White-nose disease (WND) is a deadly disease in hibernating bats. Its causal agent is the fungus Pseudogymnoascus destructans (Pd) that was only recently introduced to North America, where it causes pervasive mortality. In contrast, it is native to Eurasia, where bats do not experience mass mortality. The pathogen successfully survives for long periods in cave substrates (e.g., mud, soil) as well as on contaminated caving clothes and equipment. This emphasizes the importance of limiting its human-mediated spread, particularly over long distances and from areas where it is native and genetically highly diverse. To assess associated conservation risks more precisely, we conducted a global online survey on cavers’ WND knowledge, travel frequencies, and hygiene habits regarding caving equipment, and we analysed the relationship between those variables. We interpreted our results based on respondents’ continents of origin, separating them into 3 categories: continents where Pd was recently introduced and continues to cause significant bat mortality (North America); continents where Pd is currently not present but is hypothesized to cause bat mortality if introduced (Australia and South and Central America); and continents where Pd is native and currently not associated with bat mortality (Europe and Asia). We found significant differences between categories, showing that further awareness of WND is most needed in Europe and Asia where specific guidelines for limiting Pd spread are still to be developed. Interestingly, however, WND knowledge alone explained very little variance in respondents’ hygiene habits, showing that other factors such as habits, emotions, or personal values have a stronger influence on cavers’ behaviour.

Bat conservation, pathogen pollution, Pseudogymnoascus destructans, white-nose syndrome, wildlife disease

White-nose disease (WND), or white-nose syndrome as it is often referred to (WNS, please see (Whiting-Fawcett et al. 2025 for details on nomenclature), is a deadly disease in hibernating bats that has caused population declines of more than 90% in the colonies of most affected bat species in North America (Cheng et al. 2021). Its causal agent is the cave-dwelling and cold-loving fungus Pseudogymnoascus destructans (Pd), native to Eurasia and only recently introduced to North America, most likely by humans (Leopardi et al. 2015; Drees et al. 2018), providing a notorious example of pathogen pollution. This recent introduction explains the pervasive mortality due to WND in North America and the lack of mass morbidity or mortality in Europe (Puechmaille et al. 2010, 2011; Zukal et al. 2016; Fritze and Puechmaille 2018). Importantly, Pd only infects bats during hibernation, when their body temperature is low enough to allow the growth of a psychrophilic pathogen, but it is able to survive for multiple years in the substrate, making caves its environmental reservoir (Puechmaille et al. 2011; Fischer et al. 2020, 2022; Hoyt et al. 2020). This has two major consequences. First, healthy bats can get infected anew each autumn when they return to their hibernacula (Puechmaille et al. 2011; Hoyt et al. 2021; Fischer et al. 2022; Hicks et al. 2023; Zhelyazkova et al. 2024). Second, the unintentional transportation of cave substrate (e.g., via dirty clothes and equipment) can spread Pd spores, and therefore introduce the pathogen to new sites, countries, or continents. It can also move the pathogen between existing populations, creating favourable conditions for genetic mixing of previously isolated entities, which might lead to the emergence of potentially more dangerous pathogen variants, as classically encountered in fungi (Brasier and Buck 2001; Stukenbrock 2016).

In North America, where subterranean habitats are still in the process of Pd colonisation, bats have proved to be important vectors of its spread over short distances (e.g., Maher et al. 2012) as the fungus can only survive for several days on normothermic animals (Campbell et al. 2020). Additionally, tracking individual Pd genotypes in Europe has demonstrated very limited movement and/or establishment of the pathogen to already colonized sites within its native range of occurrence (Fischer et al. 2022). Together, these findings show that, although inevitable, “natural” bat-mediated Pd spread is relatively slow and subject to significant long-term limitations. In contrast, thanks to modern means of transportation and the ever-rising popularity of international travel, people effectively act as fast long-distance Pd vectors, even across continents. Thus, both the transcontinental introduction of the pathogen from Europe to North America (Leopardi et al. 2015) and its introduction from the East to the West Coast of the US (Lorch et al. 2006; Thapa et al. 2021) are considered human-mediated. The same is most probably true for Pd spread in other large countries such as Russia or China. Importantly, it was demonstrated that Pd spores not only easily stick to caving equipment after visiting a Pd-positive subterranean sites but can survive on such contaminated equipment at room temperature for at least 25 days (Zhelyazkova et al. 2020). This means that risks of human-mediated pathogen introductions remain even when pauses between trips are made.

