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The behaviour and ecology of the manul

  • Sibecocenter, Novosibirsk
  • Daursky State Biosphere Reserve, Russia, Zabaykalsky kray

Abstract and Figures

Though widely distributed across the cold arid steppe and semi-desert ecosystems of Central and Western Asia, the manul is uncommon and rarely seen. The habitat in which it lives is demanding and highly seasonal; the manul exhibits morphological, physiological and behavioural adaptations that meet the challenges of temperature extremes, variable food resources and the risk of predation from other carnivores. This chapter describes the ecology of the manul, drawing from field studies and what we have learned from the captive zoo population. We end the chapter by ask-ing how can the manuls ecology aid our understanding of its conservation biology?
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ISSN 1027-2992
Pallas's cat Status Review & Conservation Strategy
N° 13 | Spring 2019
CATnews Special Issue 13 Spring 2019
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Cover Photo: Camera trap picture of manul in the
Kotbas Hills, Kazakhstan, 20. July 2016
(Photo A. Barashkova, I Smelansky,
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Pallas's cat Status Review & Conservation Strategy
The behaviour and ecology of
the manul
Head and Body length: 49 cm, 55 cm;
Tail length:and 25 cm; Weight: aver-
age 4 kg seasonally up to 5.0 kg, average
4.1 kg seasonally up to 5.3 kg.
The manul is approximately the size of a do-
mestic cat but with shorter legs and thick fur,
which accentuates its stocky appearance. On
average, they weigh 4 kg, but individuals can
weigh up to 5.3 kg at the end of summer when
prey are more common and easier to catch.
There is little sexual dimorphism, with males
only slightly larger than females (100–300 g;
Ross 2009, Naidenko et al. 2014).
The manul has a distinctive appearance. The
head is broad and flattened with the ears set
on the sides rather than the top of the head,
a trait thought to be an adaptation to aid
concealment when hunting in open habitats.
The forehead is marked with distinctive black
spots, and horizontal black and white stripes
run from the eyes to the cheeks on either side
of the face. Uncommon among the cats, the
manul’s pupils are round; in felids this cha-
racteristic is associated with diurnal hunters
and/or those found in open habitats (Malm-
ström & Kröger 2006).
The manul’s coat colour varies seasonally and
geographically. The species can be found with
a silvery grey, rufous grey and dark grey coat,
and a single individual may adopt all of these
coat colours over the space of one year (Ross
2009; Fig. 1). The manul has the densest fur
cover of all cat species inhabiting temporary
climatic zone (Heptner & Sludskii 1992). The
coat is often marked by faint black striping.
In winter the pelage is longer, denser and
lighter in colour than the summer coat, with
a pale, frosted appearance, providing warmth
and improving camouflage while there is
snow cover. The manul moults in the spring,
which often produces an intermediate rufous
coat before the thinner and darker summer
coat comes in (Ross 2009). In Iran sightings
of the manul with a rufous coat colour have
occurred year-round, suggesting the red
morph is specific to the region (Farhadinia et
al. 2016). The manul’s tail is distinctly band-
ed, with narrow stripes ending in a dark tip.
Coat colouration is supremely camouflaged
on a rocky steppe background. When motion-
less the manul resembles a small stone, or
blends onto the stone it is crouching upon. Its
background matching characteristics allow it
to vanish in rocky habitats (Ross 2009, Ross
et al. 2012; Fig. 2). In addition, the manul’s
white belly and under parts improve camou-
flage by balancing the effects of sunlight on
the top darker fur and self-shadowing on the
white underside, making it particularly diffi-
cult for aerial predators to spot while it is on
the move (Ruxton et al. 2004).
Although once included within the genus Fe-
lis, the manul is now classified as the sole
species in the genus Otocolobus. Its classifi-
cation is based on its unique morphology and
its distant genetic relationship to both the
Felis (wildcat) and Prionailurus (leopard cat)
lineages. Evidence suggests that Otocolobus
manul diverged from a common leopard cat
ancestor during the late Miocene approx-
imately 5.9 million years ago. The manul is
grouped within the leopard cat lineage along-
side the leopard cat Prionailurus bengalensis,
fishing cat P. viverrinus, flat-headed cat P.
planiceps and rusty-spotted cat P. rubiginosus
(O’Brien et al. 2008). Three subspecies of the
manul have been described, but only two of
these are said to be feasible based on geo-
graphic distribution (Kitchener et al. 2017).
Although not verified by molecular analysis
and not formally recognised, the subspecies
are known as Otocolobus manul manul and O.
m. nigripectus (Kitchener et al. 2017).
Distribution and Habitat
The manul has a wide but patchy distribu-
tion across Eurasia’s high altitude montane
grasslands and steppe, from western Iran to
eastern Mongolia. The largest populations
can be found in Mongolia, southern Siberia
and China, with the distribution becoming
progressively discontinuous further west. It
has been recorded in mountain steppe and
semi desert foothills in Kazakhstan and eas-
tern Kyrgyzstan. Populations in the southwest
of its range (the Caspian Sea region, Afghan-
istan, Pakistan and northern India) are dimin-
ishing, isolated and sparse. Recent records
from Bhutan, Nepal and Pakistan suggest
its occurrence across the Himalaya and Ka-
rakorum mountains, but despite large snow
leopard survey efforts they are rarely found in
this region (Chapters 3–5).
