PICA Project Report (Action A2.2 & 2.3)

Home , Canada lynx, Chinese mountain cat, Clouded leopard, Iriomote cat, Ocelot, Oncilla

Investigation of Pallas’s cat activity patterns and temporal interactions with sympatric species

Authors: Katarzyna Ruta, Gustaf Samelius, David Barclay, Emma Nygren

PICA - “Conservation of the Pallas’s cat through capacity building, research, and global planning”

1. Introduction: 1.1 Activity patterns of wild felids:

Activity patterns form a part of species’ adaptation to their environment (Beltran & Delibes, 1994) and are therefore a fundamental aspect of animal behaviour (Nielsen, 1983; Weller & Bennett, 2001). Felids are generally considered to be crepuscular and nocturnal in their activity (Kitchener, 1991), although they are well adapted to function in a wide range of light conditions (Sunquist & Sunquist, 2002). Numerous abiotic pressures and biotic interactions are known to shape the temporal behaviour of (cat-like) carnivores (Marinho et al., 2018), including changes in temperature (Beltran & Delibes, 1994; Podolski et al., 2013), light (Huck et al., 2017; Heurich et al., 2014) and season (Podolski et al., 2013; Manfredi et al., 2011), sex and reproductive status of the animal (Kolbe & Squires, 2007; Schmidt, 1999; Schmidt et al., 2009), predation risk (Caro, 2005; Farías et al., 2012) and human disturbance (Wolf & Ale, 2009; Ale & Brown, 2009). Owing to the dietary constraints of carnivores whose preys have their own well-defined circadian rhythms (Halle, 2000; Zielinski, 2000), the availability and vulnerability of prey is, however, considered as one of the main influences on predator temporal activity (Zielinski, 1988; Lodé, 1995). According to Optimal Foraging Theory, predators are expected to synchronize their daily activity with the activity of their most profitable prey, increasing the probability of encounters while reducing energy expenditure (MacArthur & Pianka, 1966; Monterroso et al., 2013; Emmons, 1987). Temporal association between predator and prey species is well documented in felids- examples include jaguar Panthera onca and puma Panthera concolor (Harmsen et al., 2011; Foster et al., 2013), tiger Panthera tigris (Linkie & Ridout, 2011; Kawanishi & Sunquist, 2004), leopard Panthera pardus (Jenny & Zuberbuhler, 2005), Eurasian lynx Lynx lynx (Podolski et al., 2013), lion Panthera leo (Schaller, 1972), Sunda clouded leopard Neofelis diardi (Ross et al., 2013) and Iriomote cat Prionailurus iriomotensis (Schmidt et al., 2009). However, majority of studies to date focus on the large and medium-bodied species, with detailed knowledge of activity patterns of many small felids and factors shaping them still poorly understood (Marinho et al., 2018; Porfirio et al., 2016; Schmidt et al., 2009).

1.2 Intraguild competition and carnivore co-existence:

Intraguild competition is another important mechanism shaping temporal behaviour of mammalian carnivores (Donadio & Buskirk, 2006; Schoener, 1974; Palomares & Caro, 1999). Inter-specific competition is widespread in ecological communities with multiple carnivore species (Donadio & Buskirk, 2006; Palomares & Caro, 1999) and can take the form of exploitation, i.e. indirect competition for a shared resource, or interference, i.e. direct encounter between sympatric species (Schoener, 1983; Hunter & Caro, 2008). Being situated at an intermediate trophic level (Prugh et al., 2009) mesopredators are uniquely affected, as being both predator and prey they need to trade-off the risk of intraguild predation against the risk of starvation (Schoener, 1974; Cozzi et al. 2012; Penido et al., 2017). The dominant apex predators might exert negative influences on the subdominant mesopredators (Crooks & Soulé, 1999), who can suffer injury and harassment (Kruuk, 1972) or in extreme cases killing (Donadio & Buskirk, 2006; Palomares & Caro, 1999). When intra-guild pressure is high, mesocarnivores might reduce the risk of competition through differentiation of prey, spatial habitat use and/or temporal activity (e.g. Davies et al., 2007; Schoener, 1974; Steinmetz et al., 2013). Niche partitioning, particularly along the space and time axes, is recognised as an important mechanism allowing carnivore co-existence (Massara et al., 2018; Schoener, 1974; Davies et al., 2007; Glen & Dickman, 2005; Polis et al., 1989) and temporal segregation is widely documented in felid

PICA - “Conservation of the Pallas’s cat through capacity building, research, and global planning” communities (for example Haidir et al., 2018, Lucherini et al., 2009; Steinmetz et al., 2013; Azlan & Sharma, 2006; Li et al., 2012; Di Bitetti et al. 2010; Romero- Muñoz et al., 2010). Small-bodied species are under the greatest competitive pressure, while being at higher risk of predation by other carnivores (Hunter & Caro, 2008; Bischof et al., 2014). Yet, research on spatiotemporal interactions amongst mammalian carnivores has hitherto been biased towards large-sized flagship species, whilst knowledge about how small mesopredators deal with being both hunter and hunted is still limited (Haidir et al., 2018; Bischof et al., 2014).

1.3. Study species:

The Pallas’s cat (Otocolobus manul), is a small-sized felid with a wide but fragmented distribution across a large proportion of Central Asia, stretching from eastern Mongolia to western Iran (Ross et al., 2016; Heptner & Sludskii, 1992). It is classified as Near Threatened on the IUCN’s Red List and thought to be declining (Ross et al., 2016). Pallas’s cats are generally associated with montane grassland steppe and shrub steppe (Ross, 2009; Ross et al., 2016), however, they also exhibit habitat specificity, with a marked preference for habitats providing cover from predators, such as ravines and rocky areas (Ross, 2009; Ross et al., 2012). Their optimal habitat is believed to consist of a mixture of grassland and shrub steppe with rocky cover, ravines and hillslopes (Ross et al., 2016; Ross, 2009). Mongolia is believed to be a stronghold of the Pallas’s cat (Ross et al., 2016), where it is also known to occur across several mountain ranges in the South Gobi region (Barclay, 2019- personal communication), making the Tost Nature Reserve a suitable landscape to study the species. Owing to its rarity and elusive behaviour, little is known about Pallas’s cat ecology in the wild (Ross et al., 2010a; Murdoch et al., 2006); it is als among the world’s most understudied wild felids (Brodie, 2009; Tensen, 2018).

1.4 Study objectives:

This report summarises camera-trap results from three surveys conducted between 2012- 2015 across Tost Mountains, a protected area in Mongolia’s south Gobi. Camera- trapping technology has become one of the most important tools for studying wide- ranging, rare and elusive species (Rowcliffe & Carbon, 2008; McDonald & Thompson, 2004) and is therefore considered as the most effective methodology for surveying felids (for example Karanth et al., 2006; Azlan & Sharma, 2006; Wearn et al., 2013). Time-stamped data derived from camera traps also provides an opportunity to investigate unresolved aspects of wild animals’ ecology and community dynamics, including variations in activity patterns and segregation along the time niche axis (Frey et al., 2017), as well as those linked with rarity and elusiveness, particularly in lesser-studied, small bodied carnivore species (Bischof et al., 2014).