Considering all of the above, raising WND awareness is thought to be a successful strategy to reduce the long-distance spread of Pd and possibly other cave microbes. Yet, very little has been published regarding the effectiveness of such information campaigns and the different factors that might influence cavers’ behaviour. For example, Salleh et al. (2021) analysed WND knowledge and cleaning habits among the participants in a speleological conference in Australia and Shapiro et al. (2022) surveyed people’s attitudes and behavioural intent towards WND prevention measures applied by US National Parks. Thus, the results of these studies are biased towards people who are already interested in cave conservation and/or come from continents where WND is causing or expected to cause bat mortality (North America and Australia) (Blomberg et al. 2023). However, new Pd (re-)introductions are not less probable from places where the pathogen is currently not causing bat mortality (e.g., Europe and Asia), thus effective WND prevention can only be achieved by a global approach. As a step in this direction, we conducted a global online survey on cavers’ WND knowledge, travel frequencies, and hygiene habits, and we analysed the relationship between those variables. We used a broad definition of “cavers” referring to any people practicing any activities in natural caves of various origins, as well as mines and other artificial underground sites. We interpreted most of our results based on respondents’ continents of origin separating them into 3 categories: continents where Pd was recently introduced and continues to cause significant bat mortality (North America); continents where Pd is currently not present but is hypothesized to cause bat mortality if introduced (Australia and South and Central America, see Escobar et al. 2014; Turbill and Welbergen 2020; Blomberg et al. 2023); and continents where Pd is native and not associated with bat mortality (Europe and Asia). We hypothesized that WND knowledge would be higher at places seriously affected by the disease and that this knowledge would exert a strong influence on cavers’ hygiene habits.

In agreement with the first part of our hypothesis, our study showed a large difference in WND awareness, adherence to WND-related instructions and hygiene habits (expressed here as biosecurity scores) between cavers on different continents. In contrast, we did not find a strong support for WND knowledge influencing cavers’ hygiene habits by itself, which refutes the second part of our hypothesis. Such results align with the Theory of Planned Behaviour (TPB, Ajzen 1985), which posits that human behavioural intentions are determined by three components. These are: 1) attitude (Do I approve/like this behaviour?); 2) subjective norms (Is this behaviour socially desirable?); and 3) perceived behavioural controls (Do I find this behaviour feasible to me?). Within this conceptual framework, knowledge can indirectly influence behaviour by shaping one or more of these components, yet, habits, emotions or personal values often play a much bigger role. Thus, cavers in North America may be stricter at cleaning and decontaminating their equipment not only because they are more aware of WND but also for other more personal reasons. For example, the mere sight of hundreds of dead bats during the WND epidemics must have raised empathy and motivation for action among the caving community and the broader society (Component 1 of TPB). Additionally, once WND-related instructions were widely distributed and a certain mass of people started complying to them, it must have become socially desirable to adopt such behaviour (Component 2 of TPB). On the other hand, cavers in South America may wash their caving clothes with soap not necessarily with the intention of pathogen pollution prevention but in agreement with their everyday habits. As people from warmer countries normally go caving with everyday clothes such as pants and a T-shirt, they might find it easy to put these in the washing machine in contrast to cavers from colder countries who use heavy caving suits, often sensitive to detergents. Such behaviour aligns with Component 3 of TPB. Another example of habits’ importance is the consistency in cavers’ hygiene practices across travel distances. On the other hand, applying stricter cleaning methods when travelling further may be motivated by awareness of the higher conservation risks following long-distance Pd spread.

The same theory also explains our seemingly striking observation that cavers practicing science-related activities have better WND knowledge but not better hygiene habits than cavers who are not involved in research. This is so as professional caving for scientific purposes often involves project requirements, deadlines and budget restrictions that may require visiting multiple caves per trip, sometimes in different geographical regions. The last is especially true for monitoring activities. In contrast, recreational caving is usually practiced within the comfort zone of participants who normally spend a caving weekend within the same geographical region. Thus, cleaning equipment between sites may be generally less feasible during scientific activities the way they are standardly organized.