The manul’s range within the continental
climatic zone is characterised by aridity and
large variations in annual temperature. Tem-
perature range can reach over 100˚C, as re-
corded in Zabaikalskii krai, Russian Dauria
Fig. 1. A female manul in montane grassland/steppe habitat of Mongolia (Photo S. Ross).
Though widely distributed across the cold arid steppe and semi-desert ecosystems
of Central and Western Asia, the manul is uncommon and rarely seen. The habitat in
which it lives is demanding and highly seasonal; the manul exhibits morphological,
physiological and behavioural adaptations that meet the challenges of temperature
extremes, variable food resources and the risk of predation from other carnivores.
This chapter describes the ecology of the manul, drawing from field studies and
what we have learned from the captive zoo population. We end the chapter by ask-
ing how can the manuls ecology aid our understanding of its conservation biology?
chapter 2
CATnews Special Issue 13 Spring 2019
(+48˚C in summer to -53˚C in winter; S. Nai-
denko, pers. comm.). The manul’s habitat pre-
ferences influence the species’ distribution
within its range. Typical habitat consists of
montane grassland and shrub steppe (Fig. 1),
with a preference for areas with rocky out-
crops, ravines or other disruptive cover (Ross
2009), and within an altitude range of 450 to
5,593 m (Werhahn et al. 2018). The manul is
rarely found in very open habitats such as
short grassland and lowland sandy desert
basins, but when prey availability is very high
in open habitats it uses these habitats on a
temporary basis (V. Kirilyuk, pers. comm.). It
is also not found in areas where prolonged
snow cover exceeds 15–20 cm, for example
the manul’s north-eastern range is limited by
maximum snow depths of 16–17 cm in Trans-
baikal (Kirilyuk & Puzanski 1999). Due to their
selective use of habitats they remain patchily
distributed across their range (Heptner &
Sludski 1992, Ross 2009).
At the smaller scale, a major influence on the
manuls habitat usage is the constant risk of
predation by sympatric aerial and terrestrial
carnivores (Ross 2009). Predators of the manul
include large raptors, red foxes Vulpes vulpes,
the grey wolf Canis lupus and domestic dogs,
they are also hunted by humans (Barashkova
& Smelansky 2011, Ross et al. 2012). The ma-
nul is not a fast runner and when threatened
by other predators its best line of defence is
hiding out of sight, relying on their excellent
camouflage and taking cover in burrows (of
marmots or sympatric carnivores) or in rock
crevices (Fig. 2 & 3). In general, open areas
without suitable cover are avoided and ha-
bitats with disruptive cover such as ravines,
rocky areas, shrub-steppe and hill-slopes are
highly selected (Ross 2009, Ross et al. 2010a).
As a result, the manul uses only a small frac-
tion of habitats available within the steppe
ecosystem. Their habitat selection and spe-
cialisation is the most likely explanation for
their extremely low densities.
Manuls have a dependency on refuges or
dens. Dens are used on a daily basis to
provide important cover from predators, for
feeding, mating, giving birth, raising young,
and for thermoregulation during the extrem-
ely cold winters (Fig. 4). Den availability is
thought to be essential for manul survival,
and a critical habitat requirement for their
conservation (Ross 2009, Ross et al. 2010a).
In Mongolia they mostly use marmot burrows
in winter and rock crevices in the summer
(Ross et al. 2010a), in Southern Siberia and
Kazakhstan the den sites of sympatric carni-
vores are more commonly used (A. Barashko-
va, pers. comm.), and in Iran the manul has
been observed using aged Juniperus excelsa
tree cavities as breeding dens (Dibadj et al.
2018). Despite the range of habitats used by
the manul, the presence of suitable cavities
appears to be a standard niche requirement.
Feeding Ecology
The manul’s diet is mainly composed of small
lagomorphs and rodents. Pikas are the most
important prey across its range, typically
comprising over 50% of the diet and highly
selected over other prey species (Heptner &
Sludskii 1992, Ross et al. 2010b). As pika are
2–4 times larger than other common small
mammal prey, the manul’s preference for
them optimises hunting efficiency and ener-
gy intake. They also consume gerbils, voles,
hamsters and ground squirrels; less fre-
quently consumed prey includes small birds,
young marmots, hares, hedgehogs, reptiles
and invertebrates (Kirilyuk 1999, Ross et al.
2010b). Manuls have also been recorded eat-
ing berries (Kirilyuk 1999), scavenging from
carcases (Ross et al. 2010b), and predating on
a newborn argali sheep Ovis ammon (Read-
ing et al. 2005).
Hunting and their activity in general mostly
takes place at dawn and dusk in order to max-
imise the temporal overlap with prey while
minimising overlap with predators, such as
diurnal raptors or other competitors. Though
they may switch to a more diurnal rhythm
when temperatures are at their lowest (S.
Naidenko, pers. comm.). As a further mea-
sure to avoid predators, the manul mainly
hunts along the edges of rocky habitats and in
ravines which penetrate into open grasslands
and have high densities of pika, gerbils and
other small mammals. Long grass and thick
shrub are also used for cover when hunting
and moving through flat open grasslands in
the summer (Ross 2009).
Manuls hunt by three distinct techniques:
‘stalking’ by creeping very slowly and stealth-
ily around cover to locate and move close to
pounce on prey; ‘moving and flushing’, used
mainly in spring and summer by walking
quickly through long grass and undergrowth
to flush rodents, small birds, and grasshop-
pers which are then pounced upon; and
‘waiting in ambush’ where a manul waits
outside an active small mammal burrow for
the prey to emerge (Fig. 5), a technique used
mostly in winter to ambush pika (Ross 2009).
