>>> german version

PAAZAB – Annual Conference 2007

Title of paper: Karoo Cat Research – Introduction of an Institution

Author: Mircea Pfleiderer

Institution and address: Karoo Cat Research, Honingkrantz, Fish River 5883, South Africa

Tel.: 048 881 3907

Cell: 076 306 2343

e-mail: felis@isat.co.za


“Karoo Cat Research” is the name for a long-term project which began as a field work (started in 1993 in the Mountain Zebra National Park, continued in a private nature reserve and on some farms in the districts of Cradock, Beaufort-West, Steytlerville and Belmont). “Karoo Cat Research” is also the name of a private research facility in Fish River (founded in 2001), where we presently keep, in large, nature-type enclosures, three Karoo Cat Species: The Blackfooted or Small Spotted Cat (Felis nigripes), the African Wild Cat (Felis libyca) and the Caracal (Profelis caracal), and, additionally, one Bushveld Cat Species: The Serval (Leptailurus serval). Karoo Cat Research is based at Honingkrantz Farm in the eastern arid Karoo (350 mm/y), at an altitude of 1100 m. 2004 the institution became a supporting member of the Cat Conservation Trust, Cradock.

The main aims of Karoo Cat Research are:

  1. Research

The research project on South African Felidae is not solely based on observing the cats in captivity; it rests so to say on two pillars: Field investigations and, to have the chance of a closer look, observations on cats in captivity. The main research subjects are: Social and hunting/feeding behaviour, maternal behaviour and ontogenesis, comparative ethology (comparison of some special part of the behaviour of two or more species), ecology, systematics and taxonomy, zoo biology.

  1. Conservation

With the founding of many new nature reserves and the enlargements of National Parks the rare as well as the more common cat species are in high demand for re-introduction. The breeding program includes all above mentioned species. All the litters are mother reared and get special training for the survival in the wild, which up to now proved 100% successful. We also provide zoos with specimens of the rarer felid species.

  1. Education:

We invite local farmers, tourists and children for lectures, cat viewing and discussions, emphasising the positive ecological impact of feline predators. We also accommodate and supervise students from South African and foreign universities, and give practical advice for animal psychologists and zoo animal keepers.

African Wild Cat (Felis libyca). On the Significance of Captive Breeding.

Mircea and Jörg Pfleiderer, Karoo Cat Research, Cradock, South Africa

The African Wild Cat (Felis libyca) is not only an attractive species and quite useful in controlling small rodents. It is also a highly remarkable study and research object. The topic of finding out how and to which extend the domestication influenced and changed the behaviour of Felis libyca, especially the social behaviour, is one example, the endeavour to establish a thorough ethogram of the wild species is another.

The IUCN Cat Specialist Group regards the African Wild Cat as very endangered even though the species was not listed in 1998. While the hunting of it is prohibited in the North African countries, in Israel, Nigeria and Mozambique, and regulated in some Western African countries, the African Wild Cat gets no protection over the greater part of its range, including South Africa. This paper is a plea to all zoos and animal enterprises to help saving this animal from vanishing completely.

The species presents a very unique case of endangerment as it is still not so much threatened by extermination or by loss of habitat as most other species in danger of extinction. It is the constant, widespread hybridisation with the domestic cat that invades all natural habitats of the African Wild Cat with the ever-expanding human population which makes it increasingly difficult to find pure African Wild Cats. The domestic cat or house cat, Felis libyca forma catus, derives from up to four Egyptian and near-eastern subspecies: libyca, lowei, lynesi, ocreata, i.e., it is the domesticated form of those subspecies, and is, thus, still sufficiently closely related to the sub-Saharan subspecies (brockmanni, foxi, haussa, mellandi, taitae, tristrami, ugandae) including the Southern African ones (cafra, griselda) that interbreeding yields fertile offspring.

The hybrids, being partly domesticated even when living in the wild, have a general tendency to be slightly less afraid of humans. This unfortunately means that in the vicinity of human settlements – which today means practically everywhere – the hybrids have a slightly higher survival and reproduction chance than the true Wild Cats. Hybrids with a substantial portion of domestic cat hereditament are also less prone to die of contagious diseases like panleukopenia, because domestic cats are more resistant to this kind of infection than any of the other Felis species, which severely aggravates the situation.

The problem of hybridisation does of course not only affect the Southern African Wild Cat subspecies, which at least both are, on average, stronger and larger ( 5kg, 4kg, Stuart 1981) than their domesticated kin. Other countries that habituate this species are even more severely struck by the problem. Many of these countries are not sufficiently developed to be concerned about the conservation of smaller mammals. Countries in the Middle East, like e.g. Israel, are well alarmed about the state of their Wild Cat population, but the period of time where hybridisation did occur is much longer in this particular region where the domestication of the African Wild Cat has taken place about 4000-5000 years ago. Another problem especially of the Middle East is that the local subspecies are smaller and of more delicate built than their domesticated competitors (Mendelssohn 1989).

What can be done? The usual measures should not be neglected: Decrease extermination and loss of habitat. Unfortunately, some farmers and many bird enthusiasts still regard the African Wild Cat as “vermin”. The cats can indeed occasionally cause some damage in ostrich breeding facilities or poultry farms, they also might kill the odd lamb, but this is by far surpassed by its positive impact. The African Wild Cat very efficiently controls the food competitors of life stock – smaller and middle sized rodents – that cause a vast damage most farmers are not quite aware of. Scat analyses as well as stomach contents of destroyed animals proved invariably the African Wild Cat’s food specialisation on rodents (80-90%). In spite of this, many African Wild Cats have been killed in the past. In this respect, persuasion of farmers is as necessary as education of the general public, the latter being done most effectively in zoos.

With the development of a new conservation awareness in South Africa, the amount and size of nature-reserve-type facilities that can be kept relatively free of domestic cats are increasing. These are places where the re-introduction of pure African Wild Cats seems to be awarding.

We would like to stress the point that it is not easy to find pure-bred African Wild Cats in the Veld. 68% (n =23) of the cats we found on farms and nature reserves around Cradock in the years between 1995 to 2005 showed signs of hybridisation in fur colouring as well as in habitus proportions. This result is better than those of most other regions. Smithers, for example, claimed, in 1986, the impossibility of finding pure African Wild Cats anywhere near human settlements.

The situation concerning certain breeding programs, even those of some well-established zoos in South Africa is only slightly better. Nobody really knows how many hybridised African Wild Cats are kept in captivity and are being bred with. The only feasible remedy is controlled breeding, which can best be done in zoos and similar enterprises.

The African Wild Cat, being rather fertile (average number of offspring per year: 3.5) can produce even more and larger litters in captivity (up to 5 kittens, 2 litters/year). There should be no difficulty in obtaining enough pure breeding strains for several purposes, as exhibition, research, even hand-raised pets, and, particularly, offspring especially reared for re-release.

