Temporal activity patterns of tourists and pumas Puma concolor in public areas in the Patagonia National Park, Chile

Abstract

The presence of humans and large carnivores in shared landscapes presents unique challenges for wildlife conservation and management. This study explores the temporal activity patterns of tourists and pumas Puma concolor using camera traps to understand the potential for encounters between the two groups in Patagonia National Park, Chile. The park, characterized by a cold temperate climate with diverse wildlife and vegetation, spans over 304 527 hectares. Data collected from August 2020 to March 2022 reveal distinct activity patterns for pumas and tourists, with pumas being primarily nocturnal and tourists being more active during the day. Despite a low overall overlap coefficient (Dhat = 0.27), specific periods during twilight hours showed increased interaction potential. Seasonal variations revealed a significant increase in overlap during the autumn of 2021, with stable overlap patterns observed both before and after this period. Seasonal variations in activity patterns highlight the need for adaptive management strategies for human–puma interactions. These findings underscore the importance of enforcing park regulations, promoting responsible tourism practices, and implementing specific behavioral protocols to ensure the safety of both tourists and pumas in protected areas. This study provides valuable insights for enhancing conservation efforts in the face of increasing wildlife‐watching tourism worldwide.

Introduction

Cohabitation with humans sometimes promotes tolerance of wildlife to human activities (Bejder et al. 2009, Lowry et al. 2013, Frank 2016). Nevertheless, cohabitation and casual encounters might generate potential conflicts between wildlife and humans (Mekonen 2020, Oommen 2021, Somu and Palanisamy 2022). Encounters between wildlife and humans occur across all aspects of the urban-to-natural gradient in various regions worldwide (Shepherd et al. 2014, Gebo et al. 2022, Montero-Botey and Perea 2023). Interactions involving wild carnivores are particularly concerning because human life is at potential risk of injuries or predation (Bombieri et al. 2018). When humans enter the habitat of large carnivores for recreational purposes, during activities such as hiking or camping, casual encounters with them become common. Likewise, the rise of wildlife-watching tourism has made these encounters explicitly desirable for many (Marion et al. 2020). Human interest in documenting events for recreational or professional purposes has experienced a sharp increase in recent decades (Howlett et al. 2023, Von Essen et al. 2023). This form of tourism with carnivores, however, carries inherent risks that could threaten tourists during periods of food scarcity for predators (Newsome and Rodger 2008, Penteriani et al. 2017). For example, designated zones for public use within protected areas, such as trails or tourists' facilities, are also used by some carnivores (Lucas 2020). Situations like this lead to scenarios in which visitors and wildlife might potentially interact (Candelario 2015, Gunther et al. 2024). In recent years, predator tourism in Patagonia has increasingly centered on the apex predator, the puma, resulting in potential encounters between tourists and pumas (Cifuentes-Ibarra et al. 2023).

The puma Puma concolor is the second largest cat in South America and the most widely distributed carnivore in the Americas, inhabiting places from southern Alaska to the tip of South America (Logan and Sweanor 2024). Pumas are an adaptable species that inhabit various climates and environments, including urban and rural areas (Craighead 2019, Figel et al. 2021, Guerisoli and Schiaffini 2022). Pumas are considered an apex species that fulfills a fundamental role in the ecosystem as a top–down regulator (Zanón-Martínez et al. 2012). This species has been involved in conflictive cohabitation with humans, many of which are related to the predation of livestock and domestic animals, which have generated unfavorable perceptions among local ranchers and farmers in several countries, resulting in retaliation towards these animals (Conforti and de Azevedo 2003, Burgas et al. 2014, Ohrens et al. 2016, Guerisoli et al. 2020). On the other hand, pumas are considered an emblematic and charismatic species for conservation, generating interest in people visiting natural environments.

The presence of carnivores and their use of trails or public use areas raises concerns about possible interactions with recreational activities like hiking. In this study, we examined the activity patterns of pumas and humans across a network of hiking trails within a protected area. Our analysis focused on discerning the temporal overlap of their activities to understand the hours during which potential encounters might occur. Additionally, we calculated the activity patterns of both pumas and tourists across different seasons to determine potential variations between peak (spring–summer) and off-peak (autumn–winter) touristic periods. Determining these patterns could help to understand the potential of encounters between tourists and pumas, which is key to ensure the safety of both and the implementation of better management practices in conservation areas.