The study objectives were as follows:

1.) To enhance knowledge on daily and seasonal activity of the Pallas’s cat 2.) To enhance knowledge on temporal association/disassociation of the Pallas’s cat with sympatric species detected at the study site

2. Methods: 2.1 Study area:

The camera-trapping surveys took place within Tost Tosonbumba Nature Reserve located in the South Gobi province of southern Mongolia (43.2⁰N, 100.5⁰E; see Fig. 1 & 2). The reserve encompasses almost

PICA - “Conservation of the Pallas’s cat through capacity building, research, and global planning”

7500 km² and is comprised of Tost and Tosonbumba Mountains. It also serves as a bridge between two existing Protected Areas, the Great Gobi and the Gobi Gurvansaikhan National Park, making it one the world’s largest protected habitats for snow leopards (Snow Leopard Trust, 2016), and thus sympatric species. The temperature varies between -35⁰C in the winter and +35⁰C in the summer and the altitude ranges between 1600 and 2400 m above sea level. The study area is mostly surrounded by steppe, with few isolated rugged hillocks (see Fig. 3 & 4) and has prominent human presence towards the east, including road infrastructure, mining establishments and the Gurvantes township, followed by the Noyon Mountains (Sharma et al., 2014). The area is home to a variety of carnivore predators, including snow leopard (Panthera uncia), lynx (Lynx lynx), gray wolf (Canis lupus), red fox (Vulpes vulpes), stone marten (Martes foina) and the Pallas’s cat (Otocolobus manul), as well as several prey items including pika (Ochotona sp.), Tolai hare (Lepus tolai), Argali (Ovis ammon), Chukar partridge (Alectoris chukar) and several species of rodents (Shehzad et al., 2012)

Fig. 1. Location of Tost Mountains study site in South Gobi, Mongolia (camera trap locations are marked in red; map source- the Snow Leopard Trust)

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Fig. 2: Close-up of camera trap locations in Tost Nature Reserve (left-bottom corner). The locations within the study area were adjusted each year after conducting new surveys to ensure best placements for any given year (Sharma et al. 2014; map source- the Snow Leopard Trust)

Fig. 3 & 4: Landscape type within the study area in Tost mountains (images provided by the Snow Leopard Trust)

2.2 Survey design and camera deployment:

Current data was originally collected by the Snow Leopard Trust, as a part of the organisation’s long- term ecological study on the snow leopard Panthera uncia in South Gobi, Mongolia (SLT, 2019). The field methodology used by the SLT was previously outlined in a publication by Sharma and colleagues (2014) and is reproduced below.

The cameras were deployed across 1,648 km² within the entire mountainous area of Tost that was separated from other mountains by at least 20 km of steppe. Since the (original) study aimed to monitor abundance and population dynamics of the snow leopard (Sharma et al., 2014), a grid of 5x5 km on the mountainous part of the study area was overlaid with one camera trap in each grid cell, allowing for about 2 camera stations per home range (Karanth & Nichols, 1998; Sharma et al., 2014). Within each grid cell the cameras were installed at sites with most recent signs of snow leopard activity (e.g. urine marking or faeces) along saddles on the ridgelines, valley or cliff bases and near overhanging boulders. In cases where the grid cell did not have ample snow leopard habitat or any signs of snow leopard presence, the camera placements were rarefied. The resultant density varied between 2 and 4 camera trapping stations per 100 km² (Sharma et al. 2014). Overall, 35-40 camera stations were in operation each year, with one camera trap (Reconyx RM45, RC40 or HC500) at each station. The traps were checked every two weeks to ensure battery and memory card adequacy. At least 2 Gigabyte cards were used, allowing each camera to store more than 10,000 images. The Reconyx camera trigger sensors are calibrated to use a combination of motor and heat sensors, preventing unnecessary triggers from heated rocks or movements of grass within the camera’s field view. The traps were set to take 5 pictures per trigger, at an interval of 0.5 second between each picture and no time period set between the triggers (Sharma et al. 2014).

2.3 Data analysis:

The camera trap data was provided by the Snow Leopard Trust and was then analysed using software developed by the ZSL (Zoological Society of London) specifically to process images from camera trap

PICA - “Conservation of the Pallas’s cat through capacity building, research, and global planning” arrays (Amin et al., 2016). This requires creation of four standard format data source files in Microsoft Excel outlining 1.) individual camera locations and associated fixed habitat variables 2.) individual camera settings and configurations 3.) individual camera setup, service and recovery history, and 4.) image details for every photograph from each camera. To create the latter file, metadata (image filename, date, time, temperature) were extracted automatically from folders of the original jpg image files using Exiv2 software (Huggel 2012; http://www.exiv2.org/index.html) and entered into standard Excel formats. Date and time information in the meta-data were cross-checked against images. Details of each image content indicating image type (wildlife, livestock or preselected categories of ‘other’) and species identification (with information on number, age, sex and other behaviours where appropriate) were then added manually by visual inspection (Bruce et al., 2016).

Species trapping rates were calculated as the mean number of independent photographic “events” per trap day x 100, only using cameras that operated for more than 75% of the survey time period for each survey grid. An “event” was defined as any sequence of images from a given species occurring after an interval of ≥60 min from the previous images of that species (Tobler et al., 2008). Trapping rate provides a simple index of relative abundance (RAI), assuming that a target species will trigger cameras in relation to their density, all other factors being equal (Bruce et al., 2016).

Circadian (24 hour) activity patterns were constructed from the number of independent photographic events per hour. Observations were classified as diurnal (if activity occurred between 1 hr after sunrise and 1 hr before sunset), nocturnal (if activity occurred between 1 hr after sunset and 1 hr before sunrise) and crepuscular (if activity occurred up to 1 hr before and after sunrise and sunset), as per Porfirio et al. (2016). Times of sunrise and sunset at the time of trapping were obtained from an online tool set for the city closest to the study location in Mongolia- i.e Dalanzadgad (https://www.timeanddate.com/sun/mongolia/dalanzadgad). Due to the low sampling effort of the Pallas’s cat in current survey, no statistical analyses could be performed on the data and thus inferences about activity were made based solely on observed patterns.

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3. Results: 3.1 Camera trap survey effort:

All three camera trap surveys achieved the recommended minimum sampling effort of 1000 camera- trap days (O’Brien et al. 2003) (see Table 1 for a breakdown per survey).

Total number of camera stations: 112

Total number of days deployed: 10 751 (9724 operational)

Total number of wildlife events: 2 904

Table 1: Summary of camera trap effort per survey

Survey duration 29/05/2012- 15/07/2013- 05/08/2015- 08/10/2012 16/11/2013 06/12/2015 Total number of camera stations 40 35 37 Total number of days deployed 4391 2229 4131 Total number of days operational 4064 1966 3694 Total number of wildlife events 1006 1051 847

3.2 Species diversity:

A total of 14 (identifiable) mammal and 3 bird species were photographed in Tost Nature Reserve over the course of the study (see Table 2). Species that were not distinctive enough to be identified in the camera trap images were classified at level of family or subfamily; these are not included in the list below.