The first task is, obviously, to start with pure-bred specimens. While it was not possible in the past years to provide breeders with reliable genetic markers (actually up to now it is claimed that there is no known genome/markers for this species), there are some new developments coming up. Still, the so very close relationship of the African Wild Cat to its own domesticated form will always render such methods extremely difficult.  One kind of test that we are aware of is done by Dr. Hartley in Cape Town. This test rates, on a scale, how far removed the African Wild Cat specimen will be from the genome/marker of a domestic cat.

We still have to wait for new developments to prove the validity of their results. Some researchers argue the case for the subspecies in our region, so we presently have to rely on our eyesight and some rather crude measurements to separate the pure bred specimens from hybrids. Those methods may look to some scientists quite primitive and “untechnical”, but they prove surprisingly trustworthy.

Features of the African Wild Cat in comparison to hybrids.

Proportions: The African Wild Cats of the South African sub region are on average slightly larger than the domestic cat, but of more slender – never dainty – built, with longer legs than even those of the oriental domestic-cat breeds. This gives the impression of a very “upright” posture when the cat is sitting. The back legs are visibly longer than the front legs. Due to this fact, the tail appears shorter than that of a domestic cat as it barely covers the heels. The short, wedge-shaped but not too delicate head, which is set on a long, slender neck, shows in profile a slight rise from the straight nose bridge to the forehead. The ears are large, broad-based and upright, with moderately pointed tips that may have a small tuft. The light yellow or greenish eyes are strikingly large.

While many hybrids can be distinguished by the rounded skull, fuller cheeks, shorter nose, a comparatively small nose leather and a more sturdy body, this can be a difficult way of detecting domestic influence if it is less than 1/8. Further signs for hybridisation are: a massive, short-legged body, stocky, “compact” appearance, a short, thick neck, small ears, rounded ear tips, small eyes, a pointed muzzle, a break in the profile line between forehead and nose or a “dip” in the nose

            Pattern, colouring: The African Wild Cat of Southern Africa is distinguished by several features that renders it relatively easy to differentiate it from domestic cats. The most striking feature of the African Wild Cat is the colouring of the ears, which show a bright orange-red to orange-yellow. It is never greyish or brown, and even a dull red is a clear sign of a domestic cat within the forebears.

  The facial markings in the greyish or beige face are not very striking as they are of a pale fawn or mushroom brown but never black. The nose leather is a strong sienna red, by no means pink or grey. The chin usually is a light buff colour that can sometimes appear cream or even whitish.

The smoothly tapering tail always has a untainted black tip and three more or less clear black rings. In-between the rings the fur colour, other than that of the body, is a light cool grey (see also Fig. 2). In most hybrids, tip and rings are, if present, not so conspicuous. Hybrid’s tails also have a tendency to show more than three rings.

The back of the body is darker and greyer than the beige sides. The breast, rather than being white, is yellowish or a rich buff colour, a hue that is rare in domestic cats. The breast should not be the whitest part of the body, and it should not be too sharply set off. The patches on the soles of the feet and the markings around the elbow region of the legs are quite noticeable and of a deep black, while the spots that can flow together to stripes on the grey or fawn body are light brown and indistinct, in opposition to the black or very dark body markings in the typical tabby or mackerel (striped) domestic cat.

The coat of the African Wild Cat is dense and short, lighter and thinner in summer than in winter. The fur is smooth and shiny, and neither plushy nor woolly in texture. There should never be any evidence of longish hair.

In summary: While it is often easy to recognise a cat as being partly or fully domestic, it is sometimes very difficult to distinguish a pure African Wild Cat from an only slightly hybridised one. One should be careful to breed with specimens that lack even only one of the features that, in correct combination, make up a proper African Wild Cat.


Today, nature conservationists are dedicated to control the numbers of feral cats, especially within game reserves. Most of the terrains are, however, too small and too far apart from each other for sustaining a pure African Wild Cat population (if ever it started as pure in the beginning). The large ranges of male cats, in particular while in search for a suitable sexual partner, and their cunning in crossing even high, electrified fences, makes it almost impossible to keep them inside a reserve. Vice versa, domestic tomcats are likely to penetrate the sanctuary for the same purpose, doing severe harm to any action plan trying to save the African Wild Cat.

As already mentioned in the introduction, what we need is controlled reproduction, which only can be obtained in captivity.

The question of how strict one should be in preserving purity will certainly have to be a matter of discussion. In our experience, there are so many African Wild Cats that are not entirely pure as regards to hybridisation with domestic cats, that breeders sticking to pure breeding strains will face a so called “bottle-neck” situation, i.e. the gene pool might become to small for first-rate breeding results, or, in other words, there could emerge a severe inbreeding problem resulting in poor, weakly or sickly specimens. Thus it could be a good plan to carefully combine different but still similar southern African subspecies, viz., griselda of the Kalahari and Namibia, and cafra from South Africa. In the Northern Cape and some parts of the Karoo the appearance of the African Wild Cat is in any case intermediary. In consequence we think the avoidance of domestic cat’s influence on the genetic pool could have, nowadays and even more in future, priority over the strict separation of neighbouring subspecies.

It is a shame to say so but our efforts however hard to keep a population of African Wild Cats completely free of domestic gene influence cannot be thoroughly successful. Every meticulousness to keep that influence small is thus highly desirable. The best strategy for a successful re-introduction is to select wild life sanctuaries wherever we see a chance that hybridisation can be avoided for at least a few generations, starting with a population large enough to diminish the danger of mixing with undesirable specimens from outside. The above-mentioned fertility of the African Wild Cat could make this well possible. The strictly territorial behaviour of once well-established female African Wild Cats will additionally help in avoiding the intrusion of alien females quite successfully.

The breeding requirements for re-releasing purposes are of course much more complex than those for specimens intended for a life in human care. Re-released specimens should be accomplished in avoiding humans and their settlements as far as possible. They have to possess special skills for the survival in the wild.

How this can be done is a wide subject that requires still more thorough research. A couple of re-release experiments have already been successfully carried out by the authors, though on a small scale. Some of the individuals that have been bred in our institution and re-introduced in nature reserves are being monitored by students via telemetric means. Presently they are in the second year of their life in the wild, still appearing sound. How the offspring of those cats will look like, only the future will show.


In the gallery you will find pictures of pure-blooded African Wild Cat specimens. Take notice of the height of the walking cat and the facial features and the faint markings of the resting one. You can also see the typical ear size, shape and colouring of an African Wild Cat. Note also the lack of conspicuous fur markings on head and neck.

We developed a measuring method to calculate the proportions of a Felis libyca specimen. Our still preliminary results on photographs of standing or walking animals are: the ratio length to average height (= 1.31 in the figure) is rarely below 1.5 for domestic cats but about 1.25 to 1.4 for African Wild Cats. 