Following a successful kill, prey is routinely
taken into dens and burrows to consume in
safety in Mongolia (Ross 2009), but observa-
tions of eating prey at the capture site are
also common in Russia (S. Naidenko & V.
Kirilyuk, pers. comm.).
Movement, density and dispersal
Similar to most other cats, the manul is sol-
itary. Males do not help raise kittens and,
as a rule, they meet females only during the
mating season (Ross 2009). Males’ home
ranges encompass 1 to 4 female territories in
the typical polygynic system of solitary felids.
Research in Mongolia has shown that males
have highly overlapping ranges throughout
the year indicating little territoriality. How-
ever, aggressive encounters between males
do occur during the breeding season indicated
by fighting injuries during this time (Ross et
al. 2012) and suggesting that male territori-
ality is associated with the breeding season
and maiting rights. In contrast to male home
ranges, spatial overlap between females was
Fig. 2. A male manul showing its supreme ability to blend into a rocky background in
rocky habitat in Mongolia (cat in the centre; Photo S. Ross).
behaviour and ecology of the manul
Pallas's cat Status Review & Conservation Strategy
rare in Mongolia, but appears to be related to
their relatively small home ranges, low densi-
ty and the spacing of their preferred mountain-
ous/rocky outcrop habitat (Ross et al. 2012).
Several cues regulate the manul’s spatial
behaviour, they have been observed spraying
and cheek-rubbing (Mellen 1993), which pro-
vide temporal information for conspecifics.
The manul also effectively communicates
through vocalisations, making a strange call
sounding like a honking goose. The long-
distance calls and scent marking are likely
used by the manul for mate attraction and to
maintain spacing (Peters & Peters 2010).
Home range size is large in comparison to
other species of their size. In Mongolia, fe-
males use areas between 7.4−125.2 km2,
averaging 23.1 km2, compared to male home
ranges of 21−207 km2, averaging 98.8 km2
(Ross et al. 2012). Research has shown that
the availability and distribution of preferred
rocky habitats is one of the main stimuli af-
fecting home range size in Mongolia (Ross et
al. 2012). Home ranges appear markedly smal-
ler in Russian Dauria with male and female
home ranges averaging 27.4 km2 and 10.0 km2,
respectively (KIrilyuk & Barashkova 2011).
Density: There are no rigorous density esti-
mates for the manul, mainly due to their low
densities and cryptic behaviour resulting in dif-
ficulty in observing and surveying the species.
Ross (2009) estimated density using 3-years
of radio-telemetry data, surveys and observa-
tional data in what is considered prime habi-
tat in Mongolia. At 4–8 manuls/100 km2, the
cats occurred at extremely low density in com-
parison to other carnivores found in the area.
Much higher density estimates have been
found in Dauria, Russia, for example Naidenko
et al. (2014) captured a total of 16 manuls in
an area of 16 km2, equating to a density of
100/100 km2. Snow tracking in Russia has
also indicated that the manul can occur at
very high densities (Kirilyuk & Barashkova
2011, Barashkova et al. 2017). More research
is needed to understand regional differences
and temporal changes in manul density, but
presumably prey density and availability, and
predation pressure are the most influential
factors. Nevertheless, based the majority of
surveys and the scarcity of sightings across
the species range, evidence suggests that low
density/rarity is the more common state of
manul populations.
A number of factors may contribute towards
the manuls’ low density, including habitat spe-
cialisation, competition and prey availability.
The habitats selected by the manul only cover
10–30% of mountain steppe typically occu-
pied by the species, restricting the amount of
available habitat reduces potential density of
the species. Predation by other carnivores and
competition for scarce prey resources may
further constrain population density. For ex-
ample, in the Mongolian study area carnivore
density was measured using Distance Sam-
pling. Corsac fox Vulpes corsac density was
approximately 40–60 foxes/100 km2, red fox
density was 15–25 foxes/100 km2 and grey
wolf density was 3–20/100 km2. The area
also contained a high density of large raptors.
These predators constrain manul density, di-
rectly through predation, and indirectly by in-
fluencing the species habitat selection (Ross
2009). Prey density may also be influential,
as higher small mammal prey density should
provide better nutrition and improve kitten re-
cruitment and survival. High prey density may
also reduce predation pressure, as predators
focus on the more available and easily cap-
tured small mammal prey (e.g. Korpimäki &
Krebs 1996). These theories need to be tested
in a high density manul population, such as
those found in Dauria.
Dispersal: As is normal with all solitary ani-
mals, manuls disperse from their natal home
range after maturing. Data from Mongolia
indicate that this happens when the kittens
are approximately 4–5 months of age. Fol-
lowing emigration from their natal range,
sub-adults make exploratory movements
before settling and establishing their own
home range area about 5 to 12 km from their
natal home (Ross 2009).
Unusually large dispersal movements are
also commonly seen in adult manuls of both
sexes (Ross 2009). The sudden abandonment
of the home range and subsequent relocation
to a new area mostly occurs between August
and October. Individuals have been recorded
migrating a straight-line distance of 18 to
52 km, and journeys often entail crossing ha-
bitats that are not normally used. For exam-
ple, one adult male was observed making an
exploratory, looping excursion of 170 km over
2 months, requiring swimming across a large
river twice, before settling in a new area. The
high incidence of home range abandonment
(50% of adults, of 29 study cats) suggests
that it is an integral part of their ecology
(Ross 2009). Observations of large move-
ments have also been observed in Daurskii
Reserve, Russia (S. Naidenko, pers. comm.).