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Table 2: Species identified in Tost Nature Reserve (2012-2015)

Order Family or Species Common name IUCN No. subfamily status events Mammalia Carnivora Canidae Canis lupus Gray wolf LC 26 Carnivora Canidae Vulpes corsac Corsac fox LC 1 Carnivora Canidae Vulpes vulpes Red fox LC 676 Carnivora Felidae Otocolobus manul Pallas’s cat NT 9 Carnivora Felidae Panthera uncia Snow leopard VU 318 Carnivora Mustelidae Martes foina Stone marten LC 129 Cetartiodactyla Bovidae-Caprinae Capra sibirica Siberian ibex LC 401 Cetartiodactyla Bovidae-Caprinae Ovis ammon Argali NT 30 Lagomorpha Leoporidae Lepus tolai Tolai hare LC 220 Lagomorpha Ochotonidae Ochotona sp. Pika species LC 36 Aves Galliformes Phasianidae Alectorus chukar Chukar partridge LC 465 Columbiformes Columbidae Columba livia Rock dove LC 1 Accipitriformes Accipitridae Aquila chrysaetos Golden eagle LC 1 Non-wildlife species Cetartiodactyla Bovidae-Caprinae Shoat Sheep & goat LC 50 Carnivora Canidae Canis familiaris Domestic dog 15 Cetartiodactyla Camelidae Camelus Domestic 3 bactrianus bactrian camel Perissodactyla Equidae Equus caballus Horse 9 IUCN Red List categories: LC= Least Concern, NT= Near Threatened, VU= Vulnerable, EN= Endangered

3.3 Species abundance:

The red fox was the most commonly captured mammal on camera traps (RAI=7.99), followed by the Chukar partridge (RAI=5.07). The Pallas’s cat was the least abundant species in the current study (RAI=0.10; see Fig. 5).

Species RAI

9 7.99 8 7 6 5.07 5 4 3.06 3 2.42 2 0.58

Meantrapping rate 1 0.31 0.25 0.21 0.10 0

Fig 5. Relative abundance index (camera trap events/day x 100) of Pallas’s cat and sympatric species (in descending order)

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3.4 Pallas’s cat activity pattern: 3.4.1 Circadian:

The Pallas’s cat was captured at various times throughout the day, with a slight peak of activity around midday (11.00; see Fig. 6 & 7). Majority of detections occurred during the day (56%, n=5) suggesting a mostly diurnal activity pattern, with crepuscular and nocturnal tendencies (both 22%, n=4).

0 23 1 22 2 21 3 20 4

19 5

18 6

17 7

16 8 15 9 14 10 13 11 12

1 No.independent photographicevents 0 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 Hour of the day

Fig. 6 & 7: Circadian activity patterns of the Pallas’s cat

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3.4.2 Circannual:

Monthly: Current surveys ran between the months of May and December. Within that period, the Pallas’s cat showed a slight peak of activity in August (n=3), and stable activity levels from September to November (all n=2; see Fig. 8). No activity was detected in May, June, July or December.

2 events

No.independent photographic 0

Fig. 8: Monthly activity pattern of the Pallas’s cat

Seasonal: Following the approach of Ross (2009), two seasons were recognised based on variations in daily temperatures: summer (15 April- 10 October) and winter (15 October-14 April). Of all detections (n=9), majority took place in the summer (n=5, 56%). The Pallas’s cat did not show pronounced differences in circadian activity between seasons (Fig. 9).

1 No.independent photographicevents 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23

Summer Winter

Fig. 9: Seasonal activity pattern of the Pallas’s cat

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3.5 Temporal interactions with sympatric species: 3.5.1 Pallas’s cat and potential predators:

Pallas’s cat activity was analysed in relation to four sympatric carnivores- one felid (snow leopard, Panthera uncia) and three canids (gray wolf Canis lupus, red fox, Vulpes vulpes and domestic dog Canis familiaris). The degree of Pallas’s cat temporal segregation was the highest for the red fox, which exhibited a strong nocturnal activity pattern, followed by the snow leopard that was mostly nocturnal with crepuscular tendencies. The gray wolf seemed to exhibit a mostly crepuscular-nocturnal activity pattern, only slightly overlapping with that of the Pallas’s cat around sunset. The degree of temporal disassociation was the lowest for the domestic dog, which similarly to the Pallas’s cat was mostly active during the day (see Fig. 11 for a breakdown per species).

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Pallas’s cat:

0 23 1 22 2 21 3 20 4 19 5

18 6 17 7 16 8

15 9 14 10 13 11 12

Snow leopard: Gray wolf:

0 0 23 1 23 1 22 2 22 2 21 3 21 3 20 4 20 4 19 5 19 5 18 6 18 6 17 7 17 7 16 8 16 8 15 9 15 9 14 10 14 10 13 11 13 11 12 12

Red fox: Domestic dog:

0 0 23 1 23 1 22 2 22 2 21 3 21 3 20 4 20 4 19 5 19 5 18 6 18 6

17 7 17 7 16 8 16 8

15 9 15 9 14 10 14 10 13 11 13 11 12 12

Fig. 10: Daily activity pattern of the Pallas’s cat in relation to potential predator species

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3.5.2 Pallas’s cat and potential prey:

Predator-prey interactions of Pallas’s cat were analysed in relation to four potential prey species- three mammal (pika Ochotona sp., Tolai hare Lepus tolai, Argali Ovis ammon) and one bird species (Chukar partridge, Alectoris chukar). The temporal peaks of Pallas’s cats seemed to match the closest the activity of pika, with the highest bout of Pallas’s cat activity overlapping the highest bout of activity of pika (at 11.00). Some degree of overlap was also found with the activity of Chukar partridge and Argali, both of which also exhibited mostly diurnal activity. The degree of temporal association was the smallest for the Tolai hare, which was predominantly nocturnal (see Fig. 11 for a breakdown per species).