Behavioural Enrichment for Blackfooted Cats (Felis nigripes) in Zoos

Mircea and Jörg Pfleiderer, Honingkrantz, P.O.Box 20, Fish River 5883, RSA

1. Introduction:

Behavioural Enrichment may be a new expression but has for a long time been a well recognised task. Nearly all that can be said on smaller cats in zoos has already been mentioned by the late Paul Leyhausen forty years ago. We can only fill in details. That the concept need not be a purely theoretical one and certainly is not in the Johannesburg Zoo, has been – for small carnivores – nicely demonstrated last year by Rynette Coetzee (2000).

It is only too well known that Felis nigripes, the Blackfooted Cat (BfC), generally makes a difficult and rather dull zoo exhibit, at least in European and American zoos. Only very few zoos keep that animal, examples being the Wuppertal Zoo in Germany, or the San Francisco Zoo. In these zoos, we learned of the same problems in virtually the same words. Keeping is difficult. There are health problems, especially of the respiratory system and the kidneys. But there are behavioural problems as well: The cats are very shy and not a bit tame, not even towards the personnel, they hide away and sleep during visiting hours, thus making a poor zoo exhibit, and they have a pronounced inclination to stereotypic movements. In short: BfCs are dull and boring, sick, non-social, and generally don’t reproduce well: a nuisance for any zoo.

Our own investigations in the veld and in enclosures tell quite a different story and suggest that the situation could be dramatically changed by moderate changes in the keeping.

2. Methods and Material

Our project was started about 7 years ago by M.P. and by Leyhausen who had also been, in the 1960s in his institute at Wuppertal (Germany), one of the first to do any research on BfCs. The original aim of the project was to gain more insight into the wild life of these cats (and other small cat species of the Karoo). That is, veld observations were to be the dominating task. These were done mainly in the Eastern (High Rainfall) Karoo, first at the Mountain Zebra National Park, then at the farm Gannahoek south of Cradock, then in the Agtersneeuberg Nature Reserve, and since recently at the farm Honingkrantz near Fish River. While it is difficult actually to see BfCs in the wild, tracks can easily be found. Together with scat and scent marks, such observations can give a good notion of what the animals are doing. Thus, we studied the whereabouts of an approximate total of 8 BfCs, of which probably 5 were males. These studies were complemented by some zoo observations and other observations of captive animals, as well as close studies of 2 males in captivity. A first detailed report was given by Pfleiderer & Leyhausen (1998). For the present purpose, the captivity observations were, of course, particularly elusive.

We started with a small movable outside enclosure of size 3m x 1.5m which proved to be satisfactory when filled with sufficient accessories. We have now a fixed outside enclosure of 3.5m x 7m.

Jock, found in the veld near Douglas (Northern Cape) at the end of 1997, came to us when he was about 3 months old. We kept him from January through May 1998. We then had to leave him at another place and regained him only in February 2001.

Koos, found in the Cradock region when very young, was more or less hand raised. He was about one year old when he was lent to us in January 2000. He was very shy at that time and not accustomed to people spending time with him. He stayed with us about 4 ½ months.

3. Results

  1. Shyness and tameness

As all zoo people, but astonishingly enough not too many other people – including human psychologists – know, shyness and tameness are not at all antagonists but neither are they syn-agonists. Up to a certain degree, it is a “both” and “and”. They are somewhat reciprocally connected in the sense that an increase in the one always means a decrease in the other. But the reciprocity is by no means strict. It depends on the circumstances how much a change of one influences the other. We have the power to select the circumstances but, sadly enough, not so often the proper knowledge (which, by the way, we would call a good hand). In any case, a good balance of the two is what should be a goal in any animal keeping.

Amongst others, R.F.Ewer (1968) declares BfCs to be absolutely untameable, and many Zoo personnel would agree with that statement (pers. comm.). True enough, we know of no case where a BfC became as tame as, say, a caracal easily becomes. But neither do we know of a consequent and, especially, competent attempt from birth on. But tame they can be. Leyhausen & Tonkin (1966) who studied, 40 years ago, 3 BfCs in captivity, found them to respond quite well to food offered from the finger, and one specimen even slept sometimes on Tonkin’s lap (pers. comm.). Ours certainly did not become pets but lost most of their shyness even towards people unknown to them.

Our first BfC in captivity was a male, Jock, then a few months old. More by regular playing sessions than by hand feeding (note the importance of not only feeding!) he became, within weeks, quite fearless of humans and he has never since lost his boldness. He spent nearly three years at another place, with little attention except feeding. Nevertheless, he did not stop coming forward whenever he heard or saw someone approaching, even a dog, and even in bright sunlight.

Surprisingly enough, when we once visited him and resumed his favourite hunting game “Leather mouse at a string”, another male, fully matured and still quite wild and shy, came forward to take part.

Jock is now back with us. It took a few days in which we saw little of him before his inquisitiveness took over again. Now he does not mind our presence in the enclosure at all, provided that any too close approach is initiated by him rather than by us. When – to give an example – we sit in his cage and have lunch, even together with people he has never seen before, he certainly wants to partake, and takes food from between our feet, or directly from our hands.

Koos, the other male we tended last year when he was about 1 year old, also became quite tame after a few weeks of experimental playing and feeding sessions (where we tried to make the feeding sessions also a play). It took him, however, fully 4 weeks of daily training before he first took an active part in playing.

While BfCs seem to live extremely solitarily in the wild, they nevertheless respond very positively to attention. What they really need to get accustomed to a new situation is time. Let the cats approach you and do not try to approach them in undue haste.

b. Stereotypic Movements

These cats who walk or run, in the wild, for kilometres a night (Sliwa 1998) can run around the enclosure continuously “like a little toy train”, as Leyhausen (1961) once described it. But they do it, to our knowledge, as monotonous movement only if there is nothing else to do. We have seen our cats run around or to and fro in their enclosure more than a few times and monotonously only when they were very uneasy. Generally they run, if at all, only a few meters, then stop and investigate, or sit and look around. Sometimes, they move something around, or play hunting. But they do it only if – and that “if” counts – they have something to play with or to investigate. This is why an interesting cage topology with accessories, both which change now and then, is so essential. A variety of landmarks is useful. We put into our cage tree stumps (lying, not erect – BfCs are not very inclined to climbing) to jump on, sit on, use as lookout place, sharpen the claws upon, as well as to lie below. We also had stones to sit on or run on, as well as grass vegetation and Karoo bushes to hide away in – or bask in the sun without feeling “seen”. Grass is indispensable for their health (Leyhausen & Tonkin 1966) as they consume it daily and go off their food if they are without it for too long.

