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Mammal responses to tourism 855


values. These negative impacts could compromise the con- servation goals of protected areas and the ecosystem services they provide, including tourism (Turton & Stork, 2008; Leung et al., 2018). Therefore, biodiversity monitoring pro- grammes should be a priority in protected areas that have been opened for visitors, particularly where important bio- diversity values overlap with high tourism potential. Such monitoring can serve as an early warning system for the need to change management schemes to promote the long- term maintenance of species (Yoccoz et al., 2001). However, tourism-driven impacts are difficult to measure (Buckley, 2003) and the lack of data collected before the intensification or beginning of tourismmakes these assessments even more challenging (Butsic et al., 2017). This is the case for national parks in Brazil, where tourism has been growing at an annual rate of 10% (ICMBio, 2019) but studies assessing the impacts of visitors on biodiversity remain scarce (Cunha, 2010; Silva et al., 2018; Monteiro & Lira, 2020). Here we used a quasi-experimental setting to investigate


the potential impacts of carefully planned nature-based tourism on mammal species at Cavernas do Peruaçu National Park, a high-priority area for conservation in Brazil (Ministério do Meio Ambiente, 2018). We surveyed the mammal community using camera traps on tourist and non-tourist trails before and after the Park officially opened for visitors. To our knowledge this is the first study of this type in a Brazilian national park using baseline data collected before the intensification of tourism. According to the risk–disturbance hypothesis (Frid & Dill, 2002) and previous assessments conducted elsewhere (Rogala et al., 2011; Zhou et al., 2013), we expected that some species would avoid or limit their use of tourist trails after visitors were allowed into the Park, causing a decline in species rich- ness and their probability of trail use. Given that anthropo- genic pressure can also modify the activity patterns of species (Marchand et al., 2014;Gaynor et al., 2018)we antici- pated that the impacts of visitors could also lead to the tem- poral displacement of mammals. Notably, we expected that species would be less active and would show reduced diurnal activity on tourist trails after the beginning of tourism. Our intent with this assessment is not to jeopardize tourism but to inform effective management strategies that facilitate both biodiversity conservation and the development of low- impact tourism in the region.


Study area


This study was conducted in Cavernas do Peruaçu National Park (Fig. 1), in south-eastern Brazil in the ecotone between Cerrado (Neotropical savannah) and Caatinga (a mosaic of thorn scrub and seasonally dry forests associated with a semi-arid climate; Leal et al., 2005). The 568 km2 Park pro- tects extensive areas of dry forests and woody savannah and


supports 70% of all large mammals found in the Brazilian Cerrado (Ferreira & Oliveira, 2014). The Peruaçu River is the main source of water in the Park and its valley harbours a unique speleological system with hundreds of caves and archaeological sites with major tourism potential. Gallery forests along the river and dry forests are the main vegeta- tion types in the river valley (Oliveira-Filho & Ratter, 2002). The climate is semi-arid, with a mean annual temperature of 24.4 °C and a totalmean annual rainfall of 925mmcon- centrated in the wet season (mid October–March; Geoclock, 2005). Given that caves are the main tourist attraction in the


Park and that these are also extremely fragile ecosystems, the potential negative impacts from tourist activity have long been a concern for those managing the Park. As such, a carefully designed plan for tourism was included in the Park’s management plan (Geoclock, 2005). Tourist visitation is restricted to the Peruaçu River valley in the cen- tral region of the Park and consists of guided visits to caves and rock art panels, which are accessed via dedicated trails. Before their visit, tourists must hire a certified local tour guide. They are then registered and informed about the rules in the Park, particularly restrictions on accessing non- tourist areas and walking off-trail. Each guide can host eight visitors at a time and there is a daily limit on the number of visitors allowed on each tourist route. The Park remained closed to tourism until roads, walkways, visitor centres, and other tourist infrastructure were improved or built, but a small number of visitors (200–600 per year) were al- lowed on a few tourist trails and caves during a pre-opening pilot scheme. The Park officially opened to tourists in 2015 (Fig. 1) and visitation increased substantially, reaching almost 7,000 tourists in 2017 (Supplementary Table 1).


Methods


Camera-trap surveys To investigate the potential effects of visitors on the mammal community in the Park, we set passive infrared camera traps (Bushnell Trophycam, Bushnell Corporation, Overland Park, USA) at 16 sampling sites on pre-existing trails in tourist and non-tourist areas (mean minimum distance to the nearest sampling site was c. 0.77 km; Fig. 1, Supplementary Table 2). We conducted surveys during 2011–2017 restricted to the Peruaçu River valley, where all tourist routes are located (Table 1). At each site we deployed camera traps 1–2mfrom the trail at a height of c. 30 cm, par- allel to the ground and aimed at the trail. We set the camera traps to work continuously and record 10-s videos when triggered, with 30-s intervals between triggers. We removed thin vegetation directly in front of the cameras to prevent false triggers. We conducted maintenance to replace SD


Oryx, 2022, 56(6), 854–863 © Crown Copyright, 2022. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605321001472


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