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Pallas’s cat:

0 23 1 22 2 21 3 20 4 19 5

18 6

17 7 16 8 15 9 14 10 13 11 12

Pika: Argali:

0 0 23 1 23 1 22 2 22 2 21 3 21 3 20 4 20 4 19 5 19 5 18 6 18 6 17 7 17 7 16 8 16 8 15 9 15 9 14 10 14 10 13 11 13 11 12 12

Tolai hare: Chukar partridge:

0 0 23 1 23 1 22 2 22 2 21 3 21 3 20 4 20 4 19 5 19 5 18 6 18 6 17 7 17 7 16 8 16 8 15 9 15 9 14 10 14 10 13 11 13 11 12 12

Fig. 11: Daily activity pattern of the Pallas’s cat in relation to potential prey species

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4. Discussion: 4.1 Pallas’s cat circadian activity pattern:

Knowledge on temporal activity of smaller-bodied felid species is limited, but studies to date suggest that majority of medium- and small-sized felids are primarily nocturnal and/or crepuscular- examples include ocelot (Emmons, 1988), Geoffroy’s cat (Manfredi et al., 2011; Pereira, 2009), Iriomote cat (Schmidt et al., 2003), clouded leopard, Asiatic golden cat (Grassman et al., 2006), Pampas cat (Lucherini et al., 2009), northern tiger cat (Marinho et al., 2018) and guina (Delibes-Mateos et al., 2014). The activity of Pallas’s cats in the current study seems inconsistent with that of other small felids, who displayed a mostly diurnal activity pattern with crepuscular and nocturnal tendencies. However, numerous biotic and abiotic factors can influence activity patterns of wild felids (Marinho et al., 2018) which consequently seem flexible both between and within species (Wilson & Mittermeier, 2009; Beltran & Delibes, 1994). Leopard cat activity varies from nocturnal (Bashir et al. 2014), crepuscular (Azlan & Sharma, 2006) to arrhythmic (Grassman et al., 2005) and Geoffroy’s cat was found to switch its activity to diurnal during a period of food shortage (Pereira, 2010). An increase of activity around dusk and dawn was also noted in Iriomote cat (Schmidt et al., 2003) and guina (Freer, 2004). While early investigations of the Pallas’s cat suggested the species to be primarily nocturnal (Murdoch et al., 2006), its temporal peaks were later found to coincide with a crepuscular activity pattern (Ross, 2009). However, Pallas’s cats can be active at any point of the day (Ross, 2009), which was also noted by anecdotal accounts that spotted the species at all times of the day (Webb et al., 2014). Moreover, Pallas’s cats were found to visit a communal carnivore sign post in China mostly during daytime hours (Li et al., 2012). A scat analysis of Pallas’s cat dietary composition also confirmed them as primarily crepuscular or diurnal hunters (Ross et al., 2010b), which current results seem to partially support.

4.2 Pallas’s cat circannual activity pattern:

The Pallas’s cat was detected more commonly in summer than in winter, which could indicate a suppressed activity of this species during winter months. One of the detections also occurred only one day after the 15th October cut-off for winter season in Mongolia originally determined by Ross (2009), and thus depending on the temperature at the time of the given survey it could have fallen under the summer season, potentially emphasising that phenomenon. Felids have a high basal metabolic rate, and thus daily tasks such as body maintenance, movement or resource acquisition require a great energy expenditure (McNab, 1989). Given the additional energetic demands of thermoregulation to manage cold stress (Pereira, 2010), decreasing activity during winter could yield considerable energy savings (Zielinski, 2000). Reduced activity during winter has been noted in bears (Linnell, 2000), mustelids (Zielinski et al. 1983; Zschille et al., 2010; Zalewski, 2000), procyonids (Ewer, 1973) and small Arctic mammals (Chappell, 1980). A negative influence of temperature on activity levels was also reported for feral domestic cats (Izawa, 1983). Temperature can change dramatically between seasons in Mongolia (Ross, 2009) and wild-living Pallas’s cats show preference for marmot burrows in the winter (Ross et al. 2010a), presumably as being over 1 meter deep they provide good protection from the cold (Davenport, 1992). Winter dens of Pallas’s cats also have significantly smaller entrances than summer dens, thus restricting air circulation and increasing heat retention during cold weather (Ross et al., 2010a). Therefore, the species could be spending more time inside marmot dens during winter, as they provide them with a means to thermoregulate. Reduced activity of Pallas’s cats during months with snow cover was suggested by preliminary field data (Munkhtsog et al., 2004). Furthermore, deep snow can impede movement of Pallas’s cats due to their built and is therefore a limiting factor in the distribution (Sunquist & Sunquist, 2002)- the species is rarely found in areas with mean 10-day snow exceeding 10 cm (Ross, 2009). Pallas’s cat mortality also seems biased towards winter- in one 3-year

PICA - “Conservation of the Pallas’s cat through capacity building, research, and global planning” long study over 80% of deaths occurred between October and April (Ross, 2009) and anecdotal accounts from the field report finding several individuals in very poor condition or dead at times that coincided with excessive snowfall (Barclay, 2019- personal communication). Therefore, suppression of activity during winter could represent a behavioural adaptation of the Pallas’s cat to the harsh environmental conditions of Mongolia’s South Gobi and help explain current pattern of results.

Winter is also usually associated with depletion of prey base, and nutritional stress might lead individuals to modify their temporal behaviour to maintain their body condition and energy balance (Trites & Donnelly, 2003). Seasonal variation of prey availability was reported to alter activity of some small felids- Geoffroy’s cats (Leopardus geoffroyi) were found to be less active (Manfredi et al., 2011) as well as demonstrating an apparent shift toward increased diurnal activity during winter months, possibly compensating their food intake with alternate prey (Pereia, 2010). On the contrary, seasonality does not seem to affect activity patterns of ocelots (Leopardus pardalis), who were mostly nocturnal in both dry and rainy seasons (Porfirio et al., 2016). Results on seasonal activity of leopard cat (Prionailurus bengalensis) are mixed- whilst one study reported lower activity of the species during the cooler (dry) season (Grassman, 2000), others found none or minimal seasonal variation (Austin et al. 2007; Grassman et al., 2005). No pronounced seasonal variability in daily activity patterns of the Pallas’s cat was found in the current survey. It could therefore mean that similarly to some other small wild felids, Pallas’s cat temporal behaviour is not influenced by seasonality. Alternatively, it could suggest that the abundance and vulnerability of its prey did not vary substantially between seasons. The Pallas’s cat is known to preferentially select pikas (Ross et al., 2010b; Heptner and Sludskii 1992; Sunquist and Sunquist 2002), whose density in Mongolia seems similar across seasons (Ross et al., 2010b) and who as a non-hibernating species is also active during winter, making it an important prey item for a variety of mammalian carnivores (Flux & Angermann, 1990; Bischof et al., 2014). Nonetheless, reliance of pikas on underground food stores (haypiles) throughout winter (Dearing, 1997; Varner et al., 2016) and inclement weather might reduce the amount of time they spend on the surface during that season, thus limiting their availability to predators (Sokolov et al. 2009; Ross et al., 2010b). Whilst a pika specialist, the Pallas’s cat also utilises a broad range of other prey items, including rodents, lagomorphs, insects, reptiles, birds and carrion (Ross et al., 2010b). Analysis of Pallas’s cat dietary composition also revealed a significant increase in consumption of insects and a decrease in pika consumption during winter (Ross et al., 2010b). Therefore, the Pallas’s cat might be adjusting its diel activity to the activity of various prey species whose circadian rhythms vary seasonally, thus demonstrating a degree of dietary flexibility, in accordance with previous investigations that deemed the Pallas’s cat a facultative specialist predator of pika (Ross et al., 2010b; Ross, 2009).

