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ISSN 1027-2992
CAT
news
Pallas's cat Status Review & Conservation Strategy
Special
Issue
I
I
N° 13 | Spring 2019
CATnews Special Issue 13 Spring 2019
02
CATnews is the newsletter of the Cat Specialist Group,
a component of the Species Survival Commission SSC of the
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lished twice a year, and is available to members and the Friends of
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Associate Editors: Tabea Lanz
Cover Photo: Camera trap picture of manul in the
Kotbas Hills, Kazakhstan, 20. July 2016
(Photo A. Barashkova, I Smelansky,
Sibecocenter)
The designation of the geographical entities in this publication, and the representation of the material, do not imply the expression of any
opinion whatsoever on the part of the IUCN concerning the legal status of any country, territory, or area, or its authorities, or concerning the
delimitation of its frontiers or boundaries.
Pallas's cat Status Review & Conservation Strategy
09
STEVEN ROSS1*, ANNA BARASHKOVA2, VADIM KIRILYUK3 AND SERGEY NAIDENKO4
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).
Origin
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
10
(+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
11
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
12
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).
Disease
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
13
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
*<steveross101@yahoo.co.uk>
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.
