Wild felids in Sarawak, Malaysian Borneo 257


2010), and thus reflect the tolerance of particular species to forest disturbance (Griffiths & Schaik, 1993; Norris et al., 2010). We detected differences in activity patterns be- tween protected and unprotected areas for some species, but these differences were small. Likewise, Bernard et al. (2013) found no impact of anthropogenic activities on the activity patterns of a suite of mammals in Imbak Canyon Con- servation Area, Sabah. Lowland dipterocarp forest appears to be the preferred


FIG. 4 The influence of elevation and distance from road, river and longhouse on occupancy of felid species in Sarawak. Black dots indicate the occupancy model coefficient for each species, black horizontal lines the 89% Bayesian credible intervals. Grey horizontal lines indicate significance. Vertical lines and shaded areas show the mean and 89% Bayesian credible intervals for community-level estimates, respectively. Vertical dashed lines show the zero-effect size.


sites (Ψ= 0.411) than in protected areas (Ψ= 0.155), with most of the captures in Ulu Kapit, central Sarawak, a logging concession; although not officially protected, this area con- tains large tracts of continuous lowland forest, which could be one of the reasons for the relatively high occurrence of bay cats there. Past studies have also recorded this spe- cies in production forests such as Sela’an Linau Forest Management Unit (Mathai et al., 2014) and Muput Forest Management Unit (Hon, 2011) in Sarawak, several sites in Sabah (Wearn et al., 2013; Hearn et al., 2018) and East Kalimantan (Sastramidjaja et al., 2015). Our findings con- firm that production forests, as part of a landscape that also contains large areas of intact forest, can contribute to sustaining species of conservation importance. Our data corroborate the findings of previous studies suggesting that felids exhibit diel activity patterns ranging from strongly nocturnal to largely diurnal. The activity pat- terns of the clouded leopard, and the marbled, bay and leo- pard cats, were consistent with those observed by Hearn et al. (2018), with marbled and bay cats being diurnal, clouded leopard cathemeral and leopard cat nocturnal with minimal activity after dawn or before dusk (Fig. 3). The diel activity of the flat-headed cat in our study was consistent with that observed by Adul et al. (2015) in central Kalimantan (Fig. 3); the species is nocturnal and is often detected during surveys by boat at night (Hearn et al., 2010; Mohd-Azlan &Thaqifah, 2020). Temporal activity rhythms of larger tropical mammals can be shaped by habitat deg- radation and other anthropogenic activities (Norris et al.,


habitat type of the felids in Sarawak, with the exception of the flat-headed cat. Felids appear to be widely distributed along elevation gradients up to 1,700 m. The flat-headed cat is closely associated with water bodies and usually only recorded in lowland areas (Bezuijen, 2000; Cheyne et al., 2009; Hearn et al., 2010; Wilting et al., 2010, 2016; Wadey et al., 2014; Mohd-Azlan & Thaqifah, 2020), although the species has been detected at elevations of up to 780 min Sabah (Brodie & Giordano, 2011). Consistent with this, most detections of the flat-headed cat in our study were along riverbanks; multi-species occurrence models showed the effects of river proximity (positive) and elevation (nega- tive) on flat-headed cat occurrence (Fig. 4). Elevation is often used as a proxy for changes in habitat


characteristics, such as decreasing temperature and chan- ging forest stature from lowlands to highlands (with asso- ciated transitions in plant species composition); there are also probably changes in the density of prey for carnivores (Mohamad et al., 2015). Felids and other medium-sized and large lowland animals may use higher elevations to avoid anthropogenic activities (Tan et al., 2017); it is therefore critical to preserve mid-elevation habitats as refugia from lowland habitat destruction and climate change (Brodie, 2016; Brodie et al., 2017; Scriven et al., 2020; Williams et al., 2020). Wetlands, including peat swamp forest, are also important sites for conserving felids and numerous other species, despite frequently being considered wastelands deficient in nutrients and low in species diversity (JM-A, pers. obs., 2021). Wetlands have received relatively little conservation attention in Sarawak and many have been converted to other land uses (Yule, 2010; Posa et al., 2011; Tsuyuki et al., 2011). A previous study reported that the bay cat was associated with rivers (Mohd-Azlan & Sanderson, 2007). However, we found the occurrence of bay cats was not closely associated with rivers, but rather influenced by roads; occurrence of this species increased .10 km from roads (Figs 4 & 5a). This species also appears to be distributed mostly at intermediate elevations, consis- tent with the findings of Hearn et al. (2018), although it has been detected as high as 1,460 m in Pulong Tau National Park, Sarawak (Brodie & Giordano, 2012). Similar to the bay cat, the occurrence of the marbled cat increased sharply at distances of .25 km from roads (Fig. 5a). Occurrence of the clouded leopard appeared to be associated with rivers and elevation (Figs 4 & 5c), but not with roads. This is in


Oryx, 2023, 57(2), 252–261 © The Author(s), 2022. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605321001484


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