Pack size, home range size and activity pattern of dhole (Cuon alpinus) in Mudumalai Tiger Reserve, India: effects of anthropogenic environment use and reproduction

Pack size, home range size and activity pattern of dhole (Cuon alpinus) in Mudumalai Tiger Reserve, India: effects of anthropogenic environment use and reproduction | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Pack size, home range size and activity pattern of dhole (Cuon alpinus) in Mudumalai Tiger Reserve, India: effects of anthropogenic environment use and reproduction Shuta Sawaguri, Raman Sukumar, Shiro Kohshima This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6674422/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 9 You are reading this latest preprint version Abstract Understanding the ecology of endangered dhole and their responses to anthropogenic environments is important for mitigating conflicts with humans and ensuring conservation efforts for this species. This study investigated the pack size, home range size, reproduction and activity pattern of multiple dhole packs in Mudumalai Tiger Reserve, southern India, from September 2011 to August 2015 using direct observation and camera trapping. The findings were compared across dhole packs to assess the influence of reproduction and human-modified environments on their ecology and behaviour. Of the five packs identified (P1–P5), reproduction was observed three times in two packs (P1, P2) with an average litter size of seven and denning between December and February, while the others likely did not reproduce. Group sizes observed outside the den area (mean: 6.6 ± 3.4) decreased from November to December, likely due to pup dispersal and den defense needs. Packs inhabiting the areas with high proportion of anthropogenic environments and feeding on livestock and garbage (P1, P2) had significantly smaller home ranges (7.0–14.4 km²) compared to those in more natural environments (P3, P4: 36.1–40.3 km²), suggesting higher foraging efficiency in human-modified landscapes. The dholes in the study area were primarily diurnal, with 74.4% of activity occurring during daylight hours. They were particularly active in the morning and evening as they frequently took very long rests over six hours during midday. However, the pack with the greatest use of anthropogenic environments (P2) showed cathemeral activity pattern increasing night activity up to 50.5%, probably to avoid encounters with humans. The high mortality of pups estimated in P2 and unnatural deaths of adults observed both in P1 and P2 suggest that the nutritional benefits and increased reproduction in anthropogenic environments could be offset by the higher mortality rates, potentially due to conflicts with humans and diseases transmitted by feral or domestic dogs. habitat utilisation livestock predation garbage scavenging nocturnal activity anthropogenic impacts human-wildlife conflict. Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Figure 7 Introduction The decline of vertebrate species worldwide has been driven by a range of anthropogenic factors, including habitat destruction, fragmentation and overexploitation of natural resources. Additionally, increasing human populations have intensified conflicts with wildlife, while climate change further exacerbates biodiversity loss (Cardillo et al., 2004 ; Newbold et al., 2015 ; Ripple et al., 2015 ). To address these challenges and conserve vertebrate biodiversity, it is essential to mitigate human-wildlife conflicts by understanding the ecology of concerned species sharing their habitats with humans and how human-modified landscapes influence their ecology and survival. The dhole ( Cuon alpinus ), also known as the Asiatic or Indian wild dog, is an endangered species that requires urgent research attention and conservation efforts. This canid is currently found in India, Southeast Asia and China. According to the International Union for Conservation of Nature (IUCN), the estimated wild population of mature dholes between 949 and 2,215, with the numbers continuing to decline due to habitat loss and transformation, retaliatory killings linked to livestock predation and other anthropogenic pressures (Kamler et al., 2015 ). While understanding dhole ecology is crucial for conservation, such research particularly in human-modified habitats remains insufficient. Although fewer field studies have been conducted on dholes compared to other canid species such as wolves (Srivathsa et al., 2020 ), several investigations in India have provided valuable insights into their ecology. These include studies on the reproduction and social structure (Johnsingh, 1982 ; Venkataraman 1998 ), movement patterns (Acharya, Johnsingh & Sankar, 2010 ), daily activity cycles (Karanth et al. 2017 ; Krishnakumar et al. 2022 ; Ramesh et al. 2012), habitat utilisation (Srivathsa et al., 2014 ), population estimates using camera traps (Punjabi et al., 2022 ), foraging ecology (Venkataraman, Arumugam & Sukumar 1995 ) and the relationship between prey-predator dynamics and pack size (Bhandari et al., 2021 ). However, much of these existing research rely on either unidentified packs or a very limited number of identified packs. Moreover, there has been little research on how anthropogenic environments affect dhole ecology. To address these gaps, we conducted field research in the Mudumalai Tiger Reserve, India, from September 2011 to August 2015, utilising direct observation and camera trapping. We investigated key ecological parameters, including pack size, home range, activity patterns and reproduction. Additionally, we compared four identified dhole packs with varying reproductive statuses and degrees of anthropogenic environment use to evaluate the influence of reproduction and human-modified landscapes on their behaviour. The results of this study, which provide important ecological insights into their adaptability to changing environments, could contribute to a more effective conservation strategy for this endangered species. Methods Study area This study was conducted in the Mudumalai Tiger Reserve (MTR) in Tamil Nadu, southern India (Fig. 1). MTR is a part of the Nilgiri Biosphere Reserve situated in the Western Ghats, covering an area of 321 km 2 (11°32’–11°43’ N and 76°22’–76°45’ E), with the elevation between 800 and 1,100 m asl (Dattaraja et al., 2013). The average maximum and minimum temperatures are 26.6 ± 2.34°C and 15.9 ± 1.95°C, respectively (Suresh & Nanda, 2021). The region experiences three distinct seasons: the dry season from January to April, the southwest (summer) monsoon from June to September and the northeast (winter) monsoon from October to December (Dattaraja et al., 2013). The average annual rainfall is 1,681 mm in the western part and 721 mm in the eastern part of the reserve (Dattaraja et al., 2013). Vegetation varies by region: tropical moist deciduous forests in the west, dry deciduous forests in the central part and dry thorn forests in the east (Suresh et al., 2011, Dattaraja et al. 2018). The eastern part has more human population and human-altered environments such as agricultural fields and grazing lands than the central and western parts which adjoin coffee and tea plantations. In addition to dhole, large predators including tiger ( Panthera tigris ), leopard ( Panthera pardus ) and striped hyena ( Hyaena hyaena ) are seen in this area. Other larger mammals in Mudumalai include Asian elephant ( Elephas maximus ), gaur ( Bos gaurus ), chital ( Axis axis ) and sambar ( Rusa unicolor ) deer, black buck ( Antilope cervicapra ), wild boar ( Sus scrofa ), sloth bear ( Melursus ursinus ), bonnet macaque ( Macaca radiata ) and southern plains gray langur ( Semnopithecus dussumieri ) and many smaller mammals such as several species of squirrels, mongooses and civets. Data collection Direct observations We conducted fieldwork to observe wild dholes directly over a period of 467 days from September 2011 to August 2015 (see Tables 1a & 1b). We searched for dholes on foot or by vehicle approximately from 06:00 to 11:00 (or 06:30–11:30) and from 14:00 to 18:00 (or 14:30–18:30) based on recent sightings and field signs such as scats and footprints. Whenever we spotted dholes, we followed them as far as possible and observed them maintaining a distance of at least 10 m. Their behaviour was recorded using a video camera (SONY HDR-XR550V or PJ630V). We also recorded the location of each observation with a GPS device (Garmin GPSMAP 60CSx) every 15 minutes. Video and camera trapping We conducted video trapping over a total of 1,520 camera days from July 2014 to August 2015 (Table 1b). The video trapping was carried out using automatic recording video cameras (Bushnell 8MP Trophy Cam HD Max Black LED Trail Camera with Night Vision). We installed one or two video cameras (a total of 114 video cameras) at 37 locations: 19 junctions where dholes had recently been sighted or field signs had been found, 10 trails with scats evidence, seven ponds with footprints, and one site with a prey carcass (Fig. 1). The video cameras were securely fixed to tree trunks or other structures at 0.5–1.5 m agl, angled slightly downward. They operated 24 hours a day, with each video recording set to 60 seconds and an interval of one second between captures. We collected data and replaced batteries every two weeks to one month. During these visits, we also replaced SD cards and noted the start and end times of recording. If the SD card was full, or in the event of battery depletion, the timestamp on the last recorded video was considered as the end time of the recording. We also used the camera trapping data collected by the Tamil Nadu Forest Department for over 12,884 camera days from November 2014 to March 2015 (Table 1b). They installed a total of 1070 camera traps (Cuddeback, Attack) at 107 locations within the study area, with two devices set up at each site (Fig. 1). Similar to our setup, these cameras were fixed to tree trunks or other structures at 0.5–1.5 m agl, facing slightly downward and operated continuously 24 hours a day. We specifically used the photo data that included images of dholes for this study. Identification of animals and packs We identified at least one adult from each pack based on their distinct morphological features such as facial patterns and body colour (Supplementary Information SI1). If the identified animal was observed in a group, we treated the observed group as an identified pack (P1–P4). Among the five packs we identified, one group had no identifiable individuals but it was treated as an identified pack (P5) because the number of adult individuals was clearly higher than in any other identified packs. All the other groups we observed were categorised as unidentified packs. Data analysis Definition of group size, pack size and pups Group size was defined as the number of animals observed during each direct observation outside the vicinity of the den. Pack size, on the other hand, was defined as the number of animals in the identified group including the number of pups in the den at the time of observation. Individuals with a body length of approximately 30–90 cm were considered as pups less than one year old. Small pups with a body length of about 20–30 cm and a dark brown body colour were considered as pups less than one-month-old just starting to come out of the den. Estimation of the denning season The denning season, the period when new born pups stay in or around the den, was estimated to occur from December to February based on our observation of pups in and around the dens (P1: 24 December, 2014–19 February, 2015, P2: 8–22 January, 2014 and 17 January–15 February, 2015) and previous reports on their birthing season (November–March, peaking in December; Davidar, 1975) and den usage (10–11 weeks after birth, with pups peeking out at three weeks; Johnsingh, 1982). Estimation of home ranges We plotted GPS points obtained from direct observations and from camera and video traps using QGIS, an open-source GIS software (Ver. 3.4.9, Madeira). During the direct observations, we recorded the location every 15 minutes from the initial sighting until the animals were no longer visible. We then mapped the home ranges using the minimum convex polygon (100% MCP) method. Activity pattern To analyse the dhole activity patterns, we calculated relative abundance indices (RAIs, the number of camera capture events in each time period of the day × 100 / total camera trap days; O’Brien et al., 2003). In our analysis of the camera trap records, all camera capture records occurring within a 30-minute window were treated as a single camera capture event. We classified behaviours observed through direct observation and video trapping into six behavioural categories: resting, locomotion, feeding, excretion-associated, social and others (see Table 5). In direct observation, we defined a long resting behaviour of more than one hour, carried out by all members of each pack, as 'long resting'. When analysing the diurnal activity pattern, if any member of the observed pack exhibited behaviour from any above categories, we recorded it as an event for that behavioural category by that pack. We then calculated the ratios of each behavioural category during different time periods of the day (the number of events of each behavioural category / total recorded events). In the analysis of the diurnal activity pattern, daytime (light period) was defined as the period from sunrise to sunset, while nighttime (dark period) was defined as the period from sunset to sunrise (refer to Fig. 1b). In the graphs illustrating the diurnal activity patterns, we standardised the sunrise and sunset time to 06:30 and 18:30 hr, respectively, for convenience. Statistical analysis For the comparisons of activities between daytime and nighttime of the local population and each pack (Table 4), we applied the Chi-square test (two-sided; for multiple comparisons, with p-values corrected using the Benjamini-Hochberg method). Mann-Whitney U test was applied for the analysis of the seasonal change in average number of adults in observed group (Table 3) and in the mean distance between sighting locations (Table 4). Wilcoxon signed-rank test was applied for the comparison of active and inactive behaviours between day and night (Fig. 6). We used R Statistical Software (Ver. 3.1.1) for all analyses. The average values are represented as mean ± standard deviations and range values are represented as (lowest–highest). Results Observed group/pack size, breeding events, pups and adults In