Wild felids in Sarawak, Malaysian Borneo 255
2013). We used the occurrence data and clipped environ- mental layers to model the distribution of all species using maximum entropy with a jackknife analysis in MaxEnt 3.4.1 (Phillips et al., 2017). MaxEnt assesses the distribution probability of the maximum entropy of target species by inferring the occurrence data even with incomplete information (Phillips et al., 2006; Elith et al., 2011).
Results
Detections in protected and unprotected sites We identified 3,545 photographs of felids from a total of 2,628,993 images (c. 0.1%); 863 of these were independent captures. Among the five felids, the highest number of inde- pendent captureswas of the leopard cat (681), followed by the marbled cat (94), clouded leopard (56), Bornean bay cat (21) and flat-headed cat (11; Supplementary Table 2). Felids were detected at 332 of 845 camera stations (39.3%). The leo- pard cat was detected most often (85 stations in protected and 141 in unprotected areas), followed by the marbled cat (23 stations each in protected and unprotected areas), cloud- ed leopard (21 stations in protected and 14 in unprotected areas), bay cat (eight stations in protected and 13 in un- protected areas) and flat-headed cat (six stations in protected areas). Felids were detected in 26 of the 31 study areas (83.9%; 19
protected and seven unprotected sites with at least one spe- cies; Supplementary Table 2). We did not detect any felids in SamunsamWildlife Sanctuary, Sama Jaya Forest Reserve, and Kuching Wetland, Maludam and Similajau National Parks. In none of the study sites did we record all five Bornean felids. Four felids (all except the flat-headed cat, a wetland specialist) were recorded in two unprotected areas (Kapit and Baram). The flat-headed cat was recorded in only two protected peat swamp forests (Loagan Bunut and Ulu Sebuyau National Parks). Cameras in six sites re- corded the leopard cat only: Bako, Bungoh Range, Dered Krian, Santubong and Tanjung Datu National Parks and Ulu Trusan. In Sungai Meluang National Park only the marbled cat was recorded. The bay cat was detected at nine study sites (29.0% of the total), with no more than two independent photographs per site, except in Kapit, where there were 11 independent detections. The largest felid, the clouded leopard, was detected in nine study areas (29.0% of the total), with the highest number of in- dependent detections (N = 19) in Pulong Tau National Park (Supplementary Table 2). The leopard cat was detected at the highest percentage of study sites (32.9% of locations), followed by bay cat (28.3%), marbled cat (10.7%), clouded leopard (10.6%), and flat-headed cat (4.6%; Fig. 2). The marbled cat and clouded leopard had higher probabilities of occurrence in protected than unprotected areas, and vice versa for the leopard cat and bay cat (Fig. 2).
FIG. 2 Estimated probabilities of occurrence of the five felid species in Sarawak in protected and unprotected study areas. Black points indicate mean occurrence, horizontal lines show the interquartile range, and vertical lines indicate the mean occurrence of all species.
Activity patterns
The marbled and bay cats weremostly diurnal, and the leo- pard cat was predominantly nocturnal. The clouded leopard and flat-headed cat were cathemeral, although clouded leo- pard activitywasmostly at night,with only fewcaptures during the day, and the small number of flat-headed cat detections (N = 11) peaked around midnight (Fig. 3). Except for the flat- headed cat, whichwas only recorded in protected areas, activ- ity patterns for the felids did not differ substantively between protected and unprotected study sites, with activity overlap ranging from Δ
^1 = 0.61 to 0.80 (Fig. 3). Occurrence patterns
Although the detection rates of all species except the leopard cat were relatively low, we found some statistically signifi- cant associations between habitat variables and species occurrence. The occurrence of the marbled (mean model coefficient = 1.60, 89%CI = 0.51, 2.92) and bay cats (mean = 2.50, 89%CI = 0.92, 4.65) increased at greater distances from the nearest road (Figs 4 & 5a). The occurrence of the leopard cat (mean = 6.03, 89%CI = 2.29, 10.04), flat-headed cat (mean = –5.94, 89%CI = –12.61, 0.00) and clouded leopard (mean = –6.30, 89%CI = –10.85, –1.98) were signifi- cantly affected by distance to river (Fig. 4). Leopard cats and flat-headed cats were more likely to occur near rivers, whereas clouded leopards were more likely to occur away from rivers (Fig. 5b). Elevation influenced the occurrence of the flat-headed cat (mean = –5.47, 89%CI = –10.98, –1.33) and clouded leopard (mean = 1.49, 89%CI = 0.81, 2.29; Fig. 4), with flat-headed cats having a higher probabil- ity of occurrence at low elevation and the probability of oc- currence of the clouded leopard increasing slightly above 900 m(Fig. 5c). The occurrence of none of the felids was af- fected by the distance to the nearest longhouse (mean = 0.11, 89%CI = –1.44, –1.70). Except for the flat-headed cat, Bornean felids were
distributed across a relatively wide range of elevations (up to and above 1,000 m; Fig. 6). The flat-headed cat
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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