Methods

Study area

The study area is located in the Patagonia National Park (PNP), 46°55ʹS 72°12ʹW, Aysén district, Chile, with the research conducted within the Chacabuco Valley sector (Fig. 1). This national park was established in 2018 by combining the donation of the Chacabuco Valley ranch, adjoining lands (83 733.41 ha), the adjacent Jeinimeni and Tamango national reserves, and public land without specific purpose (BNC 2018). The Chacabuco Valley ranch was acquired by Tompkins Conservation in 2004 with the goal of creating this national park (Rewilding Chile 2022). The former ranch was one of the largest in Patagonia, with over 30 000 sheep and 3000 cattle, which by 2017 were gradually reduced to zero following restoration efforts of Patagonian ecosystems (Herrera et al. 2017). The 304 527-hectare national park is characterized by a cold temperate climate with dry summers, with an average annual precipitation of approximately 800 mm (Wittmer et al. 2010). The vegetation transitions from deciduous Nothofagus forests to a Patagonian steppe, reflecting a gradient of the decreasing precipitation from the western mountainous regions to the eastern flat landscapes (Villa-Martínez et al. 2012, Luebert and Pliscoff 2017). The park is home to a variety of native carnivores and wildlife mammals, including the puma Puma concolor, culpeo fox Lycalopex culpaeus, chilla fox Lycalopex griseus, Geoffroy's cat Leopardus geoffroyi, pampas cat Leopardus pajeros, lesser grison Galictis cuja, and skunk Conepatus chinga (Iriarte and Jaksic 2017). Pumas are the apex predator in this ecosystem, having a density ranging from 0.3 to 1.04 individuals per 100 km² (Elbroch and Wittmer 2012). While these works report data from the last decade or earlier, they represent the most recent literature about the area, emphasizing the need for further research.

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The Chacabuco Valley's infrastructure and trails began to be built in 2006 and nowadays includes an extensive network of trails totaling almost 260 km, five camping zones, six parking areas, and three viewing points. This valley is traversed by the international vehicle route X-83, connecting Chile and Argentina.

Sampling design

Data on the activity patterns of pumas and tourists within the park were collected using camera traps positioned near trails frequented by tourists, from August 2020 to March 2022, resulting in a total of 9118 trap days. Seventeen sampling stations were established and equipped with a camera trap positioned at approximately 70 cm above the ground. Stations were spaced approximately 500 m apart and no attractors were used. Camera traps operated continuously, capturing three photographs per second with a 30-second interval between activations. Sampling stations were monitored every three months to download images and replace batteries.

Data analysis

Renaming photographs, organizing data, and classifying images adhered to the guidelines outlined by the Small Wild Cat Conservation Foundation (). In this process, four trained technicians were responsible for identifying the species, and unidentifiable images were discarded from our analysis. R-project software ver. 4.1.2 () and RStudio were employed for the analysis of data. To determine the activity patterns of pumas and tourists, independent events were calculated by grouping all images collected during intervals of 60 seconds (Kays and Parsons 2014). Independent events grouping and the hourly activity analysis of pumas and tourists were performed with the ‘camtrapR' package (Niedballa et al. 2016). The detection rate (DR) was calculated by dividing the number of independent events by the number of days that the cameras were active per 100 camera trap days.

Prior to overlap analysis, clock hours were adjusted for solar time to account for sunlight availability, using the sunTime function from the ‘overlap' package (Ridout and Linkie 2009, Nouvellet et al. 2012). Non-parametric kernel density estimation was used to compare the activities of puma and tourists, reported through an activity overlap plot and its overlap coefficient (Dhat1). Temporal overlap of species was conducted to determine the periods during which encounters between puma and tourists could have occurred, ranging from 0 (absence of overlap) to 1 (complete overlap) (Ridout and Linkie 2009). Confidence intervals at a 95% confidence level were estimated based on 1000 bootstrap samples. This analysis was applied using the ‘overlap' package (Ridout and Linkie 2009). For data homogeneity, we calculated Watson's homogeneity U2 test using the function watson.two from the ‘CircStats' package (Zar 1999).

To examine seasonal variations in activity patterns, our study spanned approximately 18 months, capturing data across all four seasons – summer, fall, winter, and spring. Data were collected according to Southern Hemisphere seasons: summer (21 December from the previous year to 20 March from the next year), autumn (20 March to 21 June), winter (21 June to 21 September), and spring (21 September to 21 December). Seasonal variations were analyzed both on their temporal sequence and also on a seasonal basis (independently of the calendar year) to assess potential differences in activity patterns between pumas and tourists across distinct seasonal cycles.