the direct observations, 467 days of observation effort resulted in 342 tracking events, totaling 492.69 hours of direct observation (Table 1a; Table 1b). A total of 55 individuals including 41 adults and 14 pups were observed during the camera trap survey period (Table 2). During the direct observation, we identified eight adults. Based on the pack sizes and the presence of the identified animals in the pack, we could identify five packs (P1–P5) in the study area (Table 2). The mean group size of the local population observed outside the vicinity of the dens in the study area including unidentified packs was 6.3 ± 3.4 individuals (range: 1–16, N = 268) with an average of 5.7 ± 3.0 adults (range: 1–16, N = 268) and 0.7 ± 2.1 pups (range: 0–6, N = 268). The mean group size of the five identified packs was 6.9 ± 3.4 individuals (range: 1–16, N = 262) with an average of 5.8 ± 2.9 adults (range: 1–16, N = 262) and 0.8 ± 2.5 pups (range: 0–6, N = 262). Three breeding events in packs P1 (one event) and P2 (two events) were confirmed (Table 2). A total of 21 pups were observed including six pups in P1 on 24 December, 2014, seven of P2 on 8 January, 2014 and eight of P2 on 22 January, 2015. The average litter size was estimated to be seven (range: 6–8). In P1, where six newborn pups were born in December 2014, five pups (aged 9–10 months) that had grown to near adult size (third-forth) were observed joining adults by 14 August, 2015. Thus, it is likely that P1 had successfully bred. In contrast, P2 had seven and eight newborn pups in January 2014 and 2015, respectively, but only four and six pups (aged 3-4 months) were observed in March (22th in 2014 and 18th in 2015), respectively. As there have been no observations since then to confirm the survival of these pups, it is likely that P2 failed to raise any pups to adulthood in both 2014 and 2015. In P3 and P4, no pups or immature individuals with the body size remarkably smaller than adults were observed, suggesting that these packs did not breed or failed to breed at the early stage of their reproduction during the study period. In P1 and P2, the average number of adults in the groups observed outside the vicinity of the den decreased during the denning season (December–February) (Table 3). This value was also significantly reduced in the local population of the study area, including other identified and unidentified packs. These results suggest that one or two adults stayed around the den with their pups during the denning season. During the denning season in 2014 and 2015, the video trapping cameras at P2's den frequently recorded a female (probably the mother) and a male staying around the den, apparently on guard duty. The number of adults in P1 and P2 decreased during the study period as a result of deaths of unknown causes (Table 2). In P1, the carcass of an adult member vomiting a large amount of blood from the mouth was found beside a dead wild boar (k1 in Fig. 3a) on 24 December, 2014. Boar meat fragments were found on the ground and no external injuries were observed in the dead body. Subsequently, a reduction in the number of adults in P1 was confirmed on the same day. Although we could not observe directly, we were also informed by local people that carcasses of other carnivorous animals such as striped hyenas and vultures were found near the spot where we found the dead dhole during the same period. In P2, we found the carcass of three identified male adults without any external injuries in a small area within 20–30 m 2 (k2 in Fig. 3a) on 25 July, 2015. They also vomited blood from their mouths. After this event, we could never observe P2. Both sites where we found the carcasses were close to villages (within 50 m from each village). These facts and circumstances suggest that the cause of their deaths may have been unnatural ones such as artificial poisoning. In P1 and P2, reductions in the number of adults were also observed, possibly due to causes other than those mentioned above. In P1, the number of adults decreased from 11–12 to 4–5 on 23 November, 2014 before their breeding in 2014 (24 December, 2014). In P2, it decreased from 9 to 4–6 on 7 December, 2013 before their breeding in 2014 (8 January, 2014). However, such reduction in adult number was not observed before the breeding of P2 in 2015 (22 January). In contrast, no reduction in the number of adults for any reasons was observed in P3 and P4. Home ranges We mapped the home ranges of four identified packs (P1–P4) (Fig. 2) based on a total of 1,247 recorded location points in total: 806 points from direct observation (Table 2), 227 points from video capture (Table 1b) and 214 points from photo capture (Table 1b). For P5, however, we could not obtain enough location points to determine its home range or estimate its size. Additionally, there were many recorded points of unidentified packs in the western and northern areas adjacent to the home ranges of P1–P4 (Fig. 2), suggesting the presence of one or two more packs other than the five identified packs in the study area. The mean home range size of the four packs was 24.45 ± 16.27 km 2 (range: 6.97–40.29 km 2 ) (Table 2). The home range size largely differed by packs; the largest home range (P4, 40.29 km 2 ) was over five times larger than the smallest home range (P2, 6.97 km 2 ). Fig. 2 also suggests that the home ranges located in the eastern part of the study area (P1 and P2) were smaller than those in the western part (P3 and P4): Specifically, the mean home range size of P1 and P2 (10.68 km 2 , range: 6.97–14.38 km 2 ) was less than half that of P3 and P4 (38.22 km 2 , range: 36.14–40.29 km 2 ) (Table 2). As P1 and P2 had more recorded points than P3 and P4 and very few recorded points of unidentified packs around their home ranges, the home range sizes of P1 and P2 were less likely to be underestimated. Even if we assume that the recorded points for the unidentified packs around the P1 and P2 home ranges are also the recorded points for each pack, the home range sizes of P1 and P2 were only 20.57 km 2 and 7.71 km 2 respectively. In contrast, the home range sizes of P3 and P4 may be underestimated as these packs had fewer recorded points than P1 and P2 and more recorded points of unidentified packs around their home ranges. The relationship between the estimated home range sizes and cumulative observation days for each pack (SI2) also supports this view. Thus, it is evident that the home ranges of P1 and P2 were remarkably smaller than those of P3 and P4. The environmental conditions of the home range differed between packs. For P1 and P2, 17.8% (12.8% residential areas, 5.0% farmlands, with less than 500 people) and 30.4% (4.6% residential areas, 25.8% farmlands, with more than 5,000 people) of their home ranges were anthropogenic environments (Table 2; Fig. 3a). On the other hand, for P3 and P4, less than 1.0% (0.6% and 0.0%, respectively, with less than 500 people) of their home ranges were in human-influenced environments (residential areas and farmlands), with most areas being covered by forests with minimal human influence (Table 2; Fig. 3b). The home ranges of P1 and P2, particularly that of P2, encompassed a greater proportion of anthropogenic environments and a higher human density than those of P3 and P4. Daily activity patterns We analysed and compared the number of video and camera trap records during day (light) and night (dark) periods (Table 4) using 227 video events (170 events from identified packs) recorded in 1,520.1 camera days and 214 photo events (19 events from identified packs) recorded in 12,884.0 camera days (Table 1b).The ratio of events (an index of activity level) indicated that daytime activity (74.4%) was significantly higher than that of nighttime activity (25.6%) ( χ 2 (1) = 104.8, p < 0.001, Table 4). The number of record locations during the day (77.9%) was also significantly higher than that at night (22.1%) ( χ 2 (1) = 37.9, p < 0.001, Table 4). These results indicate that the local population in the study area was significantly more active in the day (light period) than the night (dark period). The same analysis for each identified pack, however, revealed that one of the four identified packs (P2) showed no significant tendency to be more active during the day than at night, unlike the other packs (Table 4). Additionally, P2 had a significantly larger ratio of night records compared to P3 (adjusted P = 0.033) and P4 (adjusted P = 0.000). Analysis of the number of recorded locations revealed that all the identified packs had more locations during the day (63.2–82.6%) than at night (17.4–36.8%). However, significant differences were only detected for P3 and P4, which had larger sample sizes (Table 4). No significant differences in the activity ratio were observed between packs. Analysis of the average distances between the recorded locations (an index of activity range) revealed that all the four identified packs showed greater movement during the day (2.49 ± 2.09 km range: 2.20–3.34 km) compared to the night (0.54 ± 0.74 km, range: 0.53–1.56 km), although significant differences were detected only for P2 and P3 with larger sample sizes (Table 4). The result suggests that P2 showed more nighttime activity than other packs, but also used a smaller area in the night than in the day. The variation in the relative abundance indices (RAIs) of camera/video trap recordings with time of day (Fig. 4) also shows that the local population in the study area was mainly active during the day; peak activity occurring in the early morning around 07:00 and again in the evening around 18:00–19:00 and reduced activity from noon to 16:00, although overall, they were more active during the day than at night. The RAI peak in the morning was higher than that in the evening, suggesting greater activity in the morning. The RAI variation with time of day for the identified packs (Fig. 5) suggests that all the packs tended to be more active around sunrise and sunset, although slight differences were observed in the degree of daytime activity reduction. P1 and P2 showed higher activity in the evening compared to the morning, while P3 and P4 showed higher activity in the morning than in the evening (Fig. 5). However, the differences in RAIs between the morning and evening were not significant. Daily behavioural pattern Both active and inactive behaviours (Table 5) were recorded through video trapping showed a significantly higher frequency during the day compared to the night (active; Z = 2.041, P = 0.041; inactive; Z = -2.949, P = 0.003; Fig. 6). Active behaviours were more frequent in the morning and evening while inactive behaviours, particularly long-term resting, were most common during the day (Fig. 6). These results were consistent with the RAI data for the local population presented earlier (Fig. 4). Direct observation revealed that 79.1% of the daytime (129.3 hours out of 163.5 hours) was spent resting. It also revealed that 96.3% of the daytime rest (124.5 hours out of 129.3 hours) was long-term rest, resting by all pack members for more than 60 minutes. In most cases, long-term resting continued for more than six hours, with an average duration of 363.3 ± 246.4 minutes (range: 63.0–685.0 min, N = 39). We observed three instances of long-term resting from the beginning to the end (135.0 min; 559.0 min; 573 min). Such resting was observed on 63% (range: 50–73%) of observation days. In P1 and P2 with numerous observation events, it was noted that they frequently used specific locations (Fig. 3a; Fig. 3b) including shaded areas such as the bases of trees and underbrush on multiple occasions (5.78 ± 5.61 times, 1–15 times). We have no record of the camera/video trapping at these repeatedly-used long-term resting sites, because no instruments were installed there. Long-term rests were predominantly observed between 10:00 and 17:00 (Fig. 6), indicating that the reduced activity observed around midday, as shown by the camera/video trapping data (Fig. 4), corresponded to this resting behaviour during the midday (Fig. 6). During direct observations conducted in the daytime, a total of 19 events of hunting and feeding behaviours were recorded, along with four identified feeding signs (see Table 6 and Fig. 3a for location details). The target animals hunted or fed by the dholes were identified through the direct observation or the presence of associated signs (Table 6). These animals included four species of wild animals (spotted deer: seven instances; sambar deer: three instances; wild boars: three instances; Indian hares: two instances) and three types of livestock (cattle: two instances; goats: two instances; chicken: one instance; Table 6). Apart from wildlife and livestock predation, we observed two instances of garbage scavenging behaviour and one instance of foraging for unidentified food items on the ground. Livestock predations and garbage scavenging were only observed in P2 members (Table 6). In this region dholes primarily hunted and fed in the morning and evening. Excluding garbage scavenging, 31.3% of hunting and feeding (five out of 16 instances) were conducted in the morning and 25.0% (four out of 16 instances) were conducted in the evening. Including garbage scavenging, hunting and feeding were equally conducted in the morning and evening (31.6%, six out of 19 each). We observed scavenging three times in P2; once in the morning and twice in the evening (Table 6). In addition, we sighted P2 roaming around the garbage deposition site within a 100 m radius 23 times during dusk and dawn. This suggests that they begin scavenging during the dark period. Nighttime behaviour Regarding nighttime behaviour, we had no direct observation data and very few trap data. In addition, the large part of the data was obtained from P2; 42.5% (48 of 113 instances) and 78.7% (48 of 61 instances) of nighttime events from the local population and the identified packs were observed from P2, respectively (Table 4, Figures 5 and 7). So, it is difficult to estimate their nighttime behaviour. The reduced activity at night (Table 4) and the fact that all nighttime observation points were within the daytime ranges (Figures 2, 3a and 3b) suggest that they rested somewhere within their home range. However, exact locations used for their rest could not be identified. Interactions with humans In the two packs with high anthropogenic environment use (P1, P2), 15 incidents were observed where packs were chased away by livestock herders or villagers collecting livestock faeces (SI3). All the incidents occurred