The motives for such moves are unclear, but
most likely include a process of disturbance
or prey depletion, where their home area be-
comes unviable, followed by emigration and
subsequent colonisation of a new ‘better’
area. Potential home-range disturbances may
include competition with other carnivores
resulting in displacement, or localised prey
depletion (Ross 2009).
Reproduction and demography
The manul lives in areas of the world sub-
ject to temperature extremes, thus it is un-
surprising that reproduction in the wild is
highly seasonal. In Mongolia, mating occurs
between December and March; this is the
only time of the year that females exhibit
ovarian activity (Brown et al. 2002). Male
sperm production also peaks during this time
and dramatically drops off at other times of
the year (Swanson et al. 1996). Experiments
Fig. 3. Pallas's cat showing its typical behaviour when threatened. It remains perfectly
still relying on its camouflage for protection (Photo S. Ross).
Ross et al.
CATnews Special Issue 13 Spring 2019
in captivity using different treatments of day-
light have found that the reproductive cycles
of manuls are entirely controlled by day length
(Brown et al. 2002).
During the mating period males pursue fe-
males to such an extent that it appears to take
precedence over hunting and feeding. Extreme
records have included males losing a total of
1,050 g (22%) over the course of only 14 days
during the mating period, and a second male
losing 800 g (19%) over 24 days (Ross 2009).
Weight loss over the course of winter is also
common in females, but most likely due to
the scarcity of prey (Ross 2009, Naidenko et
al. 2014). When females enter oestrus, males
‘shadow’ females for 2–3 days, protecting
her from advances by other males. Mating
appears to occur within marmot burrows or
other crevices, presumably to protect the cou-
ple from predators (Ross 2009).
Gestation is 66−75 days and litter size aver-
ages 3−4 kittens in captivity (Swanson 1999),
but females may give birth up to 8 kittens.
Kitten mortality in the wild is high with ap-
proximately 68% of kittens dying before dis-
persal. Surviving kittens reach independence
and disperse at 4−5 months. A radiotracking
study in Mongolia showed that sub-adult fe-
males may mate and reproduce at 10 months
of age (Ross 2009). Their reproductive lifespan
in captivity is approximately 9 years, but there
is a decrease in fecundity after 6 years and
very few females give birth after 8 years of
age (Barclay 2013).
The lifespan of the manul in the wild may be
up to 6 years, though they can survive up to 12
years in captivity. Predation is the main cause
of mortality in the wild. Most predation occurs
in winter, from January to April, when vege-
tation cover and prey density is low, increas-
ing their exposure to predators (Ross 2009). In
Mongolia large raptors accounted for 38% of
known deaths, while predation by domestic
dogs and hunting by people accounted for an
additional 53% of known mortalities, wolves
are also a known predator, and smaller car-
nivores such as badger Meles meles and red
fox occasionally kill manuls most likely on
a competitive basis (Ross 2009, V. Kirilyuk,
pers. comm). Mortality due to predation by
domestic dogs has also been recorded in Iran,
Russia and China, and appears to be a major
threat to the wild population (Ross 2009, Ba-
rashkova & Smelansky 2011, Farhadinia et al.
2016). In Mongolia survival data showed that
on reaching maturity at 1 year of age, adults
have approximately 50% chance of surviving
until 3 years (Ross 2009).
Captive manuls, particularly kittens, have a
unique and marked susceptibility to infectious
agents, especially Toxoplasma gondii. The
manul is suspected to be naïve and suscep-
tible to the agent due to lower occurrence of
toxoplasma in the wild. Though 2 cases of T.
gondii antibodies were found in manul popu-
lations in the Chita region of Russia and cen-
tral Mongolia (Brown et al. 2005, Naidenko et
al. 2014). Naidenko et al. (2014) also recorded
antibodies to Mycoplasma, Influenza A virus
and Feline leukaemia virus in a sample of
16 cats. The manul is also exposed to feline
immunodeficiency virus FIV in the wild. This
virus does not cause death but is related to
immune depletion. Interestingly the manul
harbours a unique strain of the virus most
closely related to the African cheetah and
leopard FIV strains (Brown et al. 2010).
The conservation biology of the manul
The manul has a very wide range across cen-
tral and western Asia, and because of this the
population is very unlikely to go extinct in the
short term. However, of more concern is loca-
lised and regional extinction, as the manul’s
ecology naturally disposes them to threats
(Chapter 8).
The manul is a naturally rare species, they
are dependent on specific habitats and prey,
and are easily killed on open ground. As the
manul is a habitat specialist this is likely to
result in increased vulnerability to the effects
of habitat fragmentation and degradation. Its
large home-ranges increase the probability
that their ranges will overlap with human acti-
vities, disturbances and associated mortality,
and be more difficult to cover by protected
areas. For the manul, the availability of bur-
rows, rock crevices and other cavities is ne-
cessary, as these are critical resources, used
on a daily basis and essential for breeding.
Fig. 4. A rock crevice den-site with manul kittens (top) and a marmot burrow den (bot-
tom). Dens are used on a daily basis by the manul and are essential for raising young
(Photo S. Ross).
behaviour and ecology of the manul
Pallas's cat Status Review & Conservation Strategy
This dependency on burrows means that the
decline of burrowing species such as marmot
and small carnivores poses a threat to the
manul (Ross et al. 2016). Overall, land use
changes across the manul’s range are increas-
ing due to habitat destruction and fragmenta-
tion, declines in their prey base, and a rise in
mortality associated with increased contact
with herders and their dogs (Chapter 8).