Results

In total, 2606 photographs captured the presence of pumas, corresponding to 794 independent events and a detection rate of 8.7 per 100 trap days. On the other hand, tourists were identified in 11 049 photographs, resulting in 2801 independent events with a detection rate of 30.7 per 100 trap days. Seasonal variations were observed for the puma records, with the lowest number of events recorded in winter 2020 (67) and the highest in summer 2021 (144). For tourists, the smallest number of events occurred in winter 2020 (9) and the largest in summer 2022 (1246). Observations of pumas varied from solitary individuals to groups of up to four animals (a female accompanied by three cubs). In contrast, tourists comprised the passage of large groups exceeding 20 individuals, family excursions with minors, unauthorized bicycle use on designated paths, tourists traversing outside permitted hours, and tourists accompanied by pets, such as dogs off-leash.

Differences in daily activity patterns were observed between pumas and tourists. Pumas exhibited consistent activity across all seasons, with a peak observed at 19:00 hours, and minimal activity during midday hours. In contrast, tourists were more active between 10:00 and 18:00 hours, with fewer events recorded between 06:00–08:00 and 18:00–23:00 hours, when park trails are typically closed for hiking.

Temporal activity overlap (Dhat1) between pumas and tourists was 0.27 (CI: 0.23–0.28), which is considered low according to Monterroso et al. (2014). However, there are times of the day in which contact between both species could be more likely, which would be between 08:00–09:00 and 17:00–18:00 hours (Fig. 2; Watson's U2 value = 32.67, p < 0.001). Figure 2 shows the usage of walking trails by wildlife and visitors at multiple sampling points.

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The analysis of hourly activity overlap between pumas and tourists revealed a higher degree of overlap during autumn 2021, contrasting with a lower overlap during winter 2020 (Fig. 3). Furthermore, the autumn season of 2021 marked a pivotal moment in our study, characterized by a shift in overlap coefficients from lower values before this season to slightly higher values after it. It is important to note that the confidence interval for the winter of 2020 was considerably wider than the rest of the seasons due to the sparse data available associated with sampling commencing towards the season's end. Confidence intervals for subsequent seasons showed stability, underlining a consistent pattern of interaction across different seasons. The difference overlaps plots between sequential seasons and general seasons are shown in the Supporting information. The primary seasonal difference observed was that, at the beginning of the monitoring period, puma events outnumbered tourist events. However, following the summer of 2021, tourist events increased substantially, reaching nearly four times the number of puma events. This trend intensified by the summer of 2022, when tourist records were recorded up to 11 times more frequently than puma records, marking the summer as the season with the highest prevalence of tourists. In contrast, autumn and winter seasons displayed nearly equal numbers of records for both pumas and tourists.

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Discussion

Our study examined the activity patterns of pumas and tourists on trails within the Chacabuco Valley sector in the Patagonia National Park (Chilean Patagonia) providing insights on the temporal overlap between the two species. Pumas were predominantly nocturnal, while tourists were mainly active during daytime. Reports in Colombia and the central Andes (Peru and Bolivia) have shown that pumas can exhibit cathemeral behavior, with activities both during the day and night (Cáceres-Martínez et al. 2016, Viscarra et al. 2022). However, our study reports a predominance of nocturnal habits in pumas, possibly influenced by the presence of tourists or the availability of prey such as guanacos.

Before becoming a national park, the study area was one of the largest livestock ranches of Chilean Patagonia, where conflicts arose due to pumas preying on mostly sheep, leading to retaliatory killing of predators by ranchers (Zanón-Martínez et al. 2012, Herrera et al. 2017). This may have led pumas to develop avoidant behavior towards humans (Ordiz et al. 2021). For over 20 years there has been an absence of livestock farming and persecution, resulting in a relationship between pumas and humans that could be considered neutral so far, eventually evolving towards tolerance or habituation (Ohrens et al. 2016).

A notable overlap concentration between puma and tourists during twilight hours, particularly around dawn and dusk, was observed. This temporal overlap suggests specific times when interactions, such as visual encounters or sightings, might occur. Although stalking and attacks by pumas have not been reported within the park, the possibility of such incidents cannot be entirely dismissed. While puma attacks are considered rare, with a higher prevalence in North America and Canada (Beier 1991, Chester 2003, Larabee et al. 2010) than in South America (Ramírez-Álvarez et al. 2021, Taylor et al. 2023), reports of people attacked or killed in Chile can be found in the local press. Therefore, reducing interactions between pumas and tourists during dusk and after dawn hours could be an effective strategy to ensure the safety of park visitors.