In terms of monthly activity, the Pallas’s cat demonstrated a slight increase in August, and stable levels throughout September through to November. This pattern could be simply a result of the duration of the current survey, which only covered the latter part of the year from May to December. Nevertheless, studies on captive Pallas’s cats carried out at Moscow Zoo also reported heightened activity around October-November (Alekseicheva, 2009), which was associated with an increase in daily food consumption that in captive Pallas’s cats can start as early as June (Demina, 2006). October, however, is said to mark the onset of ‘food craving’ behaviour, when the animals are observed longingly waiting for arrival of food, occasionally even grabbing it from the keepers’ hands. It was suggested that this is due to the inherent tendency of Pallas’s cat to increase their body mass prior to the breeding season (Alekseicheva, 2009). This could potentially explain detections of Pallas’s cats during the months of October-November in the current survey, although a longitudinal study is needed to determine that with more certainty. Furthermore, since activity patterns form a part of species’ adaptations to their environment (Beltran & Delibes, 1994), activity patterns of captive

PICA - “Conservation of the Pallas’s cat through capacity building, research, and global planning” animals are likely to differ from those of their wild-living counterparts. Nonetheless, increased activity between September to November was also recorded in wild-living Pallas’s cats, which coincided with times of energetic stress (Ross, 2009) and some studies on captive ocelots showed that majority of cats retained their natural activity patterns (Resende et al., 2014; Weller & Bennett, 2001). Captive studies can therefore provide valuable complimentary information on temporal behaviour of wild- living animals when conducting field surveys is unfeasible, as sometimes could be the case in Tost Mountains due to the severity of climate.

Sex and reproductive status could also have affected patterns of Pallas’s cat seasonal activity observed in this study. Guina (Leopardus guigna) was significantly more active in spring than during autumn, possibly due to searching behaviour of available mates (Freer, 2004) and the Iriomore cat (Prionailurus iriomotensis) showed seasonal variation in activity modulated by its reproductive status (Schmidt et al., 2009; Schmidt et al., 2003). Studies on captive Pallas’s cats indicate that they exhibit a pronounced reproductive seasonality, with distinct reproductive patterns between the breeding (December- April) and non-breeding (June-October) seasons (Swanson et al., 1996). Activity patterns of the wild-living individuals also seem to vary seasonally between males and females (Ross, 2009). Future studies could therefore benefit from running year-round surveys, encompassing the breeding season (January- March in wild Pallas’s cats; Ross, 2009), while accounting for the effects of sex and reproductive status whenever possible.

Human disturbance may also exert influence on both daily and seasonal activity patterns of the Pallas’s cat. A recent meta-analysis has revealed a strong effect of human disturbance on temporal dynamics of wild-living animals, with 62 mammal species across 6 continents found to alter their diel activity patterns towards increased nocturnality to avoid contact with humans (Gaynor et al., 2018). Much of Pallas’s cat home range is inhabited by nomadic pastoralists, who move their livestock several times a year seeking seasonal pastures (Marin 2010; Mearns, 1993) and aspects of Pallas’s cat den selection patterns appear suggestive of human-avoidance behaviour- the animals chose dens significantly further away from human camps only during winter months when they were occupied, but not during summer months when they were empty (Ross et al., 2010a). Therefore, Pallas’s cats might need to alter their seasonal spatial (Munkhtsog et al., 2004) and/or temporal behaviour (Farhadinia et al., 2016) in avoidance of seasonally occupied areas. However, anecdotal evidence suggests that the species is also capable of inhabiting areas subject to heavy human disturbance (Webb et al., 2014). Such tolerance could nonetheless put it at more risk of anthropogenically-induced behavioural changes, which will be discussed in the next section.

4.3 Temporal interactions of the Pallas’s cat and potential predator species:

The snow leopard in the current survey displayed a nocturnal/crepuscular activity pattern, consistently with previous studies conducted on the species in southwest Mongolia that found it to be nocturnal with a slightly crepuscular trend (McCarthy et al., 2005) or strongly crepuscular in Nepal and elsewhere in Mongolia (Schaller et al., 1994). The peaks of Pallas’s cat activity appeared to coincide with the lowest activity of snow leopard during daytime hours, potentially demonstrating temporal disassociation to reduce the chances of interspecific encounters. Smaller species are more likely to be predated upon by other carnivores (Hunter & Caro, 2008; Donadio & Buskirk, 2006; Bischof et al., 2014) and time partitioning from a larger carnivore was previously reported in Neotropical canids- the crab-eating fox and the pampas fox were both nocturnal while in allopatry, whereas in sympatry the smaller species, the pampas fox, reduced its activity when the activity of the crab-eating fox was high (Di Bitetti et al., 2009). Given the morphological and ecological similarity of cat species (Johnson et al., 2006; Morales & Gianini, 2010) and their territoriality (Palomares & Caro, 1999), the

PICA - “Conservation of the Pallas’s cat through capacity building, research, and global planning” potential for negative interactions could be expected to be even higher between felids. Niche partitioning is well documented in felid communities (e.g. Haidir et al., 2018, Lucherini et al., 2009; Steinmetz et al., 2013; Azlan & Sharma, 2006; Li et al., 2012; Di Bitetti et al. 2010; Romero- Muñoz et al., 2010). Intra-guild pressure from a larger felid specifically was also found to affect the activity on oncilla, who was primarily nocturnal in absence of larger cat species- margay, ocelot and puma, but became more diurnal when they were present, a temporal activity flexibility that was suggested to reflect avoidance of interspecific conflict in a felid guild (Oliveira- Santos et al., 2012); in another study oncilla also appeared to separate itself temporally from jaguar and larger mesopredators (Penido et al., 2017). Interspecific killing, the most extreme form of interference competition, is common among mammalian carnivores, including felids (Palomares & Caro, 1999; Donadio & Buskirk, 2006). The snow leopard is not amongst known predators of the Pallas’s cat (Ross, 2009) and investigation of snow leopard prey preferences in South Gobi, Mongolia found no Pallas’s cat material in its faeces (Shehzad et al., 2012), which could suggest that the potential for intraguild predation by the snow leopard is low. Nonetheless, a dominant killer species may not necessarily consume its victim (Palomares & Caro, 1999; Ritchie & Johnson, 2009)- for example cougars were found to kill coyotes and bobcats but not to eat them, presumably as they had access to an alternative or preferred food source (Koehler and Hornocker 1991). Furthermore, even when interspecific killing is rare (lethal effects), apex predators can still exert negative influence on mesopredators by instilling fear (non-lethal effects), which can also motivate changes in behaviour to diminish interspecific encounters (Ritchie & Johnson, 2009; Mukherjee et al., 2009) and/or by reducing their effectiveness as predators (mesopredator suppression- Prugh et al., 2009; Bischof et al., 2014) The threat of predation alone, or perceived predation risk, can translate to equally or even more profound impacts on mesopredator behaviour and ecology than direct kills (‘ecology of fear’- Brown et al., 1999; Ritchie & Johnson, 2009; Sergio et al., 2007). Those risk effects can in turn induce the subordinate animals to avoid habitats and time periods at which the dominant species is active (Lima & Dill, 1990; Ritchie & Johnson, 2009; Steinmetz et al., 2013). Activity patterns of Pallas’s cat in the current study could therefore be suggestive of a temporal adaptation in response to (perceived) predation risk from the snow leopard. A study of a communal sign post on Tibetan Plateau, China, sniffed and marked by the snow leopard and sympatric carnivores also found evidence of temporal segregation between the snow leopard and the Pallas’s cat – the snow leopard visited the site mostly throughout the night and occasionally at dawn or dusk, whereas the Pallas’ cat was found to only come throughout the day (Li et al., 2012).