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1 Office for Conservation of the Environment, Di-
wan of Royal Court, Muscat, Oman
2 Sibecocenter, Novosibirsk, Russia
3 Daursky State Nature Biosphere Reserve, Kom-
somolskaya str., Nizhnii Tsasuchei, Ononskii
District, Zabaikalskii Krai, Russia
4 Severtsov Institute of Ecology and Evolution,
Russian Academy of Sciences, Leninsky, Mos-
cow, Russia
Ross et al.
... It is currently listed as 'Least Concern' in the IUCN Red List of Threatened Species TM (Ross et al. 2020) and is the only living species within the Genus Otocolobus (Kitchener et al. 2017). Similarly to other small cats (Macdonald and Loveridge 2010;Anile and Devillard 2016), the Pallas's cat has received very little attention from the scientific community (Lanz et al. 2019), likely because the low density and the remote areas typically inhabited by this species (Ross et al. 2019a) make scientific study particularly challenging (Hunter 2015). As a result, few studies have been conducted on Pallas's cats; thus, its ecology and status are still poorly known (Lanz et al. 2019). ...
... As a result, few studies have been conducted on Pallas's cats; thus, its ecology and status are still poorly known (Lanz et al. 2019). The main threats to Pallas's cats are illegal poaching and habitat loss (Ross et al. 2019a), with this latter threat being further exacerbated by livestock grazing. Livestock presence can have a negative impact on Pallas's cats because of exploitation of pastures, hence reducing availability of primary prey items such as lagomorphs and rodents (Ross et al. 2019b). ...
... Livestock presence can have a negative impact on Pallas's cats because of exploitation of pastures, hence reducing availability of primary prey items such as lagomorphs and rodents (Ross et al. 2019b). In addition, herding dogs and human killings represent the major causes of mortality of Pallas's cats (Hunter 2015;Ross et al. 2019a). Fur trade, especially in Mongolia, is still considered a major threat because this species is legally hunted for domestic use (Hunter 2015). ...
Context. The ranges of many small, at-risk felid species occur almost entirely in unprotected areas, where research efforts are minimal; hence data on their density and activity patterns are scare. Aims. We estimated density and activity patterns of Pallas’s cats on unprotected lands in central Mongolia during two periods (May–August and September–November) in 2019. Methods. We used spatially explicit capture–recapture models to estimate population density at 15.2±4.8 individuals per 100 km2. Key results. We obtained 484 Pallas’s cat images from 153 detections during 4266 camera-days. We identified Pallas’s cats using pelage markings and identified 16 individuals from 64 detections. Pallas’s cat activity was consistent between the two survey periods (~0.50), with cats mainly active during crepuscular hours in the first period and strictly diurnal in the second. Conclusions. We provide the first estimation of a Pallas’s cat population density using camera-trapping. Compared with other methods used, densities were high in our study area, which was likely to be due to a combination of highly suitable habitat and abundant prey. Seasonal shifts in the activity patterns of Pallas’s cats indicated a likely adaptive response to reduced risk of depredation by raptors. Implications. We recommend August to November as the best time for camera-trapping surveys for Pallas’s cats, given their high daily activity and the easiest interpretation of images used for individual identification collected during this time. We also suggest that future camera-trapping surveys of Pallas’s cat be mindful of potential camera-trap avoidance through time.
... Focal area models included a relative abundance index (RAI) of domestic livestock, dogs, and humans (domestics), and of prey species indicate that this area is suboptimal for this felid, likely because it is generally higher in elevation and has more extensive glacial and snow cover relative to the other areas (Ganyushkin et al. 2018). The Pallas's cat appears to prefer areas that are steep, with natural vegetation and higher occurrence of prey, matching the high habitat and dietary specialization known for this species (Ross et al. 2019a, Chimed et al. 2021, Greenspan and Giordano 2021. ...
... When considering that livestock numbers have markedly increased across Mongolia over the last decades (Berger et al. 2013, Pfeiffer et al. 2018, and that we also recorded diffused livestock presence into protected areas (this study, Augugliaro et al. 2020, our findings are of conservation concern for a number of reasons. First, the Pallas's cat is specialized in its habitat choice, and only uses a small fraction of area available within the steppe ecosystem (10-30%; Ross et al. 2019a); it is dependent on areas that are protected but also increasingly used for grazing, which may eventually threaten the cat from habitat degradation affecting the abundance of small rodents, hence progressively reducing prey availability (Cao et al. 2016, Schieltz andRubenstein 2016). Second, the proximity of preferred areas to herders' houses and camps implies higher chances of Pallas's cat predation by dogs, considered one of the most important causes of human-related deaths of these cats (Ross 2009;Barashkova 2012Barashkova , 2017, and poisoning, directly as a predator control method or as consequence of poisoning of Pallas's cat's primary prey for pest or disease control (Tseveenmyadag and Nyambayar 2002). ...