between 06:10 and 18:16, with the highest frequency observed between 07:00–08:00 (six incidents) and 16:00–17:00 (five incidents), likely coinciding with the start and end of grazing periods. We did not find any conflicts and carcasses of dholes within the home ranges of P3 and P4. Discussion Our findings on pack dynamics, reproduction, home range size, activity pattern and associated behaviours of wild dholes provide detailed insights into the ecology of this species that has been relatively understudied. In addition, we compared these parameters between four identified dhole packs with varying reproductive statuses and degrees of anthropogenic environment use to evaluate the influence of reproduction and human-modified landscapes on their behaviour. Observed group size The mean group size observed outside the vicinity of the den of wild dholes in India reported from direct observational studies show significant variation ranging from smaller averages of three individuals (Cohen et al., 1978 ) to much larger averages such as 14 to 17 individuals (Bhandari et al., 2021 ; Majumder et al., 2011 ). However, excluding these extreme cases, the majority of studies have reported group sizes between five and nine individuals (Bhandari et al., 2021 ; George et al., 2021 ; Johnsingh, 1982 ; Ramesh, 2010 ; Rice, 1986 ), consistent with the value of this study (6.3 ± 3.4 individuals). The smaller mean group size reported by Cohen et al. ( 1978 ) (3.1 ± 2.0 individuals) can be attributed to their very short study period less than two months (from October to December), coinciding with the season of pup dispersal (Johnsingh 1982 ) and beginning of the denning. Camera trap studies in India, Thailand and Malaysia also reported lower pack sizes (two to six individuals; Bashir et al., 2014 ; Jenks et al., 2012; Kawanishi & Sunquist, 2008 ; Selvan et al., 2014 ), likely due to the potential underestimation inherent in the methodology (Jenks et al., 2012). In contrast, the larger group sizes reported from central India (14 to 17 individuals: Bhandari et al., 2021 ; Majumder et al., 2011 ) could be explained by ecological factors, such as lower tiger density. Bhandari et al. ( 2021 ) demonstrated that a region with a lower tiger density (0.46 individuals / 100 km 2 ) supported larger dhole packs (16.8 ± 3.1 individuals, N = 5) compared to a region with a higher tiger density (5.36 individuals / 100 km 2 ), where group sizes averaged 6.4. ± 1.3 individuals (N = 7). This suggests that increased predation pressure and kleptoparasitism from tigers may drive smaller dhole group sizes in high-density tiger areas. The high tiger density (6.07–9.72 individuals / 100 km 2 ; Kalle et al., 2011) in our study area and comparatively lower group size (6.3 ± 3.4 individuals) observed in this study are consistent with this hypothesis. Although the results of many studies in India are consistent with this hypothesis, there is an exception. Majumder et al. ( 2011 ) reported a high mean pack size (13.9 ± 1.4 individuals) in Pench, India, where tiger density was also high (4.94 individuals / 100 km 2 ) (Karanth et al., 2004 ). This deviation indicates that other factors, such as prey availability, habitat characteristics or social dynamics, may also influence dhole group sizes. Denning season, litter sizes and timing of pack splits and/or pup dispersal The denning season (December–February) and the average litter size (seven pups, range: 6–8, N = 3, from two packs) observed in this study is consistent with the previous reports from the same study site by Venkataraman et al. ( 1995 ) and Venkataraman ( 1998 ) (December–March, 6.4 ± 1.7 pups, range: 4–10, N = 10, from two packs), as well as with those from an adjacent site (Bandipur Tiger Reserve) by Johnsingh ( 1982 ) (December–February, 8.5 pups, range: 6–8, N = 2, from one pack). Decrease in the number of adults in the groups observed outside the vicinity of the den during the denning season and video tapping records at the den suggest that one or two adults probably including the mother stayed around the den with her pups apparently on guard duty. At Bandipur, Johnsingh ( 1982 ) also reported that a mother dhole remained at the den with the pups until they were 58 days old. The timing of the decline in the adult numbers observed in the packs P1 and P2 just before their breeding (November-December) coincides with the timing of pack splits and/or dispersal of pups reported in previous studies conducted at the same region as our study (November: Johnsingh, 1982 ; December: Venkataraman et al., 1995 ). Therefore, the seasonal changes in the adult number in the observed groups can be important information to estimate the timings of their reproduction, pack splits and/or pup dispersal of the local population. Home range size The mean home range size observed in this study (24.5 ± 16.3 km 2 , range: 7.0–40.3 km 2 , N = 4, eight-month period) is notably smaller than values reported in previous studies conducted in the same and nearby areas (50.2 ± 22.0 km 2 ranging between 23.4–83.3 km 2 , N = 4, 23–37-month period) (Karanth & Sunquist, 2000 ; Johnsingh, 1982 ; Venkataraman et al., 1995 ). In particular, home range sizes as small as those of P1 (14.4 km 2 ) and P2 (7.0 km 2 ) have never been reported in the previous studies based on direct observation of the wild dholes. In contrast, On the other hand, the home range sizes of P3 (36.1 km2) and P4 (40.3 km2) fell within the home range sizes reported in the previous studies. The remarkably smaller home ranges observed for P1 and P2 compared to P3 and P4 could be the result of reproduction, as P1 and P2 were found to breed, whereas P3 and P4 most likely did not. Johnsingh ( 1982 ), who studied dholes in the same region as this study reported that the home range size of a pack decreased by approximately half (from 40 km 2 to 20km 2 ) during the breeding season. Venkataraman et al. ( 1995 ), however, observed a similar reduction even in non-breeding packs during the dry season (also the breeding season), so this reduction is unlikely to be due to breeding. Although this study did not have sufficient data to statistically compare home range sizes between the denning and non-denning seasons, the home range during the denning season tended to slightly increase in P2 compared to the non-denning season (from 3.3 km2 to 4.0 km2), whereas it drastically decreased in P1 (from 14.1 km 2 to 0.9 km 2 ) (Table 2 ). The result also suggests that factors other than reproduction caused small home ranges in P1 and P2. As the home ranges in P1 and P2 contained more anthropogenic environments than the other packs (Fig. 3 a), this may have resulted in their smaller home ranges. Home range reduction associated with anthropogenic resource use has been reported from various canids, including red foxes (Coman, Robinson & Beaumont, 1991 ; Walton et al., 2017 ), raccoon dogs (Saeki et al., 2007 ) and golden jackals (Rotem et al., 2011 ). Similarly, the remarkably small home range of P2 (7.0 km 2 ) is likely due to its dependence on anthropogenic food resources, as evidenced by observed garbage scavenging and livestock predation (Table 6 ). Venkataraman et al. ( 1995 ), who studied dholes in the same study site as ours, reported that the home range size of a pack occupying an area roughly overlapping the P2's home range (54.2km 2 ) was smaller than that of a pack occupying an area roughly overlapping the P3's home range (83.3km 2 ). They also analysed faeces and estimated that the former pack obtained 25.6% of their food by livestock predation, compared with only 5.8% for the latter pack. Their results also support the view that use of anthropogenic food resources may reduce the need for extensive foraging thereby resulted in the smaller home ranges in P1 and P2. Livestock predation is widely documented in dholes (Acharya, 2007 ; Barnett et al., 1980 ; Cohen et al., 1978 ; George et al., 2021 ; Johnsingh, 1983 ; Kumaraguru et al., 2011 ; Ramesh, 2010 & Ramesh et al., 2012b ; Selvan, et al., 2013a & 2013b ; Thinley et al., 2011 ; Venkataraman et al., 1995 ; Wang & Macdonald, 2009 ) and other canids, including gray wolves (Jenairo-Otero et al., 2020 ) and coyotes (McInturff et al., 2019). Livestock predation and garbage scavenging in artificial environments may not only reduce the necessity for extensive foraging leading to smaller home ranges but also provide sufficient nutritional intake. The fact that reproduction was observed only in P1 (once) and P2 (twice) during the study period suggests better nutritional conditions in these packs compared to P3 and P4. Venkataraman et al. ( 1995 ) and Venkataraman ( 1998 ) also reported that the pack occupying an area roughly overlapping the P2's home range bred for consecutive five years, while the pack occupying an area roughly overlapping the P3's home range did not breed every year for eight years. These facts suggest that in packs where the home range contains more anthropogenic environments, the home range becomes smaller because livestock predation and garbage scavenging can provide enough foods for breeding within a narrow home range. Daily pattern of activity and behaviour The results of this study revealed that the dholes in the study area have a diurnal activity pattern, mainly active during the light period, but most active in the early morning and the evening as they take extended rest periods around noon. It was also found that one of the four studied packs (P2) was more active at night, unlike the other packs, and showed a cathemeral activity pattern, with almost equal activity during the light and dark periods. Previous studies in various regions have also reported that the activity pattern of wild dholes is diurnal (Kamler et al., 2012 ; Karanth et al., 2017 ; Palei et al., 2021; Ramesh, 2012a; Vinitpornsawan & Fuller, 2020 ; Widodo et al., 2022 ), crepuscular (Ghaskadbi, Habib & Qureshi, 2016 ), diurnal or crepuscular (Grassman et al., 2005 ; Jenks et al., 2015 ; Nurbianto et al., 2015) or cathemeral (active both day and night) (Krishnakumar et al., 2022 ; Pudyatmoko, 2019 ). The reason why many reports describe the activity pattern of wild dholes as crepuscular is probably because they are essentially diurnal but their activity declines significantly around midday. Such activity declines around midday were also observed in 15 out of 18 previous studies that showed graphical activity data on dholes (eg. Jenks et al., 2015 ; Karanth et al., 2017 ), although most authors did not mention about it. The decline in activity around midday may reflect a behavioural adaptation to avoid the heat, as reported for other canids such as African wild dogs (Fuller, Nicholls & Kat, 1995 ), coyotes (Andelt & Andelt, 1981 ) and two species of foxes (Sunquist, 1989 ). The observations that the dhole packs in ‘long-term resting’ shifted their locations little by little, likely to follow shaded place, support this view. Interestingly, in the activity data of three studies from Bangladesh (Zakir et al., 2021 ), Bhutan (Vernes, Rajaratnam & Dorji, 2022 ) and Indonesia (Rahman et al., 2018 ), we could not identify any midday activity declines. In Bhutan, the lack of a midday activity declines could be attributed to cooler temperatures at higher altitudes. In addition, the research in Bangladesh (Zakir et al., 2021 ) and Indonesia (Rahman et al., 2018 ) were conducted in forested areas probably with more shaded places than our research area. The midday long resting behaviour similar to the ‘long-term resting’ observed in this study was also reported in maned wolves ( Chrysocyon brachyurus ) living in open and semi-open habitats, especially grasslands with scattered bushes and trees, where the average midday rest periods estimated from continuous GPS collar data ranged from 7.1 to 9.7 hours (Emmons et al., 2012 ). The timing of the hunting behaviour observed in this study (80% of the hunting events were recorded during the early morning and evening hours, Table 6 ) is consistent with the results of previous research which indicated peak hunting activity during the early morning and evening hours and discussed possible synchronisation with prey activity patterns (Johnsingh, 1982 ; Karanth & Sunquist, 2000 ). Vinitpornsawan & Fuller ( 2020 ) and Zakir et al. ( 2021 ) reported that dhole activity patterns synchronised with those of their main preys, such as sambar deer and wild boar. Pudyatmoko ( 2019 ), who studied in Indonesia also reported the synchronisation of the activity pattern between the dholes and prey animals such as rusa and barking deer. These studies suggest that, alongside midday heat avoidance, prey activity can be an important factor influencing dhole activity patterns, including midday activity decline. Although both P1 and P2 bred, but only P2 showed a significantly higher percentage of nocturnal activity (50.5%) than the other packs. The fact suggests that P2's high proportion of nocturnal activity is not the result of reproduction alone. Rather, it is likely to be related to the fact that P2 had a prominently more anthropogenic environment within its home range than the other packs and was highly dependent on anthropogenic food resources such as livestock and garbage; P2 often engaged in garbage scavenging and livestock predation (Table 6 ). Although garbage scavenging has been reported in various canid species, such as fennec foxes (Brahmi et al., 2012 ) and maned wolves (Silva & Talamoni, 2003), it has not been reported in this species. P2's dependence on anthropogenic food resources such as garbage and livestock may explain its larger rate of night activity. Since residents disposed food waste, such as chicken skins, in the morning and evening hours at a garbage dump near residential areas. Observations showed that P2 visited the area with the garbage dump in the morning and evening, or remained there throughout the night. In addition, grazing of livestock by villagers was conducted during the light hours (from around 06:00 to 18:00). All human-dhole encounters (SI3) were recorded during light hours especially in the morning between 07:00–08:00 (six incidents) and in the evening between 16:00–17:00 (five incidents), likely coinciding with the start and end of grazing periods. Hostile reactions of residents against P2 (SI3) suggest that P2 increased the activity during the dark period, shifting their activity time to the early morning (before 06:00) and/or later evening (after 18:00) to avoid possible encounters with residents. Similar increase of night activity has been reported in other