The degree of temporal partitioning between the snow leopard and the Pallas’s cat, however, was lower than between the snow leopard and gray wolf in the aforementioned study, suggesting that the degree of segregation along the time niche may be dependent on the position of the species in the food chain and its body size (Li et al., 2012). This is consistent with the idea that inter-specific competition should increase as species become more similar (Morin, 1999; Donadio & Buskirk, 2006). Dietary overlap is suggested as a strong factor motivating interspecific killing and aggression in carnivore communities (Donadio & Buskirk, 2006; Schaller, 1972) and the dietary habits of snow leopard and Pallas’s cat differ substantially- whilst the former preferentially hunts the Siberian ibex (Capra sibirica) followed by domestic goat (Shehzad et al., 2012), the latter disproportionately selects pika followed by small rodents (Ross, 2009; Ross et al., 2010b). The only prey item they could be competing over is Argali (Ovis ammon), however it is only the third most common prey consumed by the snow leopard (Shehzad et al., 2012) and predation of Argali lambs by the Pallas’s cat is sporadic (Reading et al., 2005; Murdoch et al., 2006) and thus the potential for food competition is low.

PICA - “Conservation of the Pallas’s cat through capacity building, research, and global planning”

Furthermore, similarity in body size is a strong motivator for inter-specific competition (and interspecific killing) (Morin, 1999; Donadio & Buskirk, 2006), and with a body weight ranging between 25- 75 kg (Sunquit & Sunquist, 2002) the snow leopard is over 10 times the size of the Pallas’s cat (2.5-5 kg; Heptner & Sludksii, 1992; Ross, 2009). Therefore, an alternative explanation for the current result could be that the niches of the Pallas’s cat and the snow leopard are distinct enough to prevent competition, thus allowing them to live in sympatry with one another. The Pallas’s cat was also observed to co-exist with another species of felid, the Chinese mountain cat (Felis bieti) on the Tibetan Plateau in Sichuan (Webb et al., 2016). Furthermore, due to the regulating effects of apex predators on trophic cascades, presence of a large predator such as the snow leopard might signal a healthy, well-functioning ecosystem (Ripple et al., 2014), thus allowing other species such as the Pallas’s cat to thrive.

The activity of gray wolf is generally nocturnal (Karamanlidis et al., 2017; Kusak et al., 2005; Ciucci et al. 1997) although it can vary across sites (Theuerkauf et al., 2003). Albeit based on few observations, the activity peaks of gray wolf in this study seem to indicate a crepuscular-nocturnal pattern. Increased activity at dawn and dusk was previously noted in gray wolf in Poland, which was also associated with periods where they killed the most prey (Theuerkauf et al., 2003). Gray wolves across Mongolia are heavily reliant on wild and domestic ungulates (Hovens and Tungalaktuja, 2004), and thus the risk of dietary competition between them and Pallas’s cats is low. However, gray wolves are known to occasionally predate on Pallas’s cats (Ross, 2009) and intraguild predation is also a strong driver of mammalian behaviour that can lead to shifts in activity patterns by the subordinate species (Caro 2005; Farías et al., 2012). Therefore, Pallas’s cat mostly diurnal activity might be a manifestation of a temporal mechanism minimising the risk of lethal encounters with the gray wolf. Whilst the Pallas’s cat showed crepuscular tendencies, its activity only slightly overlapped with that of the gray wolf at dusk but not at dawn, potentially indicating a degree of crepuscular segregation linked with avoidance of conflict.

The red fox displayed a strongly nocturnal activity pattern, consistently with findings from many parts of its range, e.g. Japan (Takeuchi et al., 1992), Bulgaria (Georgiev et al., 2015), Spain (Díaz‐Ruiz et al., 2016) and England (Doncaster & Macdonald, 1997). Diet of red foxes in Mongolia was found to consist mainly of insects, followed by small rodents and lagomorphs (Murdoch et al., 2010); on the Tibetan Plateau in China they also preyed heavily on small rodents and pikas, followed by marmots and hares (Schaller, 1998). Therefore, red foxes and Pallas’s cats have considerate dietary overlap, increasing the risk of competitive exploitation and interference. It has also been suggested that the risk of intraguild killing due to competition is the highest when the larger species is 2.0-5.4 times the size of the smaller one (Donadio & Buskirk, 2006). Among carnivores living in sympatry with the Pallas’s cat in Tost Nature Reserve, the red fox would therefore be expected to constitute the greatest competitor. This seems congruent with current results, as the Pallas’s cat showed the strongest temporal disassociation with the red fox. Since the red fox is a habitat generalist, spatial avoidance may not be a viable option for smaller carnivores (Bischof et al., 2014) who may be forced to adapt their temporal behaviour instead. This was previously reported for the Altai mountain weasel (Mustela altaica) that was observed frequenting areas used by its main prey, pika, while exhibiting diurnal activity contrasting with that of its main predators, stone marten and red fox (Bischof et al., 2014). Pallas’s cats are also known to be predated on by the red fox (Ross, 2009) and to preferentially hunt pika (Ross et al., 2010b)- given those similarities in trophic level and ecology, the Pallas’s cat might display similar behavioural adaptations to the Altai mountain weasel. Activity of oncilla was also found to be

PICA - “Conservation of the Pallas’s cat through capacity building, research, and global planning” dissimilar from larger sympatric mesopredators, i.e. ocelot and crab-eating fox, possibly reflecting temporal niche segregation (Penido et al., 2017). Previous field studies of Pallas’s cat activity indeed suggest that the species uses time to separate itself from predators, thereby minimising competition and predation risk (Ross, 2009). Moreover, some of Pallas’s cat observations in the wild were noted to occur in an area with few canids- whilst only anecdotal, this evidence could demonstrate that the Pallas’s cat prefers areas where foxes and wolves occur at low densities (Webb et al., 2014), thus indicating niche segregation along the space dimension as well.