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Biased research and conservation efforts result in some faunal groups (e.g., small felids) being understudied, and hence these groups are often declining without adequate knowledge to manage for threat reduction. The Pallas's cat (Otocolobus manul) occurs across central and western Asia with declining populations and the largest population is likely in Mongolia. A potential threat to this felid is livestock encroachment across its range, including within protected areas, yet we lack a clear understanding of the impact of livestock husbandry on this cat. We used motion-sensitive camera data from 216 sites in 4 study areas in western Mongolia to study the occurrence probability of Pallas's cat in relation to habitat characteristics and occurrence of livestock, and conducted a local assessment within a strictly protected area where we obtained the highest number of detections. We estimated a relatively low occupancy (0.33 ± 0.10), which is associated with sites with natural vegetation, steeper slopes, and greater prey abundance. Occupancy also increased with increasing livestock occurrence, particularly large herds of sheep and goats. Such co-occurrence was partially adjusted by diel activity segregation, presumably to limit direct encounters. Our results suggest that the preferred habitat by Pallas's cat in the study region coincides with areas encroached by livestock. The Pallas's cat's habitat is specialized and its dependence on areas that are increasingly used for grazing may eventually threaten the cat with habitat degradation, prey depletion, predation by dogs, and poisoning from pest control. Relevant conservation actions should regulate livestock encroachment within protected areas and improve grazing regimes. The Pallas's cat is an indicator species of mountainous and steppe ecosystems in central Asia; hence, further research towards the preservation of its populations would also benefit other key species across its range.
... The high occurrence of pikas in Pallas's cats' diet is not surprising, since pikas (Ochotona spp.) in general often occur in locally high densities on the arid steppe of Russia, Mongolia, and China (Smith et al., 1990). Somewhat surprisingly, researchers also found that Pallas's cats less frequently consume berries and scavenge from carcasses (Ross et al., 2019;Reading et al., 2009). ...
Pallas's cats range across Central Asia and commonly consume small mammals. Most studies on Pallas's cats' diets found that they strongly selected particular prey species, such as pika (Ochotona spp.). Pika occurrence in Pallas's cat diet was disproportionately high compared to availability in Central Mongolia. We conducted research into Pallas's cat ecology in Khalzan Soum, Sukhbaatar Aimag since 2018. Brandt's vole density started to increase in 2017, peaked in 2018–2019, and decreased in 2020. We collected feces of Pallas's cats every month from May 2019 to May 2020. During this time, the monthly percent occurrence of Brandt's voles in Pallas's cats' diet ranged from 82.6 to 89.9%. We found a significant correlation between rodent abundance and occurrence in scats of Pallas's cats during the kitten rearing season. Brandt's vole (Lasiopodomys brandtii), Daurian ground squirrel (Ochotona dauurica) and Mongolian hamster (Allocricetulus curtatus) were significantly preferred. Our research provides insights into how Pallas's cats play an important role in biological control in this ecosystem during a Brandt's vole's population irruption.
Carnivores play key roles in maintaining ecosystem structure and function as well as ecological processes, understanding how sympatric species coexistence mechanism in natural ecosystems is a central research topic in community ecology and biodiversity conservation. In this study, we explored intra- and interspecific niche partitioning along spatial, temporal, and dietary niche partitioning between apex carnivores (wolf Canis lupus , snow leopard Panthera uncia , Eurasian lynx Lynx lynx ) and mesocarnivores (Pallas’s cat Otocolobus manul , red fox Vulpes vulpes , Tibetan fox Vulpes ferrilata ) in the Qilian Mountain national park of China using camera trapping data and DNA metabarcoding sequencing data. Our study showed that apex carnivore species had more overlap temporally (the value of time overlap from 0.661 to 0.900) or trophically (the value of diet overlap from 0.458 to 0.892), mesocarnivore species had high dietary overlap with each other (the value of diet overlap from 0.945 to 0.997), and apex carnivore and mesocarnivore species had high temporal overlap (the value of time overlap from 0.497 to 0.855). Large dietary overlap was observed between wolf and snow leopard (Pianka’s index=0.892) and Pallas’s cat and Tibetan fox (Pianka’s index=0.997) and suggesting increased resource competition for these pair species were existed. We conclude that spatial niche partitioning playing a key role in facilitating the coexistence of apex carnivore species, spatial and temporal niche partitioning facilitate the coexistence of mesocarnivore species, and spatial and dietary niche partitioning facilitate the coexistence between apex and mesocarnivore species. Our findings address, for the first time, niche partitioning was considered across temporal, spatial and dietary dimensions and diverse coexistence patterns of carnivore species were presented in the Qilian Mountain national park of China. These finding will contribute substantially to current understanding of carnivore guilds and effective conservation management in fragile alpine ecosystems.
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Olfactory communication is important for many solitary carnivores to delineate territories and communicate with potential mates and competitors. Pallas’s cats (Otocolobus manul) are small felids with little published research on their ecology and behaviour, including if they avoid or change behaviours due to dominant carnivores. We studied their olfactory communication and visitation at scent-marking sites using camera traps in two study areas in Mongolia. We documented four types of olfactory communication behaviours, and olfaction (sniffing) was the most frequent. Pallas’s cats used olfactory communication most frequently at sites that were not visited by snow leopards (Panthera uncia) and when they used communal scent-marking sites, they were more likely to use olfactory communication when a longer time had elapsed since the last visit by a snow leopard. This suggests that Pallas’s cats may reduce advertising their presence in response to occurrence of snow leopards, possibly to limit predation risk.