We cannot, of course, completely exclude monotonous running in the night when the cats are not attended. We have, however, occasionally observed that our cats, after a tiring play session, retreated to a resting place which they apparently did not leave for several hours. In the small enclosure of about 5 sqm we had originally for Jock and later for Koos, we could not detect any sign of frequent monotonous running in soft ground but they might have used a low stone wall on one side of the cage. In the larger enclosure of some 25 sqm we use now, Jock often takes the same path from a stone house to the gate, which has, by frequent use, been cleared of loose sand. But the position, more or less diagonally through the enclosure, as well as the form which is far from straight and avoids the little Karoo bushes growing in the enclosure, reminds us much more of a path in a dry-riverine sand bed which we found BfCs to use frequently and repeatedly in the wild, than of a path made by stereotypic movements as seen in zoos. Also, there is apparently no difficulty for him to change the path immediately whenever we put little obstacles on it.

Thus it seems to us that stereotypic movements need not be a problem as long as excessive monotonous running is avoided by allowing sufficient other activities.

c. Play

All cats like to play sometimes, younger ones more than older ones. BfCs are no exception. A number of objects to play with will help to keep the cats occupied for a considerable part of their active time. They enjoy investigating new objects: a piece of cloth, or wood, a rubber pipe, a leather skin. A worn-out sweaty jersey is of great olfactory interest. Don’t expect to get it back undamaged. Koos liked to drag around comparatively heavy and sturdy objects, the largest being our observing chair, measuring about 45 cm square and 15 cm high, and weighing as much as the cat. When dragging it he looked like an busy ant.

Wild BfCs walk preferably in dry sand beds of little rivers. Most scent marks we found were adjacent to such beds. Sometimes we found evidence of a cat having not only walked but jumped around in a way which we think must have been playful movements, followed by a – probably short – rest.

No wonder both our cats were extremely fond of heaps of sand and gravel which should be replaced or shovelled around frequently not only because they use part of it for their excretions (a well-defined part of it, by the way, and scat and urine separately) but also because they like playing around with it. Both cats visibly enjoyed the scent and feeling of fresh dry sand. They used it as their preferred walking surface, they dug in it and shoved it around. Jock used to scrape out moulds in the sand and to flatten himself into them when lying in wait for his toy prey. Such moulds we also found often in the sand beds of wild cats’ territories.

As hunting is an essential part of a wild cat’s life, hunting games are welcome even for older cats. A piece of leather on a string, perhaps with a little stone sewed inside to give it extra weight and speed, has proved to be the source of a large variety of different games. It is not always necessary but generally a particular attraction if a person is on the other side of the string. Both our cats could be kept busy for hours.

Food intake is also much more interesting if it does not just mean eating. In the wild, cats have to work for their food. Ours enjoyed ripping off meat from bones, chewing on bones, and eating nearly everything they could cut into pieces. They did not mind getting a whole Eland’s rib in one piece. From a small chicken, nothing remains but part of the legs. Bones and feathers seem to be vital. The occasional arthropod has the advantage that it can be fed live, even in zoos. Another well-accepted procedure is that we held fast a large piece of food. The cat got it only if it was able to rip the whole thing from our hands, or rip off a part, or chew it off in parts. Whoever fed him could easily lift up the whole cat without him loosing his grip on the meat he fed on. Both cats liked it even if they looked very furious. Don’t forget to wear strong gloves! Both cats sometimes showed signs of disappointment if the food access was too easy, and both grew easily bored if we offered the same kind of food several times in succession. They liked a frequent change of the feeding place as well as hiding the food under bushes or putting it at places with difficult access.

It definitely helps for changing the nocturnal habits a bit if feeding and playing is often done in daytime. Our cats learned to be alert at any time of the day when we attended them regularly at certain times. This, however, worked noticeably better in the morning and in the afternoon than around noon, which latter time they apparently considered as somewhat ill-chosen. But if something interesting occurs at noon, they do not hesitate to wake up and come forward and look.

d. Resting Places

Sliwa (1998) observed BfCs to sleep every day at a different place within their territory, coming back to one particular place only after about 3 weeks. We have similar indirect evidence. A BfC came to a particular place (water hole) in rather strict 3-week intervals. Sleeping or resting places which we found to have been recently used were not used again for quite some time (however, it was of course impossible for us to completely avoid leaving some evidence of our presence there!). We also found places that had been used as sleeping places but not recently. In most cases, the places were hollow termite hills, illustrating the cat’s Afrikaans name “miershooptier”.

Both our cats used essentially all possible hiding places: A very protected one under a hollow tree stump, a stony compartment, a more open place under a tree stump, or below our observing chair. Note that we supplied this number of hiding places even in the original small enclosure (5 sqm). Later, we added a hollow termite hill which was very well accepted. When Jock was young he used to change his favourite resting place rather regularly every 2 – 3 weeks. It was particularly interesting to see him when he had chosen the observing chair which is only about 10 cm high in the middle. While he grew, he soon barely fitted under it so he more or less pressed himself in. He did not stop using it when it rained and the place became somewhat wet. He even accepted our sitting on the chair while he was under it. The only situation when he agreed to sit in our hand was when the observer sat on the chair and put one hand under it before Jock came because then he had no other choice, except not to come at all and staying elsewhere. Now, being much older, he often changes resting places several times per day, depending on weather, time of the day (direction to sun), and what might happen around. After a few weeks of adaptation to the new enclosure, he has again detected the observing chair as a good and warm hiding place and uses it wherever we place it in the enclosure.

The subadult Koos changed his favourite place at irregular intervals, sometimes within a day, but was then again rather strict in keeping to it for a while. He never accepted the observing chair as a hide, assumedly because he found it a little too open (but he used it as a toy). As compared to Jock, he generally hid more – i.e., less of him could be seen – in all different hiding places.

We have never seen our cats sit in the open for more than a rather short time, particularly in day time, or to sleep in the open. But with a little protection from behind, they were quite content to be at one side in the open, be it for sleeping, or for looking around and noticing what was going on, or for basking in the sun.

It seems mandatory to us to provide captive BfCs with a choice of several resting and sleeping places offering varying protection. Now and then a new place, or an altered one, will be welcome. One dry place should offer complete, others partial hiding. If there is a possibility for them to hide completely, these inquisitive cats often will not do so – provided, of course, that the other circumstances are good -, or they come out, fully or partly, just to see what is happening outside. But if such possibility is not given, the cats will unsuccessfully try to hide which will drive them into what is called defensive sleep (Pfleiderer 1990; 2001), or into monotonous and even stereotypic movements.

e. Health problems

It has been suggested that a genetic defect in the world-wide zoo population of BfCs is responsible for the respiratory tract and kidney diseases, even when it is known that specimens that are caught in the wild also develop these problems. Instead we feel that some old traditions of zoo-hygiene might be responsible: a water bowl in which the water is frequently replaced (so there is no control over the fluid intake), and, worse, a concrete floor in the inside enclosure that is frequently sprayed with water to flush out dirt and germs, combined with heating, especially floor heating.