canid species living in human-dominated landscapes, such as European wolves (Ciucci et al., 1997 ) and coyotes (Kitchen et al., 2000 ). Influence of anthropogenic environments on dholes Our findings indicate that dhole packs inhabiting areas with a high proportion of anthropogenic environments and frequently utilising human-associated food sources, such as livestock and garbage (P1, P2), maintained significantly smaller home ranges than those in more natural environments (P3, P4). This suggests that anthropogenic landscapes provide higher foraging efficiency, reducing the need for extensive movements. Additionally, the pack with the highest reliance on anthropogenic food (P2) exhibited a shift in its activity pattern, increasing nocturnal activity, presumably to avoid hostile encounters with humans. Packs that frequently consumed anthropogenic food (P1, P2) also exhibited a higher frequency of reproduction, likely due to improved nutritional conditions. However, they experienced higher mortality rates among both adults and pups. In contrast, no reproduction or adult deaths were recorded in the packs living in more natural environments (P3, P4) during the study period. The increased mortality in packs relying on anthropogenic resources may be linked to human-wildlife conflicts, including retaliatory killings, as well as disease transmission from domestic and feral dogs. The circumstances surrounding adult deaths in P1 and P2, along with documented human-hostile interactions (Table S1 ), suggest that some deaths resulted from illegal retaliatory killings. Retaliatory killings, such as those by poisoned bait targeting carnivores, including canids and felids, are widespread human responses to livestock predation (Sillero-Zubiri et al., 2004). Furthermore, diseases such as rabies (RABV; Rabies lyssavirus ), canine distemper (CDV; Canine morbillivirus ) and parvovirus (CPV; Carnivore protoparvovirus 1 ) and mange ( Sarcoptes scabiei var. canis ), which are often transmitted by domestic dogs (Durbin et al., 2004; Kamler et al., 2015 ), have been reported to impact dhole populations (Karanth & Sunquist, 2000 ). Packs that frequently visited garbage sites and residential areas with high feral dog populations, such as P2, were likely at greater risk of disease exposure. Notably, P2 bred twice during the study period, producing 15 pups, but none survived beyond 3–4 months. In contrast, P1, which had slightly less anthropogenic resource use, successfully raised five pups to at least 9–10 months. This suggests that disease transmission from feral or domestic dogs may have played a role in P2’s reproductive failure. Overall, while anthropogenic environments offer dholes access to high-energy food resources, leading to increased reproductive rates, these benefits are counteracted by elevated risks of human conflicts and disease. This trade-off is exemplified by the eventual collapse of P2 during the study period, whereas packs in more natural environments (P3, P4) maintained stable pack sizes with minimal disruptions. For effective long-term conservation of dholes in human-modified landscapes, it is essential to implement strategies that minimise human-wildlife conflict. Potential measures include compensation programmes for livestock losses, improved waste management to prevent dholes from scavenging in garbage sites, and controlling feral dog populations to reduce disease transmission. By addressing these challenges, coexistence between dholes and human communities can be improved, ensuring the survival of this endangered species. Declarations Supplementary information The online version contains supplementary materials available at Acknowledgements The first author had received generous support from the Masinagudi field station staff during the stay in MTR. Fieldwork would have been impossible without immense help from Trackers Shiva, Mohan and the late Bomma. We would like to express our gratitude to Dr Daisuke Muramatsu for statistical advice and to Drs Sanjeeta Sharma Pokharel and Nachiketha Sharma for proof-reading. RS was a JC Bose National Fellow during the tenure of this work. Author contributions SS conceived the ideas, designed the methodology and collected the data; SS performed analyses and SS and SK interpreted the data; SK and RS supervised the work. SS wrote the draft of the manuscript. All authors have reviewed and contributed to writing the final version of the manuscript. Funding This work was financially supported by ITP-HOPE and AS-HOPE (Primate Research Institute of Kyoto University) and JSPS Core-to-Core Program, A. Advanced Research Networks (Wildlife Research Center of Kyoto University) from the Japan Society for the Promotion of Science. Availability of data and material The datasets analysed during the current study are available from the corresponding author on reasonable request. Code availability Not applicable. Permits The Consulate General of India in Japan and the National Biodiversity Authority kindly permitted us to work in India. The Tamil Nadu Forest Department kindly granted us permission to work in MTR and provided us with its camera trap data on dholes. Ethics approval The authors confirmed that the ethic policies of the journal, as noted on the journal’s guidelines page, have been adhered to. No ethical approval was required as there was no sample collection from animals or humans. Consent to participate Not applicable. 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Supplementary Files 2025.05.15TablesS.Sawaguri.docx 2025.05.15SupplementarymaterialsS.Sawaguri.docx SI1. Photo identification of pack members of the study. SI2. Relationship between the estimated home range sizes (MCP) and cumulative observation days for each pack. SI3. List of Human-dhole interactions (human to dhole only) Cite Share Download PDF Status: Under Review Version 1 posted Editorial decision: Revision requested 05 Aug, 2025 Reviews received at journal 24 Jul, 2025 Reviews received at journal 04 Jul, 2025 Reviewers agreed at journal 17 Jun, 2025 Reviewers agreed at journal 16 Jun, 2025 Reviewers invited by journal 16 Jun, 2025 Editor assigned by journal 19 May, 2025 Submission checks completed at journal 19 May, 2025 First submitted to journal 15 May, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. 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Sawaguri","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA2UlEQVRIiWNgGAWjYBADHnn2BiBlYEGCFsOeAyAtEqTYcyMBRBKhRb79jOHHr233ZBhnPr+64UeBBAN/e3cCXi0GZ3KMpWXbinnYpXPKbvYAHSZx5uwG/FoYcjdIS7Yl8DDOzkm7wQPUYiCRi1+LfP/bzb9BWhhunkm7+YcYLQw3crdJfgRpucF+7DZRthjceP/NmuFcAjCQc9huyxhI8BD0i3x/WvLNH2UJ9vLsx5/dfPPHRo6/vZeAw4CAmQdM8RiASYLKQYDxB5hif0CU6lEwCkbBKBh5AAD/CUZLPnp4SgAAAABJRU5ErkJggg==","orcid":"","institution":"Kyoto University","correspondingAuthor":true,"prefix":"","firstName":"Shuta","middleName":"","lastName":"Sawaguri","suffix":""},{"id":472496010,"identity":"e372bcfb-8e88-4928-8e8c-40beefb7fef5","order_by":1,"name":"Raman Sukumar","email":"","orcid":"","institution":"Indian Institute of Science Bangalore","correspondingAuthor":false,"prefix":"","firstName":"Raman","middleName":"","lastName":"Sukumar","suffix":""},{"id":472496011,"identity":"d18b4862-b7b5-469e-926d-918892c30668","order_by":2,"name":"Shiro 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13:37:10","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":40662,"visible":true,"origin":"","legend":"","description":"","filename":"2025.05.15TablesS.Sawaguri.docx","url":"https://assets-eu.researchsquare.com/files/rs-6674422/v1/44e095ed46abba5059598225.docx"},{"id":84895544,"identity":"0ecf77ae-e4ba-4c24-a083-18c6fa3c1c88","added_by":"auto","created_at":"2025-06-18 13:53:11","extension":"docx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":2823578,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eSI1\u003c/strong\u003e. Photo identification of pack members of the study.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eSI2\u003c/strong\u003e. Relationship between the estimated home range sizes (MCP) and cumulative observation days for each pack.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eSI3\u003c/strong\u003e. List of Human-dhole interactions (human to dhole only)\u003c/p\u003e","description":"","filename":"2025.05.15SupplementarymaterialsS.Sawaguri.docx","url":"https://assets-eu.researchsquare.com/files/rs-6674422/v1/7ef4ce01a8e2ffe8869deccd.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Pack size, home range size and activity pattern of dhole (Cuon alpinus) in Mudumalai Tiger Reserve, India: effects of anthropogenic environment use and reproduction","fulltext":[{"header":"Introduction","content":"\u003cp\u003eThe decline of vertebrate species worldwide has been driven by a range of anthropogenic factors, including habitat destruction, fragmentation and overexploitation of natural resources. Additionally, increasing human populations have intensified conflicts with wildlife, while climate change further exacerbates biodiversity loss (Cardillo et al., \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Newbold et al., \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Ripple et al., \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). To address these challenges and conserve vertebrate biodiversity, it is essential to mitigate human-wildlife conflicts by understanding the ecology of concerned species sharing their habitats with humans and how human-modified landscapes influence their ecology and survival.\u003c/p\u003e \u003cp\u003eThe dhole (\u003cem\u003eCuon alpinus\u003c/em\u003e), also known as the Asiatic or Indian wild dog, is an endangered species that requires urgent research attention and conservation efforts. This canid is currently found in India, Southeast Asia and China. According to the International Union for Conservation of Nature (IUCN), the estimated wild population of mature dholes between 949 and 2,215, with the numbers continuing to decline due to habitat loss and transformation, retaliatory killings linked to livestock predation and other anthropogenic pressures (Kamler et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). While understanding dhole ecology is crucial for conservation, such research particularly in human-modified habitats remains insufficient.\u003c/p\u003e \u003cp\u003eAlthough fewer field studies have been conducted on dholes compared to other canid species such as wolves (Srivathsa et al., \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), several investigations in India have provided valuable insights into their ecology. These include studies on the reproduction and social structure (Johnsingh, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1982\u003c/span\u003e; Venkataraman \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e1998\u003c/span\u003e), movement patterns (Acharya, Johnsingh \u0026amp; Sankar, \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), daily activity cycles (Karanth et al. \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Krishnakumar et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Ramesh et al. 2012), habitat utilisation (Srivathsa et al., \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), population estimates using camera traps (Punjabi et al., \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), foraging ecology (Venkataraman, Arumugam \u0026amp; Sukumar \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e1995\u003c/span\u003e) and the relationship between prey-predator dynamics and pack size (Bhandari et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). However, much of these existing research rely on either unidentified packs or a very limited number of identified packs. Moreover, there has been little research on how anthropogenic environments affect dhole ecology.\u003c/p\u003e \u003cp\u003eTo address these gaps, we conducted field research in the Mudumalai Tiger Reserve, India, from September 2011 to August 2015, utilising direct observation and camera trapping. We investigated key ecological parameters, including pack size, home range, activity patterns and reproduction. Additionally, we compared four identified dhole packs with varying reproductive statuses and degrees of anthropogenic environment use to evaluate the influence of reproduction and human-modified landscapes on their behaviour. The results of this study, which provide important ecological insights into their adaptability to changing environments, could contribute to a more effective conservation strategy for this endangered species.\u003c/p\u003e"},{"header":"Methods","content":"\u003ch2\u003eStudy area\u003c/strong\u003e\u003c/h2\u003e\n\u003cp\u003eThis study was conducted in the Mudumalai Tiger Reserve (MTR) in Tamil Nadu, southern India (Fig. 1). MTR\u003csup\u003e\u0026nbsp;\u003c/sup\u003eis a part of the Nilgiri Biosphere Reserve situated in the Western Ghats, covering an area of 321 km\u003csup\u003e2\u003c/sup\u003e (11\u0026deg;32\u0026rsquo;\u0026ndash;11\u0026deg;43\u0026rsquo; N and 76\u0026deg;22\u0026rsquo;\u0026ndash;76\u0026deg;45\u0026rsquo; E), with the elevation between 800 and 1,100 m asl (Dattaraja et al., 2013). The average maximum and minimum temperatures are 26.6 \u0026plusmn; 2.34\u0026deg;C and 15.9 \u0026plusmn; 1.95\u0026deg;C, respectively (Suresh \u0026amp; Nanda, 2021). The region experiences three distinct seasons: the dry season from January to April, the southwest (summer) monsoon from June to September and the northeast (winter) monsoon from October to December (Dattaraja et al., 2013). The average annual rainfall is 1,681 mm in the western part and 721 mm in the eastern part of the reserve (Dattaraja et al., 2013). Vegetation varies by region: tropical moist deciduous forests in the west, dry deciduous forests in the central part and dry thorn forests in the east (Suresh et al., 2011, Dattaraja et al. 2018). The eastern part has more human population and human-altered environments such as agricultural fields and grazing lands than the central and western parts which adjoin coffee and tea plantations. In addition to dhole, large predators including tiger (\u003cem\u003ePanthera tigris\u003c/em\u003e), leopard (\u003cem\u003ePanthera pardus\u003c/em\u003e) and striped hyena (\u003cem\u003eHyaena hyaena\u003c/em\u003e) are seen in this area. Other larger mammals in Mudumalai include Asian elephant (\u003cem\u003eElephas maximus\u003c/em\u003e), gaur (\u003cem\u003eBos gaurus\u003c/em\u003e), chital (\u003cem\u003eAxis axis\u003c/em\u003e) and sambar (\u003cem\u003eRusa unicolor\u003c/em\u003e) deer, black buck (\u003cem\u003eAntilope cervicapra\u003c/em\u003e), wild boar (\u003cem\u003eSus scrofa\u003c/em\u003e), sloth bear (\u003cem\u003eMelursus ursinus\u003c/em\u003e), bonnet macaque (\u003cem\u003eMacaca radiata\u003c/em\u003e) and southern plains gray langur (\u003cem\u003eSemnopithecus dussumieri\u003c/em\u003e) and many smaller mammals such as several species of squirrels, mongooses and civets.