Yet, our results do not mean that increasing cavers’ WND knowledge is useless in WND prevention. On the contrary, although it is not enough to motivate action by itself, knowledge provides the necessary instrumentation to perform a particular behaviour, once it becomes desired. This is exemplified by our results where the main reason for cavers following WND-related instructions is personal responsibility but the main reason for people not following such is the lack of awareness. In this regard, the results of our study provide useful clues on where the gaps in WND awareness are and what might be the reasons for them. Thus, cavers from North America, where Pd has been decimating bat populations in the past two decades, are particularly knowledgeable about WND and the specific instructions designed for its mitigation. This is expected given the significant ecological and economic consequences of reduced bat populations due to the disease that motivated the rapid testing and dissemination of Pd decontamination protocols (https://www.whitenosesyndrome.org/mmedia-education/national-wns-decontamination-protocol-u-s). Additionally, a variety of information resources such as articles, infographs, short videos, or whole documentaries, was developed and made accessible to the general public (see the Resources tab at https://www.whitenosesyndrome.org/). As evidenced by the high percentage of correct answers to WND-related questions among our North American respondents, these resources seem to have been effective at increasing awareness about the disease. As for continents such as South and Central America and Australia where Pd is not yet reported but is expected to cause serious bat morbidity or mortality if introduced, cavers are also well aware of WND, with approximately half of them preventively adhering to some WND-related instructions. In Australia, this is most probably explained by the fact that specific WND-response guidelines have already been developed and disseminated there https://wildlifehealthaustralia.com.au). Additionally, the country has a strict biosecurity policy in general and offers useful information resources to different target groups in order to prevent the spread of invasive pests and diseases (https://www.biosecurity.gov.au/). In contrast, we are not aware of formal biosecurity protocols in South American countries but our data is insufficient to determine whether responses from this continent reflect local conservation campaigns, broader access to information from the US, or simply local climatic conditions. Such local contexts should be taken into account when designing field hygiene protocols.

In Europe and Asia, where Pd is currently not associated with bat mortality, cavers are least aware of WND and usually clean their equipment solely with water, which can remove some but not all fungal spores (Zhelyazkova et al. 2020). This is expected both due to the lack of an immediately apparent problem and to the fact that Pd was discovered and proven native to Europe (Puechmaille et al. 2010) and Asia (Hoyt et al. 2016) after the disease had manifested in the US. Indeed, some WND-related instructions have been developed in Europe such as the BCI guidance for bat workers in UK (https://cdn.bats.org.uk/uploads/pdf/About%20Bats/WNS/UK-Guidance-for-bat-workers-Nov-2024.pdf?v=1734025851). However, these do not seem to be so popular in other countries. IUCN has also published detailed guidelines for field hygiene (https://www.iucnbsg.org/guidelines-for-field-hygiene.html), which are particularly valuable for researchers handling bats but not so applicable to recreational cavers. Lastly, the European Speleological Federation has created a WND factsheet containing recommendations for cavers (https://www.eurospeleo.eu/ECPC/wp-content/uploads/2022/09/ECPC_WNS_in_Europe_cavers_ENFR.pdf?), however it dates back to 2010 and does not reflect current advancements of the disease knowledge. Thus, we suggest to continue increasing WND awareness among the Eurasian caving community and to better explain the difference between short- and long-distance Pd spread on this continent. More precisely, according to the latest scientific evidence, moving Pd between sites in the same geographical region in Europe or Asia (e.g., at distances below 100 km) is indeed associated with minimal conservation risks. This is so as the fungus is already widely distributed (Puechmaille et al. 2011; Zukal et al. 2016; Bloomberg et al. 2023); bats have had sufficient evolutionary time to adapt to it, mostly by developing immunological tolerance (Fritze et al. 2019, 2021; Whiting-Fawcett et al. 2025); and there is probably significant intraspecific competition preventing the establishment of new Pd genotypes into new sites already colonized by the same species (Fischer et al. 2022; Zhelyazkova et al. 2024). On the other hand, moving Pd between different geographical regions can have negative consequences by facilitating genetic recombination between different strains, potentially leading to the emergence of hypervirulent pathogen variants. This has happened in Batrachochytrium dendrobatidis, the causative agent of the deadly chytridiomycosis in amphibians (Farrer et al. 2011). Particularly high attention should be paid to intercontinental travel as it risks introducing Pd into previously unaffected areas or increasing genetic diversity in already infected regions, allowing strains previously isolated geographically to interact. While not every trip with Pd-contaminated gear will result in the successful establishment of Pd, each instance raises the likelihood of its introduction by increasing propagule pressure (Lockwood et al. 2005). Hence it is better to be “safe than sorry” as it has been proven extremely challenging to find efficient management strategies to counter WND: from nearly 200 papers addressing the disease till 2023, only 10.5% included field-tested management practices and within those, only two reported solely positive impacts (Whiting-Fawcett et al. 2025). Thus, following the travel distance categories used in the present study, we advise washing caving equipment with water when changing subterranean sites situated up to 100 km apart and with water and soap for sites up to 1000 km apart. For longer distances, especially to different continents, we recommend using local equipment or following existing decontamination protocols such as those applied in North America or Australia. According to our study, the last should be particularly emphasized as a significant number of our respondents from Europe and Asia currently travel between continents without disinfecting or changing their equipment.