Evidence from our field observations as well as from those two males we kept in captivity shows that the animals are astonishingly tolerant to any weather conditions. We have seen them to be active in very hot dry weather as well as in pouring rain, even in prolonged rain, or in nightly frost. Given a really dry (i.e., including low relative air humidity) place to retreat, the cats would probably readily adapt to any climate wherever the zoo may be situated. However, the combination of wet floors and heating creates a very humid atmosphere, in which all sorts of germs thrive but certainly not BfCs. Not only the respiratory tract, but the whole system of this (semi)-desert cat is not able to cope with that sort of microclimate similar to a rain forest for any extended time. Our advice is to sluice less often, renounce heating altogether, and use what seems to be rather unknown in zoos: dehumidifiers (this is not really new, see Leyhausen & Tonkin 1966).

BfCs are used to living with next to no water at all. A BfC in the Mountain Zebra National Park used to visit a water hole only about every 3 weeks. We did not see it drink, but it ceased to return after the hole dried out. In some territories (and also in some distance from these territories’ borders) we found no fluid water. Our cats in the enclosure refused to drink any water, except very occasionally some rain water. They made a habit of burying the water bowl in sand, as they do with remains of rejected food (which, occasionally, has nevertheless been eaten up later on). We know of two male BfCs (one case reported by Leyhausen), both fully matured, that used the water bowl as a toilet.

The main source of the cats’ natural fluid intake is probably blood and meat juice of their prey. Ours liked to lick up every single drop, even when it was somewhat watered. Jock went once wild on freshly congealed blood from a slaughtered sheep (unfortunately, we were unable to repeat this experiment). Rats or chickens should, therefore, not bleed out before they are served, and the juice running from deep-frozen meat while thawing should not be wasted. Juicy meat should rather be served in bowls to minimise the loss of fluid.

Our cats were also very partial to milk, and digested it without the usual problems if given with care (temperature, concentration, quantity). They liked soured milk as well. In any case, one should not rely on their feeling so thirsty as to drink water – they may never do so, especially in humid air where they may just not notice what they need, and consequently become chronically dehydrated which, if it is not the main reason, will certainly contribute negatively to any kidney disease.

f. Intraspecific social behaviour

Free ranging BfCs are extremely solitary, living singly in huge territories that only marginally overlap. Contact with neighbours is quite infrequent and mostly indirect (by investigating each other’s scratch, urine, and scat markings). Both our cats were visited by the owner of the adjacent territory, an adult male, just 2 or 3 times each to see what was going on. We heard him calling. Especially during the extremely short mating period of some 35 hours, BfCs communicate by their surprisingly far-carrying calls. Cats kept in separate enclosures will also call each other. This does by no means mean that they want to live together. They just want to know where the neighbour is, being cut off from the indirect contact.

According to Leyhausen (1979), solitary cats kept together in larger groups will form a so-called “absolute ranking order”. Not only the display of aggression is suppressed, in order to avoid frequent, potentially dangerous fights, but also many other aspects of their social behaviour. Indeed, much of their activity, not only social activity, stops. The cats feel constantly uneasy and overproduce stress hormones. This is very contra-productive to all stages of reproduction and – as is well known – decreases the resistance to many illnesses.

We had nice demonstration of this in a group of 6 (3,3) BfCs kept together for breeding purposes on a game farm. There was no breeding for the 2 seasons for which this group existed. The animals were not sick but did not look healthy either and suffered visibly from ectoparasites. One male was often, another sometimes, the third occasionally, and all females practically never seen in daytime, save for one single occasion, where one hot summer afternoon at 4h all 6 of the cats could be seen lying openly on the concrete-like mud ground, each cat at a maximum distance from the next one. This gave the impression of the enclosure being “littered” with cats. Whether it was the heat or some other reason driving them to this unusual behaviour we could not find out. Nevertheless, the cats behaved somewhat uneasily as typical for “defensive resting” (Pfleiderer 1990, 2001). Under more “normal” conditions, the females almost never used the full range of the enclosure. Instead, they cowered closely huddled together in a den and showed every sign of permanent stress and fear. During a few night observations which we could do, the females also hardly ever made their appearance.

In many places, it is usual to keep pairs, and this does apparently often (but not always) work. We nevertheless think single keeping to be preferable. Two smaller single enclosures are better than one double-sized enclosure for a pair. If two enclosures are adjacent or close together, the cats should have places where they cannot be seen from the other side, i.e., each cat should have the opportunity to avoid mutual and one-sided eye contact, even if they might seek eye contact often. If they should happen to like each other, they will show that they would prefer a door between the enclosure to be opened.

4. Conclusions

The keeping of BfCs in some zoos can be improved. We have discussed possible reasons why difficulties arose which we – up to now – were fortunately able to avoid. We specially mentioned topology of, and accessories in the enclosure, activities of the personnel and possibilities of the cats for own activities, food and fluid intake, the need to provide them with a dry place, and finally some social questions.

We claim that there is often an imbalance between the efforts invested into physical and into psychological health. Life outside the retreat must be more interesting for the cat in order to leave the retreat and come out. Within, of course, certain limits a dirty “world” is better than a clean nothing. If necessary, we left our cats for days in their hiding and did not know where they were even in the small 5 sqm enclosure, but we never let them stay like that.

The boring appearance of some zoo enclosures would drive any decent person, let alone a BfC, into stereotypic behaviour. Any “happening” which the cat can itself induce with the right topology of and accessories in the enclosure – including changes which are always well appreciated –, is better than any static “nice-look”.

The importance of playing, even for adult cats, cannot be emphasised enough but we cannot offer a proposal applicable to every zoo. This is because it definitely takes time that may not always be available. As much as the cats like a clean enclosure and might find the cleaning procedure to be an interesting “happening”, an occasional neglect of hygiene in favour of spending some time in direct playing may well be worth while. We found it is at least a compromise to make feeding as much a play as possible. In this context, it seems necessary to us to again mention the convincing work of Rynette Coetzee (2000).

Not only cats need a time when they can very relaxedly sleep. Miershooptiers, being essentially nocturnal, need to have part of that sleep in daytime, and having large territories, need more than one choice for a good hide-away.

A good zoo is probably most interesting in night time, at least to a cat, mostly because the noise made by annoying visitors is replaced by the fascinating noise coming from companion animals. This is why changing the habits of a BfC into a more diurnal direction might be more difficult in a zoo than on a Karoo farm where daylight noises are also interesting, not to speak of the changing views which you can have on a farm. All the more it is necessary to make zoo life in daytime as interesting as it can be. Feeding, cage cleaning and other activities concerning the cats should, if possible, be done during visiting hours.