Seasonal variations in the activity patterns of both pumas and tourists determined a greater overlap in autumn 2021, with lower values before this season and a slight increase compared to previous seasons after it. This pattern indicates an interaction varying across different times of the year, suggesting that it may be necessary to adjust management practices seasonally in order to effectively minimize human–wildlife conflicts (Sweanor et al. 2008). For example, during seasons with higher overlap, park administrators might consider implementing additional measures, such as increased awareness of protocols to tourists on how to handle encounters with pumas, ensuring they remain well-informed.

The influence of the COVID-19 pandemic on tourism activities within the park provides additional context to our findings. The reduction in tourism during the initial phases of the pandemic can be attributed to widespread lockdown measures in Chile, which restricted movement and access to national parks. Subsequently, with the easing of restrictions and the progressive reopening of national borders in 2021, there was an increase in tourism activity within the park, reaching up to around 9000 visitors in a month (CONAF unpubl.). This is consistent with our results, though on a smaller scale, showing a small increased activity overlap in the seasons following autumn 2021, compared to earlier seasons (Ramírez-Álvarez et al. 2021).

Regarding the limitations of our study, the distance between cameras may have been too small to fully encompass individual puma home ranges, which in Chilean Patagonia varies between 66 and 350 km² approximately (Elbroch and Wittmer 2012). Additionally, it remains uncertain whether puma activity patterns would differ in areas devoid of human presence. These limitations underscore the need for further research with increased spacing between cameras and studies in more remote areas to better understand puma activity pattern variations in the absence of anthropogenic disturbances. Future studies could also examine sex-related differences in activity, as evidence suggests potential distinctions in hourly activity patterns between female and male pumas (Azevedo et al. 2018, Allen et al. 2024).

While our study provides valuable insights into the activity patterns of pumas and tourists within the Chacabuco Valley in Patagonia National Park, it was only conducted on trails in specific public areas. Expanding this research to other locations and protected areas could provide a broader understanding of these interactions. Additionally, this research highlights the relevance of creating specific behavioral protocols for tourists and park administrators (Arroyo-Arce and Thomson 2019). Integrating proactive, positive, and engaging conservation messages into touristic experiences may foster a stewardship ethic among tourists, supporting the transition from unregulated tourism to a conservation-oriented model (Fernández-Llamazares et al. 2020, Janvier et al. 2023).

Conclusions

This research highlights the temporal activity patterns of tourists and pumas in the Chacabuco Valley, Patagonia National Park, where the time overlap is low but more intense during dusk and after dawn. Seasonal variations revealed that, although there was a peak in overlap during autumn 2021, the overlap before and after this period remained stable. These findings are significant for tourism management and conservation efforts in Patagonia National Park, Chile. Our study provides information for developing norms and protocols to promote safe and responsible behavior among tourists towards pumas. By understanding these interactions over time, we can devise more effective strategies to ensure safe coexistence with wildlife.

Acknowledgements

– We thank park rangers and fieldwork teams from CONAF and Rewilding Chile Foundation for the arduous work of installing the camera traps and recovering images. Also, we thank the reviewers of this journal for their feedback and constructive suggestions, which have contributed to the improvement of this manuscript.

Funding

– This research received no specific grant from any funding agency in the public, commercial, or not-for-profit sectors.

Author contributions

Liliana Guzmán-Aguayo: Conceptualization (equal); Data curation (lead); Formal analysis (lead); Investigation (equal); Methodology (equal); Validation (equal); Writing – original draft (equal); Writing – review and editing (equal). Cristián Saucedo: Conceptualization (equal); Investigation (equal); Methodology (lead); Project administration (lead); Resources (lead); Supervision (equal); Validation (equal); Writing – original draft (equal); Writing – review and editing (equal). Álvaro Verdugo-Martínez: Conceptualization (equal); Data curation (equal); Investigation (equal); Writing – review and editing (equal). Ingrid M. Espinoza-León: Conceptualization (equal); Investigation (equal); Supervision (equal); Writing – original draft (equal); Writing – review and editing (equal). Lorena A. Valenzuela-Lobos: Supervision (equal); Writing – review and editing (equal). Pablo Olmedo: Writing – review and editing (equal). Andrea Bahamonde: Writing – review and editing (equal). Julio Casanova: Writing – review and editing (equal). Benito A. González: Conceptualization (supporting); Investigation (supporting); Methodology (equal); Supervision (equal); Validation (equal); Writing – original draft (equal); Writing – review and editing (equal).

Data availability statement

Data are available from the Dryad Digital Repository: (Guzmán-Aguayo et al. 2025).

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