Considering the above, an effective campaign to prevent the anthropogenic spread of Pd should both provide knowledge and stimulate positive attitudes towards bat and cave conservation and make cleaning equipment more socially desirable and more easily feasible. In terms of attitudes, positive sentiments seem to be already prevailing: according to our study, most cavers that have received information on the human impact on cave ecosystems have changed their behaviour accordingly; most cavers that had prior knowledge of WND agree that limiting its spread is needed and possible; and most cavers that follow WND-related decontamination protocols do so out of a sense of responsibility. This is not surprising as, according to the authors’ rich caving experience, speleological activities are emotionally charged and cavers may think of the subterranean environment as their homeland. In terms of making field hygiene more socially desirable, we would suggest that a personal example could be particularly effective, especially when it comes from researchers who are known and respected within the community. Lastly, to make properly cleaning caving equipment more feasible, work in two directions is required. First, cavers that are unaware of WND and the related risks of pathogen pollution in caves, should be informed about the problem and its precise solutions. This is particularly needed in Europe and Asia where an up-to-date standardised protocol for cleaning and decontaminating caving equipment according to the travel distance has yet to be established and widely distributed. Importantly, these resources should be translated into each country’s languages. Second, researchers, especially those involved in monitoring activities, should be incentivized to prioritize time for cleaning their equipment between sites and including more rest days between trips in different geographical regions. The success of this idea would mostly depend on the attitudes among the research community towards prioritizing stricter pathogen pollution prevention rather than obtaining faster results.

Last but not least, we advocate the need for further studies assessing caver’s knowledge and attitude towards updated WND and cave conservation guidelines or going deeper into questions that we could not address in this paper. For example, we did not collect data on participants’ sex, number of caving courses and workshops attended, or the exact type of science they practice in caves: all these are factors that might also influence hygiene habits. Additionally, our data might be slightly biased towards cavers that are involved in some kind of scientific activities as we distributed our survey partly through our personal contact networks containing a large number of colleagues; we also received a higher response rate from our team members’ countries than from countries where we did not have personal contacts. Finally, the specific questions we used are only one of many ways to assess WND awareness and it has been observed that question formulation can influence statistical relations between knowledge and behaviour (Ajzen et al. 2011). Nevertheless, we believe that despite these limitations, the main points of our study remain valid. We hope that our questionnaire has raised even more interest in pathogen pollution in caves among our respondents and that cavers who wished to receive updates on our project would become valuable ambassadors spreading and improving our recommendations.

Pathogen pollution is a global problem that should be addressed via broad international cooperation. Thus, although significant advancements have been made in understanding and locally managing WND in severely affected areas, more efforts are needed to prevent further long-distance introductions of its causative agent. Rising awareness among the caving community is an important step in this direction, yet it is not enough as other factors such as personal attitudes and the feasibility of the proposed conservation actions have stronger influence on human behaviour.

We sincerely thank all cavers who took the time to fill in our questionnaire and helped with its distribution. Special gratitude to Lilyana Zhelyazkova, Vesselka Zhelayzkova, Clement Roure, and Changi Wong, who translated our questions in Spanish, Russian, French, and Chinese respectively. We are grateful to Boyan Petrov, Ferdinand Salazar, and Rollin Verlinde who provided their beautiful photographs for our website. This work was supported by the Slovenian Research and Innovation Agency project V1-2140 and by the Slovenian Research and Innovation Agency research core funding program “Geography of Slovenia” (P6-0101).