We think that three causes of health problems can be avoided: Too much humidity even in the retreat (i.e., the cat is too long exposed to it without interruption), chronic dehydration from inadequate fluid intake, and unnecessary social stress. For the latter cause, we have specifically mentioned keeping too many cats together, but many other reasons for social stress would apply as well, and some can be avoided. Particularly, any kind of boredom contributes to social stress.

Finally, as is well known and perhaps need not be mentioned at all, social stress in any form may contribute severely to breeding failure.

All in all, behavioural enrichment in this widest sense is mandatory for these cats to show how lively, inquisitive and bold they really are.


Our thanks go to Brian and Renee Wayland who started our enclosure observations by providing us with Jock, and to Walene Marais who lent us Koos, as well as to the staff of many institutions and private places where we were allowed to do our research. We are particularly grateful to the late Paul Leyhausen and to Barbara Leyhausen-Tonkin who never hesitated to give necessary advice and to share their vast experience with us.


Coetzee R. 2000: A few simple ideas for enrichment for small carnivores, PAAZAB Ann.Conf.Proc.,45

Ewer R.F. 1968: Ethology of Mammals, Logos Press London

Leyhausen P. 1961: Smaller Cats in the Zoo, Intern.Zool.Yb. 3, 11

Leyhausen P. 1979, Cat Behavior, Garland STPM Press New York, London

Leyhausen P., Tonkin B. 1966: Breeding the Blackfooted Cat in Captivity, Intern.Zool.Yb. 6, 178

Pfleiderer M. 1990: Zum “Verteidigungsschlaf” von Carnivora im Zoo, Zool. Garten NF 60, 228

Pfleiderer M. 2001: Sleeping Cats in the Zoo, PAAZAB Ann.Conf.Proc., this issue

Pfleiderer M., Leyhausen P. 1998: A field study of the behaviour and ecology of three feline species, PAAZAB Ann.Conf.Proc., 39

Sliwa A. 1998: Africa’s smallest feline – the Blackfooted Cat, Endangered Wildlife 28, 10

In the meanwhile we are keeping many more BfC, males as well as females, adult as well as juvenile specimens. 2 pairs are at present involved in a successful breeding program. The oldest specimen, the above mentioned Jock, is now 10 years old and still a successful hunter, healthy and strong, and occasionally sexually active.

M. Pfleiderer, June 2007

Social Behaviour, Breeding and Territoriality of the Clouded Leopard (Neofelis nebulosa) in the Zurich Zoo

Mircea Pfleiderer, Inst. f. Zoology, Leopold-Franzens-University, A-6020 Innsbruck, Austria


The Clouded Leopard, a medium sized felid, is a rare but generally very rewarding zoo exhibit due to its beauty and agility. The Zurich Zoo (amongst others, as Frankfurt, Duisburg and Dresden) has already a good tradition in keeping and breeding the species. In spite of the profound experience of this zoo, the generous providing of space as well as the optimal structuring of both the inside and the outside enclosures, to which the cats had unlimited access, the two recent breeding pairs never mated although the hormonal condition seemed to be sound. When discussing the matter with the director and the curator, I was invited to investigate these cats and possibly to find a solution to the breeding problem.

Specimens, observation methods


 “Tutong”, born in Duisburg Zoo, 1993

 “Sawa”, born in Zürich , 1992

both kept together in the display enclosure since March 1999

 “Phan”, born in Zürich, 1989, same parents as Sawa;

 “Kiri”, born in Italy, 1990, hand raised,

both kept together in the “Cat breeding house”.


The inside display enclosure consists of 4 partitions combined to one large inside enclosure, with at least 20cm of uneven mulch ground, generously equipped with vertically and horizontally arranged tree trunks, branches, ropes and a dense “forest” of long bamboo sticks with the dry leaves left on to provide sufficient cover; the outside enclosure, a combination of two large partitions, has a deep natural soil ground allowing a dense vegetation of trees reaching to the top of the enclosure, as well as grass, fern and moss. Part of the outside enclosure and the whole of the visitor-facing side of the inside enclosure is equipped with thick glass panes. Such measures are quite efficient in giving the cats sufficient protection from the visitors (see PFLEIDERER 1990, 2001). Defensive sleep was only very rarely observed, and always in a mild form (the protocol states: “component of defensive sleep”), owing perhaps to the fact that, for both animals, the tree branches preferably used as sleeping places gave good protection. Both spent approximately half of their activity times in the inside and the outside enclosures but only rarely more than 20 minutes at a time in the latter.

The “Cat breeding house” provides outside enclosures of similar size and equipment as those of the display site, and, in addition, some thick ropes horizontally suspended between the trees for the cats to walk on. The inside rooms (one chamber and two small retreating compartments to the left and right of the chamber) are furnished with boards and cardboard boxes for the cats to rest and sleep. The concrete floor remains bare.


The fourteen days’ period of intensive investigations was done in October 1999, covering all 24 hours, with changing resting breaks in the first week, and concentrating on the night hours corresponding to the predominantly nocturnal habits of the cats in the second week, leaving part of the day observations to the zoo personnel who were very helpful and supportive, providing me with additional information about the history of the animals, previous breeding attempts etc., lending me a second, stronger torch, and equipping parts of the inside display enclosure with infrared lamps to avoid the undue disturbing of the cats by blinding them with the torch during the dark hours.

The weather conditions were rather typical for this stage of the Middle European autumn with temperatures around 0-10 °C and more or less constant drizzle, mist and very little sunlight.


The respective behaviour of the pair on display towards each other ranged from ignoring, disregard, avoidance, defensive behaviour, to rarely performed but clearly recognisable open hostility.

The behaviour of the pair in the breeding house was less hostile, but an unbiased observer could still describe the atmosphere between the two as “chilly”.

  • Activity, use of the enclosure

Similar to FURTER’s (1991) observations on 4 other Clouded Leopard specimens in Zurich, and not unlike most of the other Eurasian and African Felid species I investigated, the activity graphs of all four specimens show two definite peaks, one in the morning, one in the evening. There is a well-defined resting phase of several hours around and after midnight, and a second, somewhat more diluted one around noon. AUSTIN & TEWES (1999) found a similar pattern in two wild specimens. During most daylight hours, the cats – except Sawa when she was in heat – remained more or less inactive, sleeping and resting, grooming themselves and looking around. The main purpose of locomotion was the change of resting places.

The actual times of activity were markedly different for Sawa and Tutong, the pair on display. There was certainly a tendency for both animals to be active essentially only if the other one was fast asleep or pretended to be. In the morning, Sawa was the more active one, especially in the early daylight hours, when Tutong had already finished his morning exercise. The second time of activity occurred for Sawa essentially in the afternoon, for Tutong in the early night hours, with only a small overlap between the two. But even in these hours of overlap of the average activity times, there was very little actual overlap. Often, one animal got active for a few minutes only just when the other had an equally short rest.

