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826 M. C. Kaizer et al.


(Ribeiro et al., 2009). Over 300 species of mammals occur in the Atlantic Forest, with c. 30% of these being endemic (Paglia et al., 2012; Quintela et al., 2020). Although the mammals of the Atlantic Forest are mostly arboreal (Paglia et al., 2012), themajority of studies that have focused on mammals .1 kg have been conducted using ground- based methods, such as transect censuses and terrestrial camera traps, along with the use of indirect evidence obtained from vocalizations, tracks, faeces and carcasses (Chiarello, 2000; Srbek-Araujo & Chiarello, 2005; Oliveira et al., 2013; Geise et al., 2017). Given that loss of mammals has been widely documented throughout the Atlantic Forest biome (Canale et al., 2012; Galetti et al., 2017; Sousa & Srbek-Araujo, 2017; Bogoni et al., 2018, 2020) and prob- ably will continue to increase because of ongoing anthropo- genic activities and climate change, there is a need to gather more reliable data on the distribution and population status of arboreal mammals to inform conservation plans. Camera traps have proven to be an effective non-invasive


method to detect rare and elusive species, even in remote areas and over large spatial and temporal scales (Burton et al., 2015;Wearn&Glover-Kapfer, 2019). Although camera traps have become a ubiquitous method in ecological studies and conservation programmesfor terrestrialmammals (Glover-Kapfer et al., 2019), with great potential for global net- work monitoring (Ahumada et al., 2011; Steenweg et al., 2017), only recently has this method begun to be applied to surveying arboreal mammals in tropical forest canopies (Olson et al., 2012;Gregory et al., 2014;Whitworth et al., 2016;Bowler et al., 2017; Kaizer, 2019;Hongo et al., 2020;Moore et al., 2020). Here we present the first study using arboreal camera trapping to survey arboreal mammals in the Atlantic Forest. Our aims were to (1) assess the efficiency of camera traps for inventorying arboreal mammals in two forest types (semide- ciduous and ombrophilous forest) and (2) examine variation in the species richness, relative abundance, community com- position and functional traits of the arboreal mammal assem- blage in Caparaó National Park, Brazil. Although Caparaó National Park is one of the last significant Atlantic Forest remnants in terms of size, in south-east Brazil there is a lack of studies on the vertebrate biodiversity of the Park and its arborealmammal community is largely undocumented.


Study area


The 31,853 ha CaparaóNationalParkliesonthe borderbe- tween the states ofMinas Gerais and Espírito Santo in south- east Brazil (Fig. 1). The protected area is within the Caparaó massif, part of the northern Mantiqueira mountain range, and stretches for c. 40 kmfromnorthtosouth,withaltitudes of 630–2,892 m. (ICMBio, 2015). The vegetation types include Mountainous and High Mountainous Ombrophilous Dense Forest and Mountainous Semideciduous Seasonal Forest below1,500m, cloud forest at 1,500–1,900mand high-altitude


grasslands above 1,900 m(Veloso et al., 1991;ICMBio, 2015). Semideciduous Seasonal Forest occurs predominantly on the western side of the Park and Mountainous Ombrophilous Dense Forest mostly on the eastern side (ICMBio, 2015). The landscape surrounding the Park is dominated by coffee plantations, pastures and isolated, small forest patches. The climate is humid with a temperate summer (Alvares et al., 2013).Mean annual temperature is c. 19 °C at lower altitudes and 9.4 °C at higher altitudes (Alvares et al., 2013). Mean an- nual rainfall is c. 1,500 mm and the air relative humidity is high (.70%) during most of the year (ICMBio, 2015). There is a rainy season during October–April and a dry and cool season, with monthly rainfall ,50 mm, during May– September(Alvareset al., 2013;ICMBio, 2015).


Methods


Infrared camera traps (Bushnell Trophy Cam, Bushnell, Overland Park, USA) were deployed in Caparaó National Park during January 2017–June 2019 as part of a larger pro- ject monitoring the resident northern muriqui Brachyteles hypoxanthus population (Kaizer, 2019). We surveyed 24 sites (each with a single camera trap) in the canopy in seven valleys within the Park (Aleixo, Calçado, Facão de Pedra, Santa Marta, Rio Norte, Rio Preto and Rio Veado). Survey sites covered an altitude range of 1,000–1,768 m, with 10 sites on the western side of the Park (Montane Semideciduous Seasonal Forest) and 14 sites on the eastern side (Montane Ombrophilous Dense Forest). In 2017,we placed eight cameras each in two of these valleys (one in the west and one in the east) along an altitude gradient (Fig. 1; Kaizer, 2019). In 2018 and 2019, we placed eight more camera traps in the other valleys within the Park (Fig. 1). We placed the arboreal camera traps .250 m apart, strapping them to trees at a mean height of 12.0 ± SD 3.1m(range: 7.5–17.0 m) from the ground. We chose the ar- boreal camera-trap locations independently of tree species, based on tree connectivity; i.e. trees connected to at least three other trees where animals could cross the canopy and that were considered to offer safe access for climbers (Gregory et al., 2014; Kaizer, 2019; Whitworth et al., 2019). To prevent bias, we chose trees that were not fruiting or flowering on the day of installation, as food resources could attract certain species more than others and thus affect detection rates. We did not bait the camera traps or orientate them to the east or west (which would avoid direct sunlight and reduce shadows and false positive/negative triggers) and we placed them facing along a horizontal branch of the tree or towards a horizontal or vertical branch of an adjacent tree. The camera traps were active continu- ously and set to hybrid mode (two photographs and one 30-s video on each trigger event), with 10-s intervals between triggers and low night-time light-emitting diode intensity. We identified animals in the photographs using Wild.ID


Oryx, 2022, 56(6), 825–836 © The Author(s), 2022. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605321001563


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