Domestic dogs were also present within the Pallas’s cat-inhabited area in Tost Nature Reserve. Pastoralists live throughout a large proportion of Mongolian steppe, and most households own at least one dog, usually of large breed, for personal or livestock protection (Olson et al., 2011; Buuveibaatar et al. 2009). The detrimental effects of free-roaming dogs on wildlife are increasingly recognised (Young et al., 2011). Direct predation is the most apparent threat of dogs (Ritchie et al., 2014; Young et al., 2011) and they are a known introduced predator of the Pallas’s cat (Barashkova and Smelansky 2011, Ross et al. 2012). However, they can also negatively impact wildlife indirectly through inducing fear-mediated behavioural changes (Doherty et al., 2017), including shifts in activity patterns previously noted in response to dog presence (Gerber et al., 2012; Zapata-Ríos & Branch, 2016). Domestic dogs also occasionally ingest carnivore scats (coprophagy; Boze, 2010) and urinate/defecate on marking sites of other animals (Bekoff, 2001). Such scent marking disturbance might hinder the ability of carnivores to detect one another, making them unaware of each other’s presence and thereby increasing opportunities for interaction (Lewis et al., 2015). Surprisingly, the Pallas’s cat did not show temporal partitioning from the domestic dog, as both species were mostly active during the day. As a mesopredator living in the ‘landscape of fear’ (Ritchie & Johnson, 2009; Laundré et al., 2001), the Pallas’s cat needs to constantly trade off food for safety (Cozzi et al., 2012; Prugh et al., 2009; Penido et al., 2017). Therefore, it is possible that in this case the species might benefit more from adjusting its activity to that of its staple prey, i.e. diurnal pika, than to avoid antagonistic interactions with the domestic dog- such strategy was previously suggested to be particularly beneficial for mesocarnivores living in environments with low productivity and availability of prey (Penido et al., 2017). Another possibility is that separating itself temporally from the domestic dog could put the Pallas’s cat at risk of negative interactions from the wild-living, potentially more dangerous predators, and so the safety trade-off might work in favour of being in sympatry with the domestic dog. However, more studies are needed to investigate the exact factors shaping the temporal interactions between domestic dogs and Pallas’s cats. Regardless of the mechanisms behind it, such temporal overlap with domestic dogs can still put the Pallas’s cat at risk of fear-mediated behavioural changes though, which could have detrimental effect for fitness (Silva-Rodríguez & Sieving, 2012).

With domestic dogs also come humans, whom the dogs often accompany whilst they tend to their livestock across Mongolia (Buuveibaatar et al., 2009). Humans are disproportionally lethal hunters, who globally kill mesocarnivoes at 4.3 times higher rate than non-human predators (Darimont et al., 2015). It has therefore been suggested that they might be perceived by wild-living animals as ‘super- predators’ and presumably be far more frightening (Darimont et al., 2015; Clinchy et al., 2016). Recent playback experiments on badgers confirm that fearfulness of humans far exceeds that of other large carnivores (Clinchy et al., 2016). Hunting of the Pallas’s cat is prohibited across all range countries, apart from Mongolia (Ross et al., 2016), where permits might be obtained to hunt the species ‘for household purposes’ (Wingard & Odgerel, 2001). However, illegal hunting was found to occur frequently in central Mongolia (Murdoch et al., 2007; Murdoch et al., 2006) and illegal trade was

PICA - “Conservation of the Pallas’s cat through capacity building, research, and global planning” evident in local markets (Ross, 2009). Pallas’s cats in the current survey were detected mostly during the day, contrary to findings of mammals changing their diel activity towards increased nocturnality to minimise human encounters (Gaynor et al., 2018). This could suggest that the species is relatively tolerant of human presence- anecdotal evidence shows it is capable of inhabiting areas subject to heavy human disturbance (Webb et al., 2014). However, anthropogenic factors can also influence species interactions, with possible rippling implications throughout the ecosystem (Faeth et al., 2005). Sympatric species may shape their circadian activity patterns to avoid human disturbance, which can increase an overlap in their activity and thus potential for direct interactions- high levels of urbanization were found to increase opportunities for inter-specific interactions between pumas and bobcats (Lewis et al., 2015). Moreover, similarly to dogs, human activity can also destroy or mask scent marking signals, thus affecting intraguild communication between carnivores and potentially their coexistence (Lewis et al., 2015).

Intraguild pressures from predator species could also have contributed to the low detection rates of Pallas’s cat in the current study- it was suggested that the rarity of oncilla in Brazilian amazon could be explained by competitive intraguild interactions (de Oliveira et al., 2010). This phenomenon is also referred to as the ‘ocelot effect’, where small felid species have a lower density where the ocelot is present, and whose numbers only increase where the ocelot is absent or scarce (de Oliveira et al., 2010; de Oliveira & Pereira, 2004). Interspecific predation has been previously proposed as a likely reason for low density of Pallas’s cats, due to the restrictions it imposes on the species’ habitat use- complex habitats providing cover are preferentially selected, resulting in utilisation of only 20-30% of available landscape (Ross et al., 2016; Ross et al. 2012). High competing pressure from larger-bodied carnivores could therefore limit the abundance of Pallas’s cats in Tost Mountains.

To summarise- temporal segregation of activity is long recognised as one of the most effective mechanisms to reduce intraguild competition and promote carnivore co-existence (Massara et al., 2018; Glen and Dickman, 2005; Polis et al., 1989; Schoener, 1974) and predation pressure was proposed to determine numerous aspects of Pallas’s cat ecology (Ross et al., 2016). Observed patterns of Pallas’s cat activity seem indicative of temporal partitioning, whose degree could vary accordingly to differing lethal and/or non-lethal risk effects from sympatric predators. Current results also illustrate the complexity of intraguild interactions, which tends to increase with presence of multiple competing predators (Bischof et al., 2014). The exact dynamics of temporal interactions between the Pallas’s cat and sympatric predators, however, merit further investigation.

4.4. Temporal interactions of the Pallas’s cat and potential prey species:

Carnivores may coordinate their activity patterns with their most profitable prey in order to maximise their hunting success (MacArthur & Pianka, 1996; Foster et al., 2013; Rayan & Linkie, 2016; Steinmetz et al., 2013). Increasing body of research shows temporal overlaps of small felids with prey species- examples include Sunda clouded leopard (Ross et al., 2013), golden cat and clouded leopard (Haidir et al., 2018), ocelot (Porfirio et al., 2016), oncilla (Penido et al., 2017) northern tiger cat (Marinho et al., 2018), Iriomote cat (Schmidt et al., 2009) and guina (Delibes-Mateos et al., 2014). As both predators and prey, mesocarnivores need to constantly trade off-food and safety though (Cozzi et al., 2012; Prugh et al., 2009; Schoener, 1974; Penido et al. 2017) which may affect how they prioritise their (spatio)temporal strategies. Activity of Pallas’s cats in the current study was mostly diurnal, which could serve to facilitate co-existence with sympatric, primarily nocturnal predators. However, it also mirrored quite closely the activity of pika, which the species shows high preference for over other prey

PICA - “Conservation of the Pallas’s cat through capacity building, research, and global planning” items (Ross et al., 2010b) and which constitutes its optimal prey in terms of energetic gain due to its high biomass compared to other small mammals (Ross et al., 2010b); Pallas’s cats also do not appear to consume Siberian jerboas and Russian dwarf hamsters, both of which are nocturnal (Ross et al., 2010b), possibly emphasising its preference for diurnal prey. In addition, the distribution of Pallas’s cat strongly overlaps the distribution of pika (Ross, 2009; Ross et al., 2016), again possibly pointing to its reliance on this prey item. Therefore, Pallas’s cat activity pattern might reflect activity of its staple prey rather than regulation by intraguild predators, as was previously suggested for oncillas (Penido et al., 2017). Alternatively, predation pressure might exert a stronger influence on Pallas’s cats’ spatial behaviour, which was previously suggested to be primarily motivated by access to cover from predators rather than access to prey (Ross et al., 2012), while prey activity and availability might be more important in modulating the species’ temporal behaviour. However, this could only be established with further study on spatio-temporal dynamics of Pallas’s cat and sympatric species.