\u003c/p\u003e\n\u003ch2\u003eData collection\u003c/strong\u003e\u003c/h2\u003e\n\u003ch3\u003eDirect observations\u003c/h3\u003e\n\u003cp\u003eWe conducted fieldwork to observe wild dholes directly over a period of 467 days from September 2011 to August 2015 (see Tables 1a \u0026amp; 1b).\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003eWe searched for dholes on foot or by vehicle approximately from 06:00 to 11:00 (or 06:30\u0026ndash;11:30) and from 14:00 to 18:00 (or 14:30\u0026ndash;18:30) based on recent sightings and field signs such as scats and footprints. Whenever we spotted dholes, we followed them as far as possible and observed them maintaining a distance of at least 10 m. Their behaviour was recorded using a video camera (SONY HDR-XR550V or PJ630V). We also recorded the location of each observation with a GPS device (Garmin GPSMAP 60CSx) every 15 minutes.\u003c/p\u003e\n\u003ch3\u003eVideo and camera trapping\u003c/h3\u003e\n\u003cp\u003eWe conducted video trapping over a total of 1,520 camera days from July 2014 to August 2015 (Table 1b). The video trapping was carried out using automatic recording video cameras (Bushnell 8MP Trophy Cam HD Max Black LED Trail Camera with Night Vision). We installed one or two video cameras (a total of 114 video cameras) at 37 locations: 19 junctions where dholes had recently been sighted or field signs had been found, 10 trails with scats evidence, seven ponds with footprints, and one site with a prey carcass (Fig. 1). The video cameras were securely fixed to tree trunks or other structures at 0.5\u0026ndash;1.5 m agl, angled slightly downward. They operated 24 hours a day, with each video recording set to 60 seconds and an interval of one second between captures. We collected data and replaced batteries every two weeks to one month. During these visits, we also replaced SD cards and noted the start and end times of recording. If the SD card was full, or in the event of battery depletion, the timestamp on the last recorded video was considered as the end time of the recording.\u003c/p\u003e\n\u003cp\u003eWe also used the camera trapping data collected by the Tamil Nadu Forest Department for over 12,884 camera days from November 2014 to March 2015 (Table 1b). They installed a total of 1070 camera traps (Cuddeback, Attack) at 107 locations within the study area, with two devices set up at each site (Fig. 1). Similar to our setup, these cameras were fixed to tree trunks or other structures at 0.5\u0026ndash;1.5 m agl, facing slightly downward and operated continuously 24 hours a day. We specifically used the photo data that included images of dholes for this study.\u003c/p\u003e\n\u003ch3\u003eIdentification of animals and packs\u003c/h3\u003e\n\u003cp\u003eWe identified at least one adult from each pack based on their distinct morphological features such as facial patterns and body colour (Supplementary Information SI1). If the identified animal was observed in a group, we treated the observed group as an identified pack (P1\u0026ndash;P4). Among the five packs we identified, one group had no identifiable individuals but it was treated as an identified pack (P5) because the number of adult individuals was clearly higher than in any other identified packs. All the other groups we observed were categorised as unidentified packs.\u003c/p\u003e\n\u003ch2\u003eData analysis\u003c/strong\u003e\u003c/h2\u003e\n\u003ch3\u003eDefinition of group size, pack size and pups\u003c/h3\u003e\n\u003cp\u003eGroup size was defined as the number of animals observed during each direct observation outside the vicinity of the den. Pack size, on the other hand, was defined as the number of animals in the identified group including the number of pups in the den at the time of observation. Individuals with a body length of approximately 30\u0026ndash;90 cm were considered as pups less than one year old. Small pups with a body length of about 20\u0026ndash;30 cm and a dark brown body colour were considered as pups less than one-month-old just starting to come out of the den.\u003c/p\u003e\n\u003ch3\u003eEstimation of the denning season\u003c/h3\u003e\n\u003cp\u003eThe denning season, the period when new born pups stay in or around the den, was estimated to occur from December to February based on our observation of pups in and around the dens (P1: 24 December, 2014\u0026ndash;19 February, 2015, P2: 8\u0026ndash;22 January, 2014 and 17 January\u0026ndash;15 February, 2015) and previous reports on their birthing season (November\u0026ndash;March, peaking in December; Davidar, 1975) and den usage (10\u0026ndash;11 weeks after birth, with pups peeking out at three weeks; Johnsingh, 1982).\u003c/p\u003e\n\u003ch3\u003eEstimation of home ranges\u003c/h3\u003e\n\u003cp\u003eWe plotted GPS points obtained from direct observations and from camera and video traps using QGIS, an open-source GIS software (Ver. 3.4.9, Madeira). During the direct observations, we recorded the location every 15 minutes from the initial sighting until the animals were no longer visible. We then mapped the home ranges using the minimum convex polygon (100% MCP) method.\u003c/p\u003e\n\u003ch3\u003eActivity pattern\u003c/h3\u003e\n\u003cp\u003eTo analyse the dhole activity patterns, we calculated relative abundance indices (RAIs, the number of camera capture events in each time period of the day \u0026times; 100 / total camera trap days; O\u0026rsquo;Brien et al., 2003). In our analysis of the camera trap records, all camera capture records occurring within a 30-minute window were treated as a single camera capture event.\u003c/p\u003e\n\u003cp\u003eWe classified behaviours observed through direct observation and video trapping into six behavioural categories: resting, locomotion, feeding, excretion-associated, social and others (see Table 5). In direct observation, we defined a long resting behaviour of more than one hour, carried out by all members of each pack, as \u0026apos;long resting\u0026apos;. When analysing the diurnal activity pattern, if any member of the observed pack exhibited behaviour from any above categories, we recorded it as an event for that behavioural category by that pack. We then calculated the ratios of each behavioural category during different time periods of the day (the number of events of each behavioural category / total recorded events).\u003c/p\u003e\n\u003cp\u003eIn the analysis of the diurnal activity pattern, daytime (light period) was defined as the period from sunrise to sunset, while nighttime (dark period) was defined as the period from sunset to sunrise (refer to Fig. 1b). In the graphs illustrating the diurnal activity patterns, we standardised the sunrise and sunset time to 06:30 and 18:30 hr, respectively, for convenience.\u003c/p\u003e\n\u003ch2\u003eStatistical analysis\u003c/strong\u003e\u003c/h2\u003e\n\u003cp\u003eFor the comparisons of activities between daytime and nighttime of the local population and each pack (Table 4), we applied the Chi-square test (two-sided; for multiple comparisons, with p-values corrected using the Benjamini-Hochberg method). Mann-Whitney U test was applied for the analysis of the seasonal change in average number of adults in observed group (Table 3) and in the mean distance between sighting locations (Table 4). Wilcoxon signed-rank test was applied for the comparison of active and inactive behaviours between day and night (Fig. 6). We used R Statistical Software (Ver. 3.1.1) for all analyses. The average values are represented as mean \u0026plusmn; standard deviations and range values are represented as (lowest\u0026ndash;highest).\u003c/p\u003e"},{"header":"Results","content":"\u003ch2\u003eObserved group/pack size, breeding events, pups and adults\u003c/h2\u003e\n\u003cp\u003eIn the direct observations, 467 days of observation effort resulted in 342 tracking events, totaling 492.69 hours of direct observation (Table 1a; Table 1b). A total of 55 individuals including 41 adults and 14 pups were observed during the camera trap survey period (Table 2). During the direct observation, we identified eight adults. Based on the pack sizes and the presence of the identified animals in the pack, we could identify five packs (P1\u0026ndash;P5) in the study area (Table 2).\u003c/p\u003e\n\u003cp\u003eThe mean group size of the local population observed outside the vicinity of the dens in the study area including unidentified packs was 6.3 \u0026plusmn; 3.4 individuals (range: 1\u0026ndash;16, N = 268) with an average of 5.7 \u0026plusmn; 3.0 adults (range: 1\u0026ndash;16, N = 268) and 0.7 \u0026plusmn; 2.1 pups (range: 0\u0026ndash;6, N = 268). The mean group size of the five identified packs was 6.9 \u0026plusmn; 3.4 individuals (range: 1\u0026ndash;16, N = 262) with an average of 5.8 \u0026plusmn; 2.9 adults (range: 1\u0026ndash;16, N = 262) and 0.8 \u0026plusmn; 2.5 pups (range: 0\u0026ndash;6, N = 262).\u003c/p\u003e\n\u003cp\u003eThree breeding events in packs P1 (one event) and P2 (two events) were confirmed (Table 2). A total of 21 pups were observed including six pups in P1 on 24 December, 2014, seven of P2 on 8 January, 2014 and eight of P2 on 22 January, 2015. The average litter size was estimated to be seven (range: 6\u0026ndash;8). In P1, where six newborn pups were born in December 2014, five pups (aged 9\u0026ndash;10 months) that had grown to near adult size (third-forth) were observed joining adults by 14 August, 2015. Thus, it is likely that P1 had successfully bred. In contrast, P2 had seven and eight newborn pups in January 2014 and 2015, respectively, but only four and six pups (aged 3-4 months) were observed in March (22th in 2014 and 18th in 2015), respectively. As there have been no observations since then to confirm the survival of these pups, it is likely that P2 failed to raise any pups to adulthood in both 2014 and 2015. In P3 and P4, no pups or immature individuals with the body size remarkably smaller than adults were observed, suggesting that these packs did not breed or failed to breed at the early stage of their reproduction during the study period.\u003c/p\u003e\n\u003cp\u003eIn P1 and P2, the average number of adults in the groups observed outside the vicinity of the den decreased during the denning season (December\u0026ndash;February) (Table 3). This value was also significantly reduced in the local population of the study area, including other identified and unidentified packs. These results suggest that one or two adults stayed around the den with their pups during the denning season. During the denning season in 2014 and 2015, the video trapping cameras at P2\u0026apos;s den frequently recorded a female (probably the mother) and a male staying around the den, apparently on guard duty.\u003c/p\u003e\n\u003cp\u003eThe number of adults in P1 and P2 decreased during the study period as a result of deaths of unknown causes (Table 2). In P1, the carcass of an adult member vomiting a large amount of blood from the mouth was found beside a dead wild boar (k1 in Fig. 3a) on 24 December, 2014. Boar meat fragments were found on the ground and no external injuries were observed in the dead body. Subsequently, a reduction in the number of adults in P1 was confirmed on the same day. Although we could not observe directly, we were also informed by local people that carcasses of other carnivorous animals such as striped hyenas and vultures were found near the spot where we found the dead dhole during the same period. In P2, we found the carcass of three identified male adults without any external injuries in a small area within 20\u0026ndash;30 m\u003csup\u003e2\u003c/sup\u003e (k2 in Fig. 3a) on 25 July, 2015. They also vomited blood from their mouths. After this event, we could never observe P2. Both sites where we found the carcasses were close to villages (within 50 m from each village). These facts and circumstances suggest that the cause of their deaths may have been unnatural ones such as artificial poisoning.\u003c/p\u003e\n\u003cp\u003eIn P1 and P2, reductions in the number of adults were also observed, possibly due to causes other than those mentioned above. In P1, the number of adults decreased from 11\u0026ndash;12 to 4\u0026ndash;5 on 23 November, 2014 before their breeding in 2014 (24 December, 2014). In P2, it decreased from 9 to 4\u0026ndash;6 on 7 December, 2013 before their breeding in 2014 (8 January, 2014). However, such reduction in adult number was not observed before the breeding of P2 in 2015 (22 January).\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;In contrast, no reduction in the number of adults for any reasons was observed in P3 and P4.\u003c/p\u003e\n\u003ch2\u003eHome ranges\u003c/h2\u003e\n\u003cp\u003eWe mapped the home ranges of four identified packs (P1\u0026ndash;P4) (Fig. 2) based on a total of 1,247 recorded location points in total: 806 points from direct observation (Table 2), 227 points from video capture (Table 1b) and 214 points from photo capture (Table 1b). For P5, however, we could not obtain enough location points to determine its home range or estimate its size. Additionally, there were many recorded points of unidentified packs in the western and northern areas adjacent to the home ranges of P1\u0026ndash;P4 (Fig. 2), suggesting the presence of one or two more packs other than the five identified packs in the study area.\u003c/p\u003e\n\u003cp\u003eThe mean home range size of the four packs was 24.45 \u0026plusmn; 16.27 km\u003csup\u003e2\u0026nbsp;\u003c/sup\u003e(range: 6.97\u0026ndash;40.29 km\u003csup\u003e2\u003c/sup\u003e) (Table 2). The home range size largely differed by packs; the largest home range (P4, 40.29 km\u003csup\u003e2\u003c/sup\u003e) was over five times larger than the smallest home range (P2, 6.97 km\u003csup\u003e2\u003c/sup\u003e). Fig. 2 also suggests that the home ranges located in the eastern part of the study area (P1 and P2) were smaller than those in the western part (P3 and P4): Specifically, the mean home range size of P1 and P2 (10.68 km\u003csup\u003e2\u003c/sup\u003e, range: 6.97\u0026ndash;14.38 km\u003csup\u003e2\u003c/sup\u003e) was less than half that of P3 and P4 (38.22 km\u003csup\u003e2\u003c/sup\u003e, range: 36.14\u0026ndash;40.29 km\u003csup\u003e2\u003c/sup\u003e) (Table 2).