheng TL, Langwig KE, Lindner DL, Toothman M, Wilber MQ, Briggs CJ, 2015. Moving beyond too little, too late: managing emerging infectious diseases in wild populations requires international policy and partnerships. EcoHealth, 12(3):404-407. http://link.springer.com/article/10.1007%2Fs10393-014-0980-5
Warnecke L, Turner JM, Bollinger TK, Lorch JM, Vikram Misra, Cryan PM, Wibbelt G, Blehert DS, Willis CKR, 2012. Inoculation of bats with European Geomyces destructans supports the novel pathogen hypothesis for the origin of white-nose syndrome. Proceedings of the National Academy of Sciences of the United States of America, 109(18):6999-7003. http://www.pnas.org/content/109/18/6999.full
Wibbelt G, Kurth A, Hellmann D, Weishaar M, Barlow A, Veith M, Prüger J, Görföl T, Grosche L, Bontadina F, Zöphel U, Seidl HP, Cryan PM, Blehert DS, 2010. White-nose syndrome fungus (Geomyces destructans) in bats, Europe. Emerging Infectious Diseases, 16(8):1237-1242. http://www.cdc.gov/eid/content/16/8/1237.htm
Wibbelt G, Puechmaille SJ, Ohlendorf B, Mühldorfer K, Bosch T, Görföl T, Passior K, Kurth A, Lacremans D, Forget F, 2013. Skin lesions in European hibernating bats associated with Geomyces destructans, the etiologic agent of white-nose syndrome. PLoS ONE, 8(9):e74105. http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0074105
Wilder AP, Kunz TH, Sorenson MD, 2015. Population genetic structure of a common host predicts the spread of white-nose syndrome, an emerging infectious disease in bats. Molecular Ecology, 24(22):5495-5506. http://onlinelibrary.wiley.com/doi/10.1111/mec.13396/abstract
Willis CKR, Menzies AK, Boyles JG, Wojciechowski MS, 2011. Evaporative water loss is a plausible explanation for mortality of bats from white-nose syndrome. Integrative and Comparative Biology [Environment, Energetics, and Fitness: A Symposium Honoring Donald W. Thomas. Annual meeting of the Society for Integrative and Comparative Biology, Salt Lake City, Utah, USA, 3-7 January 2011.], 51(3):364-373. http://icb.oxfordjournals.org/
Zhang T, Chaturvedi V, Chaturvedi S, 2015. Novel Trichoderma polysporum strain for the biocontrol of Pseudogymnoascus destructans, the fungal etiologic agent of bat white nose syndrome. PLoS One, 10(10):e0141316
Zukal J, Bandouchova H, Bartonicka T, Berkova H, Brack V, Brichta J, Dolinay M, Jaron KS, Kovacova V, Kovarik M, Martínková N, Ondracek K, Rehak Z, Turner GG, Pikula J, 2014. White-nose syndrome fungus: a generalist pathogen of hibernating bats. PLoS ONE, 9(5):e97224. http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0097224
Zukal J, Bandouchova H, Brichta J, Cmokova A, Jaron KS, Kolarik M, Kovacova V, Kubátová A, Nováková A, Orlov O, Pikula J, Presetnik P, Suba J, Zahradníková A Jr, Martínková N, 2016. White-nose syndrome without borders: Pseudogymnoascus destructans infection tolerated in Europe and Palearctic Asia but not in North America. Scientific Reports, 6(19829):srep19829. http://www.nature.com/articles/srep19829
Barlow A M, Worledge L, Miller H, Drees K P, Wright P, Foster J T, Sobek C, Borman A M, Fraser M, 2015. First confirmation of Pseudogymnoascus destructans in British bats and hibernacula. Veterinary Record. 177 (3), 73. http://veterinaryrecord.bvapublications.com/archive/ DOI:10.1136/vr.102923
Blehert DS, Hicks AC, Behr M, Meteyer CU, Berlowski-Zier BM, Buckles EL, Coleman JTH, Darling SR, Gargas A, Niver R, Okoniewski JC, Rudd RJ, Stone WB, 2009. Bat white-nose syndrome: An emerging fungal pathogen? In: Science, 323 (5911) 227.
