874 P. Singh et al.


FIG. 1 North-east India, with Dampa Tiger Reserve in Mizoram and locations where the dhole Cuon alpinus has been recorded, with corresponding reliability scores (see text for details).


in newspaper reports, scientific articles, grey literature (in- cluding species checklists), and reports by forest department personnel, local informants and naturalists working in the region. For each record we noted the type of evidence (direct/indirect), date of sighting, administrative status of location (protected/non-protected), and source person or reference. We assigned reliability scores for each record (1–5; with 1 being most reliable, and 5 least reliable; Supplementary Table 1). From December 2014 to March 2015, we deployed 79


pairs of Cuddeback Ambush IR camera traps (Model 1187; Cuddeback, De Pere, USA) across 80 km2 in the north-east of Dampa Tiger Reserve’s core area. At each station we placed two cameras facing each other, c. 30 cm above the ground, on either side of forest trails or on riverbeds (Singh & Macdonald, 2017). Mean inter-trap distance was 1.02 ± SD 0.33 km, with traps remaining active for an average of 64 days (range 3–91; Singh&Macdonald, 2017). Although the stations were intended to photograph wild felids, they also photographed other carnivores. Dholes generally use forest trails and riverbeds for movement, marking territories and hunting, and our sampling design therefore incorpo- rated areas used by the species. Weexamined site-use patterns through an occupancy ap-


proach that accounted for imperfect detection (MacKenzie et al., 2002). We treated each station as a site, and each trap-day as an independent temporal replicate. During exploratory analyses we calculated Moran’s I to check for spatial dependence of detections. Spatial dependence dropped beyond 2.3 km, with,10%of distance-pairs falling within the first distance class (Supplementary Fig. 1). We considered this to be negligible with respect to the total number of trap stations and detections, and therefore treated each station as an independent site. The detection


matrix therefore contained non-detections and detections (coded as 0 and 1, respectively) for 74 sites (data from five sites could not be retrieved), with a varying number of temporal replicates (3–91 days). We used photo-capture frequencies of key prey species


(sambar n = 236, muntjac n = 145, wild pig n = 92; predicted positive influence), distance to reserve boundary (predicted positive influence), photo-capture frequencies of forest department personnel (predicted positive influence) and photo-capture frequencies of other humans (predicted negative influence) as factors likely to influence site-use by dholes. We used trap effort (number of days a trap station was active) at each site as a covariate for detection probabil- ity, predicting that higher effort would translate to higher detectability. We tested singular and additive effects of the covariates, in which each represented an ecologically plausible hypothesis. Covariates were checked for cross- correlations and z-transformed prior to analyses. Models were ranked using Akaike information criterion corrected for small sample sizes (AICc; Burnham & Anderson, 2002). Parameter estimation and model comparisons were calcu- lated with PRESENCE 11.9 (Hines, 2006). We obtained presence records from 80 locations for


1990–2018, of which we considered 41 records from 2010– 2018 with reliability scores of 1–3 (Supplementary Table 2). In the case of multiple records for the same site, we consid- ered the most recent record with the highest reliability score. Most records were from Arunachal Pradesh (n = 14) and Assam (n = 8), with five records from Mizoram and Nagaland, four from West Bengal, three from Meghalaya and two from Sikkim. There were no recent records of dholes from Manipur and Tripura. A total of 5,033 camera trap-days in Dampa Tiger Reserve generated 500 photo- encounters of dholes, comprising 92 detections (one per


Oryx, 2020, 54(6), 873–877 © 2019 Fauna & Flora International doi:10.1017/S0030605319000255


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