\u003c/p\u003e\n\u003cp\u003eAs P1 and P2 had more recorded points than P3 and P4 and very few recorded points of unidentified packs around their home ranges, the home range sizes of P1 and P2 were less likely to be underestimated. Even if we assume that the recorded points for the unidentified packs around the P1 and P2 home ranges are also the recorded points for each pack, the home range sizes of P1 and P2 were only 20.57 km\u003csup\u003e2\u003c/sup\u003e and 7.71 km\u003csup\u003e2\u003c/sup\u003e respectively. In contrast, the home range sizes of P3 and P4 may be underestimated as these packs had fewer recorded points than P1 and P2 and more recorded points of unidentified packs around their home ranges. The relationship between the estimated home range sizes and cumulative observation days for each pack (SI2) also supports this view. Thus, it is evident that the home ranges of P1 and P2 were remarkably smaller than those of P3 and P4.\u003c/p\u003e\n\u003cp\u003eThe environmental conditions of the home range differed between packs. For P1 and P2, 17.8% (12.8% residential areas, 5.0% farmlands, with less than 500 people) and 30.4% (4.6% residential areas, 25.8% farmlands, with more than 5,000 people) of their home ranges were anthropogenic environments (Table 2; Fig. 3a). On the other hand, for P3 and P4, less than 1.0% (0.6% and 0.0%, respectively, with less than 500 people) of their home ranges were in human-influenced environments (residential areas and farmlands), with most areas being covered by forests with minimal human influence (Table 2; Fig. 3b). The home ranges of P1 and P2, particularly that of P2, encompassed a greater proportion of anthropogenic environments and a higher human density than those of P3 and P4.\u0026nbsp;\u003c/p\u003e\n\u003ch2\u003eDaily activity patterns\u003c/h2\u003e\n\u003cp\u003eWe analysed and compared the number of video and camera trap records during day (light) and night (dark) periods (Table 4) using 227 video events (170 events from identified packs) recorded in 1,520.1 camera days and 214 photo events (19 events from identified packs) recorded in 12,884.0 camera days (Table 1b).The ratio of events (an index of activity level) indicated that daytime activity (74.4%) was significantly higher than that of nighttime activity (25.6%) (\u003cem\u003e\u0026chi;\u003c/em\u003e\u003csup\u003e2\u0026nbsp;\u003c/sup\u003e(1) = 104.8, p \u0026lt; 0.001, Table 4). The number of record locations during the day (77.9%) was also significantly higher than that at night (22.1%) (\u003cem\u003e\u0026chi;\u003c/em\u003e\u003csup\u003e2\u0026nbsp;\u003c/sup\u003e(1) = 37.9, p \u0026lt; 0.001, Table 4). These results indicate that the local population in the study area was significantly more active in the day (light period) than the night (dark period).\u003c/p\u003e\n\u003cp\u003eThe same analysis for each identified pack, however, revealed that one of the four identified packs (P2) showed no significant tendency to be more active during the day than at night, unlike the other packs (Table 4). Additionally, P2 had a significantly larger ratio of night records compared to P3 (adjusted P = 0.033) and P4 (adjusted P = 0.000). Analysis of the number of recorded locations revealed that all the identified packs had more locations during the day (63.2\u0026ndash;82.6%) than at night (17.4\u0026ndash;36.8%). However, significant differences were only detected for P3 and P4, which had larger sample sizes (Table 4). No significant differences in the activity ratio were observed between packs.\u003c/p\u003e\n\u003cp\u003eAnalysis of the average distances between the recorded locations (an index of activity range) revealed that all the four identified packs showed greater movement during the day (2.49 \u0026plusmn; 2.09 km range: 2.20\u0026ndash;3.34 km) compared to the night (0.54 \u0026plusmn; 0.74 km, range: 0.53\u0026ndash;1.56 km), although significant differences were detected only for P2 and P3 with larger sample sizes (Table 4). The result suggests that P2 showed more nighttime activity than other packs, but also used a smaller area in the night than in the day.\u003c/p\u003e\n\u003cp\u003eThe variation in the relative abundance indices (RAIs) of camera/video trap recordings with time of day (Fig. 4) also shows that the local population in the study area was mainly active during the day; peak activity occurring in the early morning around 07:00 and again in the evening around 18:00\u0026ndash;19:00 and reduced activity from noon to 16:00, although overall, they were more active during the day than at night. The RAI peak in the morning was higher than that in the evening, suggesting greater activity in the morning.\u003c/p\u003e\n\u003cp\u003eThe RAI variation with time of day for the identified packs (Fig. 5) suggests that all the packs tended to be more active around sunrise and sunset, although slight differences were observed in the degree of daytime activity reduction. P1 and P2 showed higher activity in the evening compared to the morning, while P3 and P4 showed higher activity in the morning than in the evening (Fig. 5). However, the differences in RAIs between the morning and evening were not significant.\u003c/p\u003e\n\u003ch2\u003eDaily behavioural pattern\u003c/h2\u003e\n\u003cp\u003eBoth active and inactive behaviours (Table 5) were recorded through video trapping showed a significantly higher frequency during the day compared to the night (active; \u003cem\u003eZ\u003c/em\u003e = 2.041, \u003cem\u003eP\u003c/em\u003e = 0.041; inactive; \u003cem\u003eZ\u003c/em\u003e = -2.949, \u003cem\u003eP\u003c/em\u003e = 0.003; Fig. 6). Active behaviours were more frequent in the morning and evening while inactive behaviours, particularly long-term resting, were most common during the day (Fig. 6). These results were consistent with the RAI data for the local population presented earlier (Fig. 4).\u003c/p\u003e\n\u003cp\u003eDirect observation revealed that 79.1% of the daytime (129.3 hours out of 163.5 hours) was spent resting. It also revealed that\u0026nbsp;96.3% of the daytime rest (124.5 hours out of 129.3 hours) was long-term rest, resting by all pack members for more than 60 minutes. In most cases, long-term resting continued for more than six hours, with an average duration of 363.3 \u0026plusmn; 246.4 minutes (range: 63.0\u0026ndash;685.0 min, N = 39). We observed three instances of long-term resting from the beginning to the end (135.0 min; 559.0 min; 573 min). Such resting was observed on 63% (range: 50\u0026ndash;73%) of observation days. In P1 and P2 with numerous observation events, it was noted that they frequently used specific locations (Fig. 3a; Fig. 3b) including shaded areas such as the bases of trees and underbrush on multiple occasions (5.78 \u0026plusmn; 5.61 times, 1\u0026ndash;15 times). We have no record of the camera/video trapping at these repeatedly-used long-term resting sites, because no instruments were installed there. Long-term rests were predominantly observed between 10:00 and 17:00 (Fig. 6), indicating that the reduced activity observed around midday, as shown by the camera/video trapping data (Fig. 4), corresponded to this resting behaviour during the midday (Fig. 6).\u003c/p\u003e\n\u003cp\u003eDuring direct observations conducted in the daytime, a total of 19 events of hunting and feeding behaviours were recorded, along with four identified feeding signs (see Table 6 and Fig. 3a for location details). The target animals hunted or fed by the dholes were identified through the direct observation or the presence of associated signs (Table 6). These animals included four species of wild animals (spotted deer: seven instances; sambar deer: three instances; wild boars: three instances; Indian hares: two instances) and three types of livestock (cattle: two instances; goats: two instances; chicken: one instance; Table 6). Apart from wildlife and livestock predation, we observed two instances of garbage scavenging behaviour and one instance of foraging for unidentified food items on the ground. Livestock predations and garbage scavenging were only observed in P2 members (Table 6).\u003c/p\u003e\n\u003cp\u003eIn this region dholes primarily hunted and fed in the morning and evening. Excluding garbage scavenging, 31.3% of hunting and feeding (five out of 16 instances) were conducted in the morning and 25.0% (four out of 16 instances) were conducted in the evening. Including garbage scavenging, hunting and feeding were equally conducted in the morning and evening (31.6%, six out of 19 each). We observed scavenging three times in P2; once in the morning and twice in the evening (Table 6). In addition, we sighted P2 roaming around the garbage deposition site within a 100 m radius 23 times during dusk and dawn. This suggests that they begin scavenging during the dark period.\u003c/p\u003e\n\u003ch3\u003eNighttime behaviour\u003c/h3\u003e\n\u003cp\u003eRegarding nighttime behaviour, we had no direct observation data and very few trap data. In addition, the large part of the data was obtained from P2; 42.5% (48 of 113 instances) and 78.7% (48 of 61 instances) of nighttime events from the local population and the identified packs were observed from P2, respectively (Table 4, Figures 5 and 7). So, it is difficult to estimate their nighttime behaviour. The reduced activity at night (Table 4) and the fact that all nighttime observation points were within the daytime ranges (Figures 2, 3a and 3b) suggest that they rested somewhere within their home range. However, exact locations used for their rest could not be identified.\u0026nbsp;\u003c/p\u003e\n\u003ch2\u003eInteractions with humans\u003c/h2\u003e\n\u003cp\u003eIn the two packs with high anthropogenic environment use (P1, P2), 15 incidents were observed where packs were chased away by livestock herders or villagers collecting livestock faeces (SI3). All the incidents occurred between 06:10 and 18:16, with the highest frequency observed between 07:00\u0026ndash;08:00 (six incidents) and 16:00\u0026ndash;17:00 (five incidents), likely coinciding with the start and end of grazing periods. We did not find any conflicts and carcasses of dholes within the home ranges of P3 and P4.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eOur findings on pack dynamics, reproduction, home range size, activity pattern and associated behaviours of wild dholes provide detailed insights into the ecology of this species that has been relatively understudied. In addition, we compared these parameters between four identified dhole packs with varying reproductive statuses and degrees of anthropogenic environment use to evaluate the influence of reproduction and human-modified landscapes on their behaviour.\u003c/p\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003eObserved group size\u003c/h2\u003e \u003cp\u003eThe mean group size observed outside the vicinity of the den of wild dholes in India reported from direct observational studies show significant variation ranging from smaller averages of three individuals (Cohen et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e1978\u003c/span\u003e) to much larger averages such as 14 to 17 individuals (Bhandari et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Majumder et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). However, excluding these extreme cases, the majority of studies have reported group sizes between five and nine individuals (Bhandari et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; George et al., \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Johnsingh, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1982\u003c/span\u003e; Ramesh, \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Rice, \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e1986\u003c/span\u003e), consistent with the value of this study (6.3\u0026thinsp;\u0026plusmn;\u0026thinsp;3.4 individuals).\u003c/p\u003e \u003cp\u003eThe smaller mean group size reported by Cohen et al. (\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e1978\u003c/span\u003e) (3.1\u0026thinsp;\u0026plusmn;\u0026thinsp;2.0 individuals) can be attributed to their very short study period less than two months (from October to December), coinciding with the season of pup dispersal (Johnsingh \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1982\u003c/span\u003e) and beginning of the denning. Camera trap studies in India, Thailand and Malaysia also reported lower pack sizes (two to six individuals; Bashir et al., \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Jenks et al., 2012; Kawanishi \u0026amp; Sunquist, \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Selvan et al., \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), likely due to the potential underestimation inherent in the methodology (Jenks et al., 2012).\u003c/p\u003e \u003cp\u003eIn contrast, the larger group sizes reported from central India (14 to 17 individuals: Bhandari et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Majumder et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) could be explained by ecological factors, such as lower tiger density. Bhandari et al. (\u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) demonstrated that a region with a lower tiger density (0.46 individuals / 100 km\u003csup\u003e2\u003c/sup\u003e) supported larger dhole packs (16.8\u0026thinsp;\u0026plusmn;\u0026thinsp;3.1 individuals, N\u0026thinsp;=\u0026thinsp;5) compared to a region with a higher tiger density (5.36 individuals / 100 km\u003csup\u003e2\u003c/sup\u003e), where group sizes averaged 6.4. \u0026plusmn; 1.3 individuals (N\u0026thinsp;=\u0026thinsp;7). This suggests that increased predation pressure and kleptoparasitism from tigers may drive smaller dhole group sizes in high-density tiger areas. The high tiger density (6.07\u0026ndash;9.72 individuals / 100 km\u003csup\u003e2\u003c/sup\u003e; Kalle et al., 2011) in our study area and comparatively lower group size (6.3\u0026thinsp;\u0026plusmn;\u0026thinsp;3.4 individuals) observed in this study are consistent with this hypothesis.\u003c/p\u003e \u003cp\u003eAlthough the results of many studies in India are consistent with this hypothesis, there is an exception. Majumder et al. (\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) reported a high mean pack size (13.9\u0026thinsp;\u0026plusmn;\u0026thinsp;1.4 individuals) in Pench, India, where tiger density was also high (4.94 individuals / 100 km\u003csup\u003e2\u003c/sup\u003e) (Karanth et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). This deviation indicates that other factors, such as prey availability, habitat characteristics or social dynamics, may also influence dhole group sizes.