CABI, Undated. CABI Compendium: Status inferred from regional distribution. Wallingford, UK: CABI
Hoyt J R, Sun KePing, Parise K L, Lu GuanJun, Langwig K E, Jiang TingLei, Yang ShuBao, Frick W F, Kilpatrick A M, Foster J T, Feng Jiang, 2016. Widespread bat white-nose syndrome fungus, northeastern China. Emerging Infectious Diseases, 22 (1), 140-142. http://wwwnc.cdc.gov/eid/article/22/1/pdfs/15-1314.pdf DOI:10.3201/eid2201.151314
Leopardi S, Blake D, Puechmaille SJ, 2015. White-nose syndrome fungus introduced from Europe to North America. In: Current Biology, 25 (6) R217-R219.
Lorch J M, Palmer J M, Lindner D L, Ballmann A E, George K G, Griffin K, Knowles S, Huckabee J R, Haman K H, Anderson C D, Becker P A, Buchanan J B, Foster J T, Blehert D S, 2016. First detection of bat white-nose syndrome in western North America. mSphere. 1 (4), 00148-16. http://msphere.asm.org/content/1/4/e00148-16
Martínková N, Bačkor P, Bartonička T, Blažková P, Červený J, Falteisek L, Gaisler J, Hanzal V, Horáček D, Hubálek Z, Jahelková H, Kolařík M, Korytár L, Kubátová A, Lehotská B, Lehotský R, Lučan R K, Májek O, Matějů J, Řehák Z, Šafář J, Tájek P, Tkadlec E, Uhrin M, Wagner J, Weinfurtová D (et al), 2010. Increasing incidence of Geomyces destructans fungus in bats from the Czech Republic and Slovakia. PLoS ONE. e13853. http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0013853 DOI:10.1371/journal.pone.0013853
Paiva-Cardoso M das N, Morinha F, Barros P, Vale-Gonçalves H, Coelho A C, Fernandes L, Travassos P, Faria A S, Bastos E, Santos M, Cabral J A, 2014. First isolation of Pseudogymnoascus destructans in bats from Portugal. European Journal of Wildlife Research. 60 (4), 645-649. http://link.springer.com/article/10.1007%2Fs10344-014-0831-2 DOI:10.1007/s10344-014-0831-2
Pavlinić I, Ðaković M, Lojkić I, 2015. Pseudogymnoascus destructans in Croatia confirmed. European Journal of Wildlife Research. 61 (2), 325-328. http://link.springer.com/article/10.1007%2Fs10344-014-0885-1 DOI:10.1007/s10344-014-0885-1
Pikula J, Bandouchova H, Novotný L, Meteyer C U, Zukal J, Irwin N R, Zima J, Martínková N, 2012. Histopathology confirms white-nose syndrome in bats in Europe. Journal of Wildlife Diseases. 48 (1), 207-211. http://www.jwildlifedis.org/content/48/1/207.full
Puechmaille S J, Verdeyroux P, Fuller H, Gouilh M A, Bekaert M, Teeling E C, 2010. White-nose syndrome fungus (Geomyces destructans) in bat, France. Emerging Infectious Diseases. 16 (2), 290-293. http://www.cdc.gov/eid/content/16/2/290.htm
Puechmaille S J, Wibbelt G, Korn V, Fuller H, Forget F, Mühldorfer K, Kurth A, Bogdanowicz W, Borel C, Bosch T, Cherezy T, Drebet M, Görföl T, Haarsma A J, Herhaus F, Hallart G, Hammer M, Jungmann C, Bris Y le, Lutsar L, Masing M, Mulkens B, Passior K, Starrach M, Wojtaszewski A, Zöphel U (et al), 2011. Pan-European distribution of white-nose syndrome fungus (Geomyces destructans) not associated with mass mortality. PLoS ONE. e19167. http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0019167 DOI:10.1371/journal.pone.0019167