In total, the female was consistently (but slightly more pronounced during her heat phase) more active than the male: 29% of the time as compared to 22%. If these activity times were distributed statistically, one would expect no activity at all in 55% of the time, and simultaneous activity in 6.3%. Actually, activity overlap was observed in only 2.6% of the time, of which no more than 0.6% were some form of interaction while the remaining 2.0% were independent activity. The difference is fully explained by the different average activity times. It was, however, not possible to decide whether this latter difference is a consequence of the strong mutual-avoidance tendency or rather an indication of different personalities. The latter interpretation is supported by radio collar observations in the wild (AUSTIN & TEWES 1999) in which the female showed quite more diurnal activity than the male.

It is interesting to compare these numbers to those for quarter-hour intervals in which Sawa was active (= activity for more than 2 minutes) in 35%, Tutong in 30% of the time, with an overlap of 9%, and both inactive in 44%. These numbers indicate that many of these intervals are not completely filled by activity. They corroborate certain facts which were already impressed on me while observing:

  1. When an animal starts being active, the activity time lasts, with only short interruptions, often for one or more hours. An average duration of some 1.3 hours explains the difference between the inactive 15-minute intervals (44%) and the inactive 1-minute intervals (52%).

  2. The female is more active than the male, especially in daytime (but less active during evening hours).

  3. Quite a large percentage of the day, namely about ¾ or about 18 hours, is spent in rest.

  4. The activity intervals of the male last shorter than those of the female. The male interrupts his activity significantly more often for a short rest.

  5. The activity times of the two animals occur essentially independently of each other if larger intervals of 15 minutes or more are considered. But they do avoid being active simultaneously on a short time scale of only a couple of minutes.

Both cats very rarely could be seen playing, and only with objects (e.g. food, rope end), never with each other. When active, they were mainly pacing around, sometimes falling into stereotypic behaviour by walking in peculiar little circles, or lifting the head, or raising on their hind legs in a certain corner etc. However, stereotypic behaviour was rare and always of short duration. Tutong showed just one single pattern, but only at 6 of the 13 days of observations altogether (with a total of 20 times). Contrarily, Sawa (with a total of 95 times) displayed stereotypic behaviour more often, sometimes only for a few seconds, up to ten times per hour during the time of heat, but with surprising flexibility of not less than 6 different patterns. These numbers probably indicate less, but not considerably less stereotypic behaviour than was observed with another male of the Zürich Zoo (Hartmann-Furter & Schiess-Meier 1994).

Both animals also climbed and jumped quite readily, and spent considerable time outside, looking around and listening intensely into the night. At least once a night, mostly in the evening hours, Sawa sat for up to 45 minutes in a corner of the outside enclosure where she watched the three tiger cubs playing in the neighbouring enclosure.

A striking feature with that pair was their avoiding each other, not only in time as described above, but also in space. Often, one cat stayed inside while the other roamed the outside enclosure, and vice versa. They interchanged their roles rather regularly, sometimes even every few seconds: One went outside the moment the other one came in. Sometimes it was clear that Tutong was the driving force, while Sawa essentially reacted to his movements, but often it was the other way round, and in the whole more mutual than one-sided. The same scene could be seen within the inside enclosure, which was used by both cats at the same time in daylight. When Sawa was asleep, Tutong restricted himself predominantly to the two right, “His” partitions, while Sawa was confined to “Her” two left partitions during Tutong’s resting periods which he invariably spent in the rightmost corner. Needless to say that these two cats never rested together, unlike the previous – more successful – breeding pair, “Kong” and “Cathay”, that was nearly always found sleeping in close contact, not only during the oestral phase (FURTER 1991). She also claims in the same paper that it may be difficult to get Clouded Leopards accustomed to each other; it just never worked with the pair “Chinkara” () and “Opai” (), then about 4 and, respectively, 5 years old.

The older pair that stayed in the cat breeding house, Kiri and Phan, had their activity phases more synchronised; in the evening hours they walked around, confining themselves almost exclusively to the chamber in spite of having free access to the large and optimally structured outside premises. In order to lure them into the trees I hid their food (large chunks of beef, whole rats or chicken) first in the lower, later in the higher, less easily accessible branches. Both cats fed only when they were left alone. The left-overs were always found inside.

In the morning hours Phan often played with the cardboard box he was given to sleep in, chewing off parts of it or shredding it to small pieces. Kiri usually walked around; unlike the pair in the larger display enclosure, neither she nor Phan showed any stereotypic movements. Both cats very strongly reacted to my presence. Phan, the usually quite lazy male, followed me wherever I went to observe the cats, trying to establish contact by, e.g., raising on the hind legs to initiate a “nose to nose control”, puffing and wheezing loudly. After a while Kiri followed, sometimes merely hissing at me, sometimes raising her voice in a long, ululating, rather weird-sounding howl of defiance.

For sleeping, the cats usually stayed each in one of the compartments; Phan sometimes preferred the cardboard box, especially if it was a new one, which happened to be every three to four days due to Phan’s habit in destroying the boxes. Like Sawa and Tutong, Kiri and Phan always seemed wary of each other, and kept each other at a distance.

  • Social and territorial behaviour

Part of this is already implicated above; with the two pairs I studied so far, one could gain the impression that the Clouded Leopard is more strictly solitary than most other feline species, which would be totally wrong as the information from the literature on this species shows. There are even records of pairs sharing a territory in the wild (LEYHAUSEN, pers. comm. 1997) just as I found with some African Wild Cats in the Karoo.

Eye contact was relatively rare; I could induce definite eye contact only 46 (76) times with Tutong (Sawa) while I could observe eye contact between the two animals only 26 times. Blinking (which is generally a friendly gesture) was still scarcer. Tutong blinked 5 times at me but only once, if at all, at Sawa. She, on the other hand, blinked 10 times at me, and I could see 2 definite cases of her blinking at him. With Phan and Kiri, I did not see any blinking at all.

Surprisingly enough, I could never hear any sort of vocal marking of the territory as it is done by many other Felid species; whether the reason for this absence lies in a certain inhibition the respective pair partners exert on each other, or this species is, like the Red Cat (Profelis caracal), one of the more silent ones, I can not say at present.

Another striking feature of the Clouded Leopard is a special way of olfactory marking not known in any other cat: When this cat, as nearly all cats do, rubs its cheek, muzzle and the side of its mouth on a hard object, then, depending on the intensity of this behavioural pattern, it often additionally opens its mouth while going on rubbing, eventually showing an impressively gaping maw disclosing all teeth. The best descriptive term probably is “Open-mouth rubbing”. Quite typically, the cat hugs the object, f.i. an upright tree stump, with the front paws holding on with the claws while standing on the hind feet; when marking horizontal branches they crouch or lie on the belly and also hold on with the claws. Occasionally, the cats end the action by rubbing intensively the nape of the neck against the marked spot.