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec16\" class=\"Section2\"\u003e \u003ch2\u003eDenning season, litter sizes and timing of pack splits and/or pup dispersal\u003c/h2\u003e \u003cp\u003eThe denning season (December\u0026ndash;February) and the average litter size (seven pups, range: 6\u0026ndash;8, N\u0026thinsp;=\u0026thinsp;3, from two packs) observed in this study is consistent with the previous reports from the same study site by Venkataraman et al. (\u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e1995\u003c/span\u003e) and Venkataraman (\u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e1998\u003c/span\u003e) (December\u0026ndash;March, 6.4\u0026thinsp;\u0026plusmn;\u0026thinsp;1.7 pups, range: 4\u0026ndash;10, N\u0026thinsp;=\u0026thinsp;10, from two packs), as well as with those from an adjacent site (Bandipur Tiger Reserve) by Johnsingh (\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1982\u003c/span\u003e) (December\u0026ndash;February, 8.5 pups, range: 6\u0026ndash;8, N\u0026thinsp;=\u0026thinsp;2, from one pack).\u003c/p\u003e \u003cp\u003eDecrease in the number of adults in the groups observed outside the vicinity of the den during the denning season and video tapping records at the den suggest that one or two adults probably including the mother stayed around the den with her pups apparently on guard duty. At Bandipur, Johnsingh (\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1982\u003c/span\u003e) also reported that a mother dhole remained at the den with the pups until they were 58 days old.\u003c/p\u003e \u003cp\u003eThe timing of the decline in the adult numbers observed in the packs P1 and P2 just before their breeding (November-December) coincides with the timing of pack splits and/or dispersal of pups reported in previous studies conducted at the same region as our study (November: Johnsingh, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1982\u003c/span\u003e; December: Venkataraman et al., \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e1995\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eTherefore, the seasonal changes in the adult number in the observed groups can be important information to estimate the timings of their reproduction, pack splits and/or pup dispersal of the local population.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec17\" class=\"Section2\"\u003e \u003ch2\u003eHome range size\u003c/h2\u003e \u003cp\u003eThe mean home range size observed in this study (24.5\u0026thinsp;\u0026plusmn;\u0026thinsp;16.3 km\u003csup\u003e2\u003c/sup\u003e, range: 7.0\u0026ndash;40.3 km\u003csup\u003e2\u003c/sup\u003e, N\u0026thinsp;=\u0026thinsp;4, eight-month period) is notably smaller than values reported in previous studies conducted in the same and nearby areas (50.2\u0026thinsp;\u0026plusmn;\u0026thinsp;22.0 km\u003csup\u003e2\u003c/sup\u003e ranging between 23.4\u0026ndash;83.3 km\u003csup\u003e2\u003c/sup\u003e, N\u0026thinsp;=\u0026thinsp;4, 23\u0026ndash;37-month period) (Karanth \u0026amp; Sunquist, \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Johnsingh, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1982\u003c/span\u003e; Venkataraman et al., \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e1995\u003c/span\u003e). In particular, home range sizes as small as those of P1 (14.4 km\u003csup\u003e2\u003c/sup\u003e) and P2 (7.0 km\u003csup\u003e2\u003c/sup\u003e) have never been reported in the previous studies based on direct observation of the wild dholes. In contrast, On the other hand, the home range sizes of P3 (36.1 km2) and P4 (40.3 km2) fell within the home range sizes reported in the previous studies.\u003c/p\u003e \u003cp\u003eThe remarkably smaller home ranges observed for P1 and P2 compared to P3 and P4 could be the result of reproduction, as P1 and P2 were found to breed, whereas P3 and P4 most likely did not. Johnsingh (\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1982\u003c/span\u003e), who studied dholes in the same region as this study reported that the home range size of a pack decreased by approximately half (from 40 km\u003csup\u003e2\u003c/sup\u003e to 20km\u003csup\u003e2\u003c/sup\u003e) during the breeding season. Venkataraman et al. (\u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e1995\u003c/span\u003e), however, observed a similar reduction even in non-breeding packs during the dry season (also the breeding season), so this reduction is unlikely to be due to breeding. Although this study did not have sufficient data to statistically compare home range sizes between the denning and non-denning seasons, the home range during the denning season tended to slightly increase in P2 compared to the non-denning season (from 3.3 km2 to 4.0 km2), whereas it drastically decreased in P1 (from 14.1 km\u003csup\u003e2\u003c/sup\u003e to 0.9 km\u003csup\u003e2\u003c/sup\u003e) (Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The result also suggests that factors other than reproduction caused small home ranges in P1 and P2.\u003c/p\u003e \u003cp\u003eAs the home ranges in P1 and P2 contained more anthropogenic environments than the other packs (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003ea), this may have resulted in their smaller home ranges. Home range reduction associated with anthropogenic resource use has been reported from various canids, including red foxes (Coman, Robinson \u0026amp; Beaumont, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e1991\u003c/span\u003e; Walton et al., \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), raccoon dogs (Saeki et al., \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2007\u003c/span\u003e) and golden jackals (Rotem et al., \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Similarly, the remarkably small home range of P2 (7.0 km\u003csup\u003e2\u003c/sup\u003e) is likely due to its dependence on anthropogenic food resources, as evidenced by observed garbage scavenging and livestock predation (Table\u0026nbsp;\u003cspan refid=\"Tab6\" class=\"InternalRef\"\u003e6\u003c/span\u003e). Venkataraman et al. (\u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e1995\u003c/span\u003e), who studied dholes in the same study site as ours, reported that the home range size of a pack occupying an area roughly overlapping the P2's home range (54.2km\u003csup\u003e2\u003c/sup\u003e) was smaller than that of a pack occupying an area roughly overlapping the P3's home range (83.3km\u003csup\u003e2\u003c/sup\u003e). They also analysed faeces and estimated that the former pack obtained 25.6% of their food by livestock predation, compared with only 5.8% for the latter pack. Their results also support the view that use of anthropogenic food resources may reduce the need for extensive foraging thereby resulted in the smaller home ranges in P1 and P2.\u003c/p\u003e \u003cp\u003eLivestock predation is widely documented in dholes (Acharya, \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Barnett et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1980\u003c/span\u003e; Cohen et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e1978\u003c/span\u003e; George et al., \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Johnsingh, \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e1983\u003c/span\u003e; Kumaraguru et al., \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Ramesh, \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2010\u003c/span\u003e \u0026amp; Ramesh et al., \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2012b\u003c/span\u003e; Selvan, et al., \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2013a\u003c/span\u003e \u0026amp; \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2013b\u003c/span\u003e; Thinley et al., \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Venkataraman et al., \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e1995\u003c/span\u003e; Wang \u0026amp; Macdonald, \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2009\u003c/span\u003e) and other canids, including gray wolves (Jenairo-Otero et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) and coyotes (McInturff et al., 2019). Livestock predation and garbage scavenging in artificial environments may not only reduce the necessity for extensive foraging leading to smaller home ranges but also provide sufficient nutritional intake. The fact that reproduction was observed only in P1 (once) and P2 (twice) during the study period suggests better nutritional conditions in these packs compared to P3 and P4. Venkataraman et al. (\u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e1995\u003c/span\u003e) and Venkataraman (\u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e1998\u003c/span\u003e) also reported that the pack occupying an area roughly overlapping the P2's home range bred for consecutive five years, while the pack occupying an area roughly overlapping the P3's home range did not breed every year for eight years. These facts suggest that in packs where the home range contains more anthropogenic environments, the home range becomes smaller because livestock predation and garbage scavenging can provide enough foods for breeding within a narrow home range.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec18\" class=\"Section2\"\u003e \u003ch2\u003eDaily pattern of activity and behaviour\u003c/h2\u003e \u003cp\u003eThe results of this study revealed that the dholes in the study area have a diurnal activity pattern, mainly active during the light period, but most active in the early morning and the evening as they take extended rest periods around noon. It was also found that one of the four studied packs (P2) was more active at night, unlike the other packs, and showed a cathemeral activity pattern, with almost equal activity during the light and dark periods.\u003c/p\u003e \u003cp\u003ePrevious studies in various regions have also reported that the activity pattern of wild dholes is diurnal (Kamler et al., \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Karanth et al., \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Palei et al., 2021; Ramesh, 2012a; Vinitpornsawan \u0026amp; Fuller, \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Widodo et al., \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), crepuscular (Ghaskadbi, Habib \u0026amp; Qureshi, \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), diurnal or crepuscular (Grassman et al., \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Jenks et al., \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Nurbianto et al., 2015) or cathemeral (active both day and night) (Krishnakumar et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Pudyatmoko, \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe reason why many reports describe the activity pattern of wild dholes as crepuscular is probably because they are essentially diurnal but their activity declines significantly around midday. Such activity declines around midday were also observed in 15 out of 18 previous studies that showed graphical activity data on dholes (eg. Jenks et al., \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Karanth et al., \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), although most authors did not mention about it.\u003c/p\u003e \u003cp\u003eThe decline in activity around midday may reflect a behavioural adaptation to avoid the heat, as reported for other canids such as African wild dogs (Fuller, Nicholls \u0026amp; Kat, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e1995\u003c/span\u003e), coyotes (Andelt \u0026amp; Andelt, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e1981\u003c/span\u003e) and two species of foxes (Sunquist, \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e1989\u003c/span\u003e). The observations that the dhole packs in \u0026lsquo;long-term resting\u0026rsquo; shifted their locations little by little, likely to follow shaded place, support this view. Interestingly, in the activity data of three studies from Bangladesh (Zakir et al., \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), Bhutan (Vernes, Rajaratnam \u0026amp; Dorji, \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) and Indonesia (Rahman et al., \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), we could not identify any midday activity declines. In Bhutan, the lack of a midday activity declines could be attributed to cooler temperatures at higher altitudes. In addition, the research in Bangladesh (Zakir et al., \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) and Indonesia (Rahman et al., \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) were conducted in forested areas probably with more shaded places than our research area.\u003c/p\u003e \u003cp\u003eThe midday long resting behaviour similar to the \u0026lsquo;long-term resting\u0026rsquo; observed in this study was also reported in maned wolves (\u003cem\u003eChrysocyon brachyurus\u003c/em\u003e) living in open and semi-open habitats, especially grasslands with scattered bushes and trees, where the average midday rest periods estimated from continuous GPS collar data ranged from 7.1 to 9.7 hours (Emmons et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2012\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe timing of the hunting behaviour observed in this study (80% of the hunting events were recorded during the early morning and evening hours, Table\u0026nbsp;\u003cspan refid=\"Tab6\" class=\"InternalRef\"\u003e6\u003c/span\u003e) is consistent with the results of previous research which indicated peak hunting activity during the early morning and evening hours and discussed possible synchronisation with prey activity patterns (Johnsingh, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e1982\u003c/span\u003e; Karanth \u0026amp; Sunquist, \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). Vinitpornsawan \u0026amp; Fuller (\u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) and Zakir et al. (\u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) reported that dhole activity patterns synchronised with those of their main preys, such as sambar deer and wild boar. Pudyatmoko (\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2019\u003c/span\u003e), who studied in Indonesia also reported the synchronisation of the activity pattern between the dholes and prey animals such as rusa and barking deer. These studies suggest that, alongside midday heat avoidance, prey activity can be an important factor influencing dhole activity patterns, including midday activity decline.