Sachanowicz K, Stępień A, Ciechanowski M, 2014. Prevalence and phenology of white-nose syndrome fungus Pseudogymnoascus destructans in bats from Poland. Central European Journal of Biology. 9 (4), 437-443. http://rd.springer.com/article/10.2478/s11535-013-0280-z DOI:10.2478/s11535-013-0280-z
Šimonovičov A, Pangallo D, Chovanová K, Lehotská B, 2011. Geomyces destructans associated with bat disease WNS detected in Slovakia. Biologia (Bratislava). 66 (3), 562-564. http://www.springerlink.com/content/16h34421556mg307/ DOI:10.2478/s11756-011-0041-2
US Fish and Wildlife Service, 2016. White-nose syndrome.org: A coordinated response to the devastating bat disease. Where is it now?, Hadley, Massachusetts, USA: U.S. Fish and Wildlife Service. https://www.whitenosesyndrome.org/about/where-is-it-now
Wibbelt G, Kurth A, Hellmann D, Weishaar M, Barlow A, Veith M, Prüger J, Görföl T, Grosche L, Bontadina F, Zöphel U, Seidl H P, Cryan P M, Blehert D S, 2010. White-nose syndrome fungus (Geomyces destructans) in bats, Europe. Emerging Infectious Diseases. 16 (8), 1237-1242. http://www.cdc.gov/eid/content/16/8/1237.htm
Wibbelt G, Puechmaille S J, Ohlendorf B, Mühldorfer K, Bosch T, Görföl T, Passior K, Kurth A, Lacremans D, Forget F, 2013. Skin lesions in European hibernating bats associated with Geomyces destructans, the etiologic agent of white-nose syndrome. PLoS ONE. 8 (9), e74105. http://www.plosone.org/article/info%3Adoi%2F10.1371%2Fjournal.pone.0074105 DOI:10.1371/journal.pone.0074105
Zukal J, Bandouchova H, Brichta J, Cmokova A, Jaron K S, Kolarik M, Kovacova V, Kubátová A, Nováková A, Orlov O, Pikula J, Presetnik P, Šuba J, Zahradníková A Jr, Martínková N, 2016. White-nose syndrome without borders: Pseudogymnoascus destructans infection tolerated in Europe and Palearctic Asia but not in North America. Scientific Reports. 6 (1), 19829. http://www.nature.com/articles/srep19829
OrganizationsTop of page
Canada: Canadian Wildlife Health Cooperative (CWHC), Western College of Veterinary Medicine, University of Saskatchewan, Saskatoon, SK, S7N 5B4, http://www.cwhc-rcsf.ca/index.php
USA: Bureau of Land Management (BLM), 20 M Street SE, Washington, DC 20003, https://www.blm.gov/
USA: National Park Service (NPS), 1849 C Street NW, Washington, DC 20240, https://www.nps.gov/
USA: US Fish and Wildlife Service (USFWS), 300 Westgate Center Drive, Hadley, MA 01035, https://www.whitenosesyndrome.org/
USA: US Forest Service (USFS), 1400 Independence Ave. SW, Washington, DC 20250, https://www.fs.fed.us/
USA: US Geological Survey (USGS), USGS National Center, 12201 Sunrise Valley Drive, Reston, VA 20192, https://www.usgs.gov/
ContributorsTop of page
13/02/17: Original text by:
David Blehert, USGS National Wildlife Health Center, Madison, Wisconsin, USA
Emily Lankau, USGS National Wildlife Health Center, Madison, Wisconsin, USA
Use of trade, product, or firm names is for descriptive purposes only and does not imply endorsement by the US government.
Distribution MapsTop of page
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