Some activity overlap with chukar partridge was also evident, which exhibited mainly diurnal activity, consistently with previous findings (Carmi-Winkler et al., 1987; Cramp & Simmons, 1979). The Pallas’s cat was observed to feed on birds, including partridge species, in both Mongolia (Ross et al., 2010b) and China (Wozencraft, 2008). Current results could therefore suggest that the partridge constitutes a larger bulk of Pallas’s cat diet than currently assumed. However, birds are generally minor components of the Pallas’s cat diet (Ross, 2009; Ross et al., 2010b) and thus it is more likely that the activity overlap between the species was in this case simply coincidental.

The incidence overlap of Pallas’s cat activity was lower for Argali and the lowest Tolai hare, possibly illustrating the order of their relative importance in diet of this felid. Predation of Pallas’s cats on Argali lambs seems to occur only occasionally during spring months (Reading et al., 2005; Murdoch et al., 2006) and the species’ scat analyses suggest that Tolai hare is rarely eaten (Ross et al., 2010b). Nonetheless, a temporal overlap of predator activity with prey does not necessarily mean that this species is key prey (Haidir et al., 2018) and thus future studies should investigate how activity of the Pallas’s cat corresponds with activity of various potential prey items.

The combined activity patterns of aforementioned and other potential prey species could also provide a continuous availability of mammalian prey in the area throughout the daily cycle, thus allowing the sympatric carnivores to segregate temporally whilst having access to prey (Monterroso et al., 2014; Monterroso et al., 2013).

5. Conclusion:

Current study aimed to use camera-trap derived data to quantify activity patterns of the Pallas’s cat and investigate its temporal interactions with sympatric species. Due to the low sampling effort and resulting lack of statistical analyses current data is insufficient to make definite conclusions about temporal activity of the Pallas’s cat. Nevertheless, the timings of Pallas’s cat camera-trap detections in the current study seem indicative of a predominantly diurnal activity pattern with crepuscular and nocturnal tendencies, potentially confirming the species as a mainly diurnal and crepuscular hunter. The seasonal patterns of Pallas’s cat temporal behaviour could suggest a suppression of activity during winter, as well as a degree of dietary flexibility, possibly promoting survival in a harsh climate of Mongolia during winter months. Activity patterns of the Pallas’s cat in relation to sympatric species also appear consistent with temporal strategies aimed at minimising competition and promoting

PICA - “Conservation of the Pallas’s cat through capacity building, research, and global planning” carnivore co-existence; however, activity and availability of prey may also play a role. Current data provides a preliminary insight into potential behavioural adaptations of the Pallas’s cat to the competing pressures in its environment, thus demonstrating its temporal plasticity, as well as enhances the overall body of knowledge about this elusive felid.

PICA - “Conservation of the Pallas’s cat through capacity building, research, and global planning”

GENERAL DISCUSSION

Current results should be interpreted with caution due to the low sampling effort of the Pallas’s cat (n=9). Probability of detection of a species is usually positively correlated with its abundance (Royle & Nichols, 2003), and thus current data could emphasise the rarity of the Pallas’s cat. Similarly to many other felids, Pallas’s cats are difficult to study due to their secretive habits, large home ranges and low densities (Porfirio et al., 2016; Ross et al., 2010a; Ross et al., 2016). Alternatively, elusiveness and rarity of the Pallas’s cat could signify expressions of ecological mechanisms, as was suggested for another small carnivore, the Altai mountain weasel (Bischof et al., 2014).

Nonetheless, non-detection cannot be directly considered absence (Li et al., 2010) and thus the low detection rate of the Pallas’s cat could simply reflect methodological limitations of using camera- trapping to study small mammals. Camera traps have been predominantly used to study large- and medium-sized animals, and thus they might be biased against detection of small, fast-moving species (Glen et al., 2013). Smaller-bodied (Kelly, 2008; Tobler et al., 2008) and more ambiguously-coated felid species are less likely to be detected by camera traps (Karanth et al., 2006). More importantly, the camera trap imagery analysed in the current study was originally collected for different purposes- to investigate abundance and population dynamics of the snow leopard (Sharma et al., 2014). Consequently, camera trap locations and heights were optimised for detection of the snow leopard, which may not necessarily have been optimal for detection of the Pallas’s cats. Small felids might also avoid travelling through pathways recently used by larger felids to minimize the risk of predation (Di Bitetti et al., 2010). Regardless of the exact reason behind it, the extremely low trapping rate of the Pallas’s cat over a period of three years (9 captures over 9724 operational camera days) emphasises the urgent need for further study of this elusive felid.

Observed relationships between the activity of Pallas’s cat and potential prey species may have also been affected by the body size detection bias- small mammals comprise a significant proportion of the Pallas’s cat diet (Ross, 2009; Ross et al., 2010b) and camera trap sensors in the current survey may not have favoured their detection (Porfirio et al., 2016). Populations of prey species living in colonies might also be patchy and separated irregularly by large distances (Murdoch et al., 2010), possibly further hindering their detection by camera traps.

It is also possible that observed patterns of temporal interactions between the Pallas’s cat and sympatric species are linked to differences in trapping rates, and thus more robust data set is needed to elucidate them further. Future studies could benefit from investigating spatial interactions of Pallas’s cats and potential predators as well- space is, along time, considered as the most important niche axis partitioned by wild-living carnivores (Di Bitetti et al., 2010). Distribution maps created by the ZSL software could provide a starting point for those investigations.

Similarly to other camera trap and observational studies, one of the main limitations of this study is that it draws inferences about biological processes simply from observed patterns derived from a remote device (Bischof et al., 2014; Haidir et al., 2018). Nonetheless, the value of using finer scale temporal data from camera traps to develop a deeper understanding of community dynamics, such as variations in activity patterns and partitioning along the temporal niche axis is increasingly appreciated in conservation research (Frey et al., 2017), as is its value in addressing ecological hypotheses linked to rarity and elusiveness of lesser-known small carnivores (Bischof et al., 2014).

PICA - “Conservation of the Pallas’s cat through capacity building, research, and global planning”

Finally, current results illustrate the benefits of collaboration between conservation organisations and sharing camera trap data across inter-species projects- the data originally collected by the Snow Leopard Trust for purposes of ecological study of snow leopard, a good proportion of which would have otherwise gone unused, also proved valuable in shedding some light on activity patterns and possible temporal adaptations of the Pallas’s cat. Moreover, the Tost Nature Reserve provides a promising setting for conducting longitudinal studies, which could produce more robust data sets in the future and effectively greatly enhance current understanding of this secretive, hitherto understudied species.

PICA - “Conservation of the Pallas’s cat through capacity building, research, and global planning”

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