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Identifying the association between the patterns of niche occupation and phylogenetic relationships among sister clades and assisting conservation planning implications are of the most important applications of species distribution models (SDMs). However, most studies have been carried out regardless of within taxon genetic differentiation and the potential of local adaptation occurring within the species level. The Pallas's cat (Otocolobus manul) is a less‐studied species with unknown biogeography and phylogenetic structure across a widespread yet isolated range from the Caucasus to eastern China. In the current study, by considering a previously proposed genetic structure and based on a cluster analysis on climatic variables, we supposed three clades for this species, including O. m. manul, O. m. ferrugineus, and O. m. nigripectus. We developed SDM for each clade separately and compared it with a general distribution model of the species to determine whether the hypothesized taxonomic resolution affects the predicted ecological niche of the within‐species structures. We assessed the effect of climate change on the future distribution of the species to detect the most sensitive clades to global warming scenarios. Our results showed that for all clades' models, the AUC and TSS were greater than the general model. Access to the preferred prey of the Pallas's cat, that is, pika, had a significant effect on the distribution of O. m. manul and O. m. ferrugineus, whereas the most influential variable affecting O. m. nigripectus habitat suitability was terrain slope. Based on our future projections, we found that future climate change likely threatens the clades O. m. ferrugineus and O. m. nigripectus more than O. m. manul, findings that were hidden in the general model. Our results highlight the proficiency of SDMs in recognizing within‐taxon habitat use of widespread species and the necessity of this procedure for implementing effective conservation planning of these species. The Pallas's cat with a widespread but isolated distribution shows a divergent habitat selection across Eurasian cold steppes. A sub‐taxon climate change projection revealed the different responses of the Pallas's cat subspecies to future global warming.
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The fishing cat’s persistence in a ‘semi-aquatic niche’ suggests the evolution of a successful hunting strategy. We describe it for the first time by analysing 197 camera-trap video-clips, collected from a participatory-science initiative, within an ethogram framework . The cats spent ∼52% of the time sitting and waiting for prey (fishes) to come nearer and took limited attempts to hunt (3.89%) in deeper waters (in which the upper portions of the cat’s body were submerged), where its hunting success was found to be 42.86%. In shallow waters, it adopted a predominantly active mode of hunting (∼96%) to flush out prey.
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Pallas’s cat is a rare felid that has a wide but patchy distribution across Eurasia’s cold steppes. The species is known to prefer rocky, mountain and shrub steppe habitats, and strongly selects habitats with good cover from sympatric predators, particularly when raising young. This is the first report of the species using Juniperus excelsa woodland habitat for foraging and breeding in Iran. We observed the use of two aged juniper tree cavities as breeding dens for a litter of four kittens. The record extends our capacity to assess the range of habitats that are used and suitable for Pallas’s cat populations.
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1. The current classification of the Felidae was reviewed by a panel of 22 experts divided into core, expert and review groups, which make up the Cat Classification Task Force CCTF of the IUCN Cat Specialist Group. 2. The principal aim of the CCTF was to produce a consensus on a revised classification of the Felidae for use by the IUCN. 3. Based on current published research, the CCTF has fully revised the classification of the Felidae at the level of genus, species and subspecies. 4. A novel traffic-light system was developed to indicate certainty of each taxon based on morphological, molecular, biogeographical and other evidence. A concordance of good evidence in the three principal categories was required to strongly support the acceptance of a taxon. 5. Where disagreements exist among members of the CCTF, these have been highlighted in the accounts for each species. Only further research will be able to answer the potential conflicts in existing data. 6. A total of 14 genera, 41 species and 77 subspecies is recognised by most members of the CCTF, which is a considerable change from the classification proposed by Wozencraft (2005), the last major revision of the Felidae. 7. Future areas of taxonomic research have been highlighted in order to answer current areas of uncertainty. 8. This classification of the Felidae will be reviewed every five years unless a major new piece of research requires a more rapid revision for the conservation benefit of felid species at risk of extinction.
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The Pallas's cat, or Manul, Otocolobus manul, belongs to the small felines. In Russia this species is located at the northern periphery of its range. Potential habitats of this species within its Russian range's part have been highlighted on the base of remote sensing data, topographic maps and digital cartographic models. Maps of the Pallas's cat's records in Russia have been compiled using literature data, authors' own data and information provided by contributors of the online database «Small Wild Сats of Eurasia». We have shown the representation of potential habitats of the Pallas's cat and its actual registrations within Protected Areas of Russia. The total area of potential habitats for this species in Russia is assessed as 145,150 km2. The presence of Manul is confirmed for five regions of Russia: Republic of Altai, Republic of Tyva, Republic of Buryatia, Krasnoyarsk Krai, Trans-Baikal Krai. The registration of the species' records mainly corresponds to the location of its potential habitats. The main categories of both federal and regional Russian Protected Areas cover 10.8% (13.5% taken with the buffer zones) of the total area of potential habitats for the Pallas's cat. The significance of each Protected Area for Manul conservation has been considered in detail. We have discussed whether the representativeness of the existing Protected Areas is sufficient for conservation of this species. We have identified the main territorial gaps that reduce the Pallas's cat protection in some parts of the Russian range.