\u003c/p\u003e \u003cp\u003eAlthough both P1 and P2 bred, but only P2 showed a significantly higher percentage of nocturnal activity (50.5%) than the other packs. The fact suggests that P2's high proportion of nocturnal activity is not the result of reproduction alone. Rather, it is likely to be related to the fact that P2 had a prominently more anthropogenic environment within its home range than the other packs and was highly dependent on anthropogenic food resources such as livestock and garbage; P2 often engaged in garbage scavenging and livestock predation (Table\u0026nbsp;\u003cspan refid=\"Tab6\" class=\"InternalRef\"\u003e6\u003c/span\u003e). Although garbage scavenging has been reported in various canid species, such as fennec foxes (Brahmi et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2012\u003c/span\u003e) and maned wolves (Silva \u0026amp; Talamoni, 2003), it has not been reported in this species.\u003c/p\u003e \u003cp\u003eP2's dependence on anthropogenic food resources such as garbage and livestock may explain its larger rate of night activity. Since residents disposed food waste, such as chicken skins, in the morning and evening hours at a garbage dump near residential areas. Observations showed that P2 visited the area with the garbage dump in the morning and evening, or remained there throughout the night. In addition, grazing of livestock by villagers was conducted during the light hours (from around 06:00 to 18:00). All human-dhole encounters (SI3) were recorded during light hours especially in the morning between 07:00\u0026ndash;08:00 (six incidents) and in the evening between 16:00\u0026ndash;17:00 (five incidents), likely coinciding with the start and end of grazing periods. Hostile reactions of residents against P2 (SI3) suggest that P2 increased the activity during the dark period, shifting their activity time to the early morning (before 06:00) and/or later evening (after 18:00) to avoid possible encounters with residents. Similar increase of night activity has been reported in other canid species living in human-dominated landscapes, such as European wolves (Ciucci et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e1997\u003c/span\u003e) and coyotes (Kitchen et al., \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2000\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec19\" class=\"Section2\"\u003e \u003ch2\u003eInfluence of anthropogenic environments on dholes\u003c/h2\u003e \u003cp\u003eOur findings indicate that dhole packs inhabiting areas with a high proportion of anthropogenic environments and frequently utilising human-associated food sources, such as livestock and garbage (P1, P2), maintained significantly smaller home ranges than those in more natural environments (P3, P4). This suggests that anthropogenic landscapes provide higher foraging efficiency, reducing the need for extensive movements. Additionally, the pack with the highest reliance on anthropogenic food (P2) exhibited a shift in its activity pattern, increasing nocturnal activity, presumably to avoid hostile encounters with humans.\u003c/p\u003e \u003cp\u003ePacks that frequently consumed anthropogenic food (P1, P2) also exhibited a higher frequency of reproduction, likely due to improved nutritional conditions. However, they experienced higher mortality rates among both adults and pups. In contrast, no reproduction or adult deaths were recorded in the packs living in more natural environments (P3, P4) during the study period.\u003c/p\u003e \u003cp\u003eThe increased mortality in packs relying on anthropogenic resources may be linked to human-wildlife conflicts, including retaliatory killings, as well as disease transmission from domestic and feral dogs. The circumstances surrounding adult deaths in P1 and P2, along with documented human-hostile interactions (Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e), suggest that some deaths resulted from illegal retaliatory killings. Retaliatory killings, such as those by poisoned bait targeting carnivores, including canids and felids, are widespread human responses to livestock predation (Sillero-Zubiri et al., 2004).\u003c/p\u003e \u003cp\u003eFurthermore, diseases such as rabies (RABV; \u003cem\u003eRabies lyssavirus\u003c/em\u003e), canine distemper (CDV; \u003cem\u003eCanine morbillivirus\u003c/em\u003e) and parvovirus (CPV; \u003cem\u003eCarnivore protoparvovirus 1\u003c/em\u003e) and mange (\u003cem\u003eSarcoptes scabiei var. canis\u003c/em\u003e), which are often transmitted by domestic dogs (Durbin et al., 2004; Kamler et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2015\u003c/span\u003e), have been reported to impact dhole populations (Karanth \u0026amp; Sunquist, \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). Packs that frequently visited garbage sites and residential areas with high feral dog populations, such as P2, were likely at greater risk of disease exposure. Notably, P2 bred twice during the study period, producing 15 pups, but none survived beyond 3\u0026ndash;4 months. In contrast, P1, which had slightly less anthropogenic resource use, successfully raised five pups to at least 9\u0026ndash;10 months. This suggests that disease transmission from feral or domestic dogs may have played a role in P2\u0026rsquo;s reproductive failure.\u003c/p\u003e \u003cp\u003eOverall, while anthropogenic environments offer dholes access to high-energy food resources, leading to increased reproductive rates, these benefits are counteracted by elevated risks of human conflicts and disease. This trade-off is exemplified by the eventual collapse of P2 during the study period, whereas packs in more natural environments (P3, P4) maintained stable pack sizes with minimal disruptions.\u003c/p\u003e \u003cp\u003eFor effective long-term conservation of dholes in human-modified landscapes, it is essential to implement strategies that minimise human-wildlife conflict. Potential measures include compensation programmes for livestock losses, improved waste management to prevent dholes from scavenging in garbage sites, and controlling feral dog populations to reduce disease transmission. By addressing these challenges, coexistence between dholes and human communities can be improved, ensuring the survival of this endangered species.\u003c/p\u003e \u003c/div\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eSupplementary information\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe online version contains supplementary materials available at\u003c/p\u003e\u003ch2\u003eAcknowledgements\u003c/strong\u003e\u003c/h1\u003e\n\u003cp\u003eThe first author had received generous support from the Masinagudi field station staff during the stay in MTR. Fieldwork would have been impossible without immense help from Trackers Shiva, Mohan and the late Bomma. We would like to express our gratitude to Dr Daisuke Muramatsu for statistical advice and to Drs Sanjeeta Sharma Pokharel and Nachiketha Sharma for proof-reading. RS was a JC Bose National Fellow during the tenure of this work.\u0026nbsp;\u003c/p\u003e\n\u003ch2\u003eAuthor contributions\u003c/strong\u003e\u003c/h1\u003e\n\u003cp\u003eSS conceived the ideas, designed the methodology and collected the data; SS performed analyses and SS and SK interpreted the data; SK and RS supervised the work. SS wrote the draft of the manuscript. All authors have reviewed and contributed to writing the final version of the manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis work was financially supported by ITP-HOPE and AS-HOPE (Primate Research Institute of Kyoto University) and JSPS Core-to-Core Program, A. Advanced Research Networks (Wildlife Research Center of Kyoto University) from the Japan Society for the Promotion of Science.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAvailability of data and material\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe datasets analysed during the current study are available from the corresponding author on reasonable request.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCode availability\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003ePermits\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe Consulate General of India in Japan and the National Biodiversity Authority kindly permitted us to work in India. The Tamil Nadu Forest Department kindly granted us permission to work in MTR and provided us with its camera trap data on dholes.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthics approval\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors confirmed that the ethic policies of the journal, as noted on the journal\u0026rsquo;s guidelines page, have been adhered to. No ethical approval was required as there was no sample collection from animals or humans.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent to participate\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent for publication\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare no competing interests.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAcharya BB (2007) The ecology of the dhole or Asiatic wild dog (\u003cem\u003eCuon alpinus\u003c/em\u003e) in Pench Tiger Reserve, Madhya Pradesh, \u003cem\u003eUnpublished PhD Dissertation. 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Mammal study 46(4):323\u0026ndash;339\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003eTable 1 to 6 are available in the Supplementary Files section.\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"european-journal-of-wildlife-research","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"ejwr","sideBox":"Learn more about [European Journal of Wildlife Research](http://link.springer.com/journal/10344)","snPcode":"10344","submissionUrl":"https://submission.nature.com/new-submission/10344/3","title":"European Journal of Wildlife Research","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"habitat utilisation, livestock predation, garbage scavenging, nocturnal activity, anthropogenic impacts, human-wildlife conflict.","lastPublishedDoi":"10.21203/rs.3.rs-6674422/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6674422/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eUnderstanding the ecology of endangered dhole and their responses to anthropogenic environments is important for mitigating conflicts with humans and ensuring conservation efforts for this species. This study investigated the pack size, home range size, reproduction and activity pattern of multiple dhole packs in Mudumalai Tiger Reserve, southern India, from September 2011 to August 2015 using direct observation and camera trapping. The findings were compared across dhole packs to assess the influence of reproduction and human-modified environments on their ecology and behaviour. Of the five packs identified (P1\u0026ndash;P5), reproduction was observed three times in two packs (P1, P2) with an average litter size of seven and denning between December and February, while the others likely did not reproduce. Group sizes observed outside the den area (mean: 6.6\u0026thinsp;\u0026plusmn;\u0026thinsp;3.4) decreased from November to December, likely due to pup dispersal and den defense needs. Packs inhabiting the areas with high proportion of anthropogenic environments and feeding on livestock and garbage (P1, P2) had significantly smaller home ranges (7.0\u0026ndash;14.4 km\u0026sup2;) compared to those in more natural environments (P3, P4: 36.1\u0026ndash;40.3 km\u0026sup2;), suggesting higher foraging efficiency in human-modified landscapes. The dholes in the study area were primarily diurnal, with 74.4% of activity occurring during daylight hours. They were particularly active in the morning and evening as they frequently took very long rests over six hours during midday. However, the pack with the greatest use of anthropogenic environments (P2) showed cathemeral activity pattern increasing night activity up to 50.5%, probably to avoid encounters with humans. The high mortality of pups estimated in P2 and unnatural deaths of adults observed both in P1 and P2 suggest that the nutritional benefits and increased reproduction in anthropogenic environments could be offset by the higher mortality rates, potentially due to conflicts with humans and diseases transmitted by feral or domestic dogs.\u003c/p\u003e","manuscriptTitle":"Pack size, home range size and activity pattern of dhole (Cuon alpinus) in Mudumalai Tiger Reserve, India: effects of anthropogenic environment use and reproduction","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-06-18 13:37:05","doi":"10.21203/rs.3.rs-6674422/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-08-05T23:37:20+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-07-24T11:55:28+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-07-04T22:02:31+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"223748278900207560806400917649535235318","date":"2025-06-17T10:30:48+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"62439479754707951688854771151448899302","date":"2025-06-16T18:47:36+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-06-16T14:42:56+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-05-19T08:28:00+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-05-19T08:24:53+00:00","index":"","fulltext":""},{"type":"submitted","content":"European Journal of Wildlife Research","date":"2025-05-15T16:08:44+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"european-journal-of-wildlife-research","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"ejwr","sideBox":"Learn more about [European Journal of Wildlife Research](http://link.springer.com/journal/10344)","snPcode":"10344","submissionUrl":"https://submission.nature.com/new-submission/10344/3","title":"European Journal of Wildlife Research","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"3d26d221-1a2a-4821-a6a4-57c7e739c1ed","owner":[],"postedDate":"June 18th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-01-26T23:38:10+00:00","versionOfRecord":[],"versionCreatedAt":"2025-06-18 13:37:05","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-6674422","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6674422","identity":"rs-6674422","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}