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Pallas’s cat is a 3-5kg species endemic to Central Asia. It is Near Threatened and thought to be at risk from habitat fragmentation, a declining prey base and consumptive hunting. Moreover, it is distributed within the temperate grasslands biome, the least protected of all biomes in the world and increasingly under pressure from economic development and population growth. While the current status of Pallas’s cat highlights its need for protection, its ecology has not been studied, impairing the development of conservation strategies. In this thesis I address the need for an ecological framework to support the species conservation. I conducted fieldwork for 30 months from 2005 to 2007. Pallas’s cats were found to maintain large home range areas and live at very low densities. They are polygamous but appear to be non-territorial. The distribution of preferred habitats and sex were the main factors in home range size variation. Pallas’s cats are highly selective feeders, preferring pika over other available prey species. As pikas are 2-3 times larger than other prey species, they may be an optimal food item which reduces foraging costs per unit energy gain. Pika eradication programmes in China and Mongolia therefore pose a serious threat to Pallas’s cats’ prey base. Pallas’s cat is also a habitat specialist. They largely confine activities in or near to rocky and ravine habitats. Both habitats were found to provide superior hiding and escape cover, indicating specialisation is an anti-predator strategy, as used habitats had comparatively low prey availability. Marmot burrows and rock crevices were used as dens on a daily basis for most aspects of Pallas’s cats’ life; similar habitats were selected for denning as for foraging. Overall, the dominant influence on Pallas’s cats’ ecology was its susceptibility to predation. Anti-predator behaviour shaped all aspects of Pallas’s cat’s ecology including its habitat use, spatial behaviour, activity and where it rested, reproduced and fed. Pallas’s cats have several ecological traits commonly associated with extinction vulnerability, painting a bleak picture for their future conservation. Considering the diverse changes that are currently sweeping through the steppe ecosystem, significant disturbance of Pallas’s cat habitat will continue to be an issue.
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Iran is most likely the western boundary of the Pallas’s cat’s, or manul Otocolobus manul global distribution range. The Pallas’s cat is amongst the least-studied felids in Iran and basic questions about its status and natural history have yet to be answered. Our review of the available information suggests significant increases in the range of the species previously known from Iran. North-eastern Iran remains a hotspot of Pallas’s cat occurrence in the country, but there are a growing number of recent confirmed records from southern slopes of Alborz Mountains, as well as the south-central provinces. Human disturbances such as mining activities and traditional pastoralism, particularly during summer when alpine and sub-alpine rangelands are occupied by flocks of livestock, might have adverse impact on the Pallas’s cat. The lack of scientific understanding of the Pallas’s cat in Iran restricts our ability to conserve the species.
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Ecologists have long sought to identify environmental and ecological traits influencing space use by individuals. Prey availability, habitat type, conspecific interactions, and sex are cited as determinants of carnivore spatial behavior, although empirical evidence of relationships between variables and home-range size are rare. We examined the relative importance of different ecological factors on the spatial behavior of Pallas’s cats (Otocolobus manul), mesocarnivores native to the montane steppes of central Asia. Between 2005 and 2007 we estimated home-range size for 9 male and 16 female Pallas’s cats. Cats used large and variable home ranges, with male home ranges 4–5 times the size of female ranges. Contrary to predictions, home-range size did not increase in response to low prey availability or seasonality. Smaller home ranges were associated with higher coverage of preferred rocky habitats in the home-range center, whereas larger home ranges were associated with higher connectivity of rocky habitats in the periphery of home ranges. This suggests that space use by Pallas’s cats is a function of sex and is mediated by the distribution and availability of habitats. Because of interspecific predation pressure experienced by Pallas’s cats, we argue that their spatial behavior originates from trade-offs between accessing food and maintaining proximity to habitats that provide cover from predators.
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Dietary plasticity and diet composition of Pallas's cat (Otocolobus manul) was quantified in central Mongolia. Diet of Pallas's cat was assessed by scat analyses, and prey surveys were used to estimate prey availability. Prey selection was calculated using multinomial likelihood ratio tests. Analysis of 146 scats identified 249 prey items. Pallas's cats ate a broad range of small mammals, insects, birds, reptiles, and carrion, but Daurian pikas (Ochotona dauurica) were the most frequently consumed prey. Multinomial likelihood ratio tests indicated nonrandom selection of prey species. Pikas were selected disproportionately to their availability, and other more numerous prey items were used less than expected, indicating feeding specialization. Specialization on pikas appears to optimize energy intake per unit foraging by Pallas's cats, because pikas are 2–4 times larger than other small mammal prey. We argue that pika control programs in China and Mongolia potentially threaten Pallas's cat populations, because no other prey species could fill the niche occupied by the pika in terms of size and year-round availability.
A male Pallas' cat (Felis manul) was housed under natural lighting conditions in an outdoor pen for 20 mo and subjected to bimonthly reproductive evaluations consisting of blood sampling for hormonal analysis, testicular volume and body weight measurement, and electroejaculation. Distinct seasonal reproductive patterns were identified; sperm production and quality in the breeding season (December-April) was significantly higher (P < 0.01) than that in the nonbreeding season (June-October). Testicular volume did not differ (P > 0.05) between seasons, but body weight gain and loss occurred (P < 0.01) 1-2 mo before the breeding and nonbreeding seasons, respectively. Although serum testosterone concentrations were similar (P > 0.05) in both seasons, serum luteinizing hormone (LH) concentrations were greater (P < 0.01) during the breeding season, and sperm concentration, total number of sperm per ejaculate, percentage of morphologically normal sperm, sperm motility, and serum LH concentration were correlated (r = 0.60-0.99; P < 0.05). The male Pallas' cat exhibits a pronounced reproductive seasonality, which has important implications for captive breeding management and genome resource banking.
Abstract We measured seasonal changes in body mass and pathogen exposure in wild Pallas' cats (Felis [Otocolobus] manul) in the Daurian Steppe of Russia in 2010-11. Pallas' cats lost about 30% of body mass over winter. Tests for antibodies to 15 potential pathogens showed that Pallas' cats were exposed to four pathogens. Two of 16 cats had antibodies to Toxoplasma gondii. Two had antibodies to Mycoplasma sp., and one each had antibodies to Influenza A virus and Feline leukemia virus. The percentage of antibody-positive wild Pallas' cats was lower than results reported for other wild felids in the Russian Far East.