“Open-mouth-rubbing” is, to my knowledge, not mentioned anywhere in the English scientific literature. A short description in German – probably the first mention at all – was given by FURTER (1991) who called it “Mundwinkelreiben”.

Even the way of territorial marking with urine is different in this odd cat species: No one ever saw them spraying urine as other cats do. Instead, they exclusively scrape out hollows in the loose ground and leave a few drops of urine in them. Sometimes they just scrape out and omit the urinating. It seems that this scraping out of (sometimes very shallow) hollows is a nearly obsessive habit at least for the display pair, since Tutong was observed to do it 62 times in the roughly 1000 hour of activity (average: once every 16 minutes) while Sawa did it 119 times in roughly 1300 hours of activity, giving an average of once every 11 minutes.

Claw sharpening or scratch marking acts I could register only 10 with Sawa, 3 with Tutong (all only in the inside enclosure) and none with the other pair (no objects apt to scratching in the chamber); as it was nearly impossible to study such behavioural patterns in the dark and densely vegetated outside enclosures, it is difficult to tell if the cats scratch outside more often. My closer investigations on the trees and other wooden objects of the cat-breeding-house enclosures to which I had access while giving food to the cats revealed only few, barely visible traces of scratching. This corresponds to the field observations of RABINOWITZ et al. (1987) and AUSTIN and TEWES (1999), who found the Clouded Leopard “being a species with secretive habits, thus disinclined to leave scrapes along forest trails”.

  • Sexual behaviour

The average duration of the females’ periods of heat was recorded in the zoo by YAMADA and DURRANT (1988) as lasting 6.2 days. AUSTIN and TEWES (1999), who obtained some telemetry data on the Clouded Leopards’ home-range use in a Thailand National Park, observed a pair staying in the same area for 6 days before moving in different directions again. As no direct observations were possible, it was assumed that the two animals stayed together for a mating period. The single oestrus I was able to observe directly also lasted about 6 days.

During that time, the activity of Sawa increased visibly. She assumed a certain restlessness, walking around, showing also an increase of stereotypic behaviour. Both types of scent marking, the scratching of troughs while urinating and the open-mouth rubbing were more and more often displayed; again, spray marking never occurred. From the second day on, reaching the peak of her heat, Sawa’s open-mouth rubbing often took into rolling. The reaction of Tutong to this was invariably rejecting: wherever he found Sawa’s scent markings, he marked “against” them on “his” side. Then he busied himself “reinforcing” the boundary lines by frequently marking the mulch near one of the three concrete thresholds that run through the inner enclosure.

Sawa never got as far as allowing closer contact, challenging the male sexually by presenting herself, or taking any sort of a mating position; in short, she never got the actual readiness to accept the male. In the few occasions Tutong looked at her or came nearer, Sawa fled, often hissing and flattening her ears. Small wonder, as she already had been bitten by him less than half a year ago.

From the information I got from the zoo keepers, Sawa came into heat, or rather showed its more conspicuous signs, about once a month, but mating never took place – not even unsuccessful trials. This was not a matter of the female not accepting the male’s attempts; the male just obstinately ignored the female.

I observed Tutong several times displaying aggressive behaviour towards Sawa, especially during her few and rather shy attempts to establish mere eye contact. In a previous breeding project, the same male killed a female (a sibling of Sawa) with a neck bite - whether in an overzealous attempt to mate or out of frustration, even the warden who saw the incident could not tell. LEYHAUSEN’s hypothesis (1966) offers another, quite plausible explanation: As the killing instinct of cats is not of the pure stimulus-response type of behaviour pattern, and develops its own internal urge if not properly released over a long period, it may then eventually grow to such strength that the cat will accept its (unloved) mate as a substitute.

Another accident happened when Tutong bit a female and the injury was not immediately detected; this cat also died from the infection that followed. Anyway, incidents like that are only too well known to anyone who keeps pairs of cats in captivity. In order to avoid another casualty , the present pair was kept in adjoining enclosures, at first allowing mere visual contact. The two enclosures were joined only later, and the cats were closely surveyed by video cameras. In spite of the fact that this had taken place more than one year ago (and with all possible care), my observations gave the general impression of a rather grudging armistice than a peaceful life together.

Conclusive remarks

An immediate solution to the breeding problem (well, actually it is a mating problem) cannot easily be found: The changing of mates might well be a success at least in one of the pairs, but is always a risk because the reaction of the cats is unpredictable. The possibility of a female being killed is particularly large in Clouded Leopards: They have the longest canines compared to the total head (and body) size of all Felidae, making the neck bite even more effective and thus more dangerous than that of others.

In the case of our two pairs there is an additional obstacle to a change of mates: Phan and Sawa originate from the same parents.

Separating the pairs for a prolonged time might induce the same risk when re-joining them because of the same reason given above, especially with Tutong whose hostile attitude towards most of the females makes him a peril to them.

Hormonal treatment does not seem necessary for Sawa, as the regular intervals as well as the intensity of her oestrus are sound; Tutong’s healthy appearance and well developed masculine habitus also does not indicate an immediate necessity of “pumping him up” with hormones that often have undesirable side effects – and could further boost his aggressiveness, too. For the older pair staying in the “cat breeding house,”, Kiri and Phan, however, a scrupulously monitored hormonal treatment may well be a cure to their lack of sexual – and other – activity.

Studies of the literature, conversations with zoo people and long discussions with my late friend LEYHAUSEN in addition to my own observations prove clearly that the Clouded Leopard (and not this species alone) is prone to a personal pair bonding. According to experience, a successful “pairing” does not only last during the periods in which the female will accept the male, but also when they sleep and rest in close contact, groom each other, not only themselves, play running, hunting and even fighting games – indeed continuously, and over a lifetime. In ranking order, nobody lags behind, or shows any fear of the partner. This is advantageous for the cats and the visiting humans, because life inside and watching life from outside the enclosure is by far more interesting for both.

However, there is a disadvantage in pair-bonding that cannot be ignored. It is their particularity selecting and accepting their partner: “Drum prüfe, wer sich ewig bindet” (Schiller, “Die Glocke”);. If the respective cats do not “like” each other, things will develop to an unpleasant, if not fatal, end.


My special thanks go to the Director of the Zoo Zürich, Dr Alexander Rübel, for the invitation, the Curators, especially Dr R. Zingg and Dr H. Schmid for their interest and support, and the staff for their readily offered help and co-operation.


Austin S.C. & Tewes M.E. 1999: Ecology of the Clouded Leopard in Khao National Park, Thailand. Cat News 31, 17

Furter M. 1991: Aktivität, Gehegenutzung und Verhalten von Nebelpardern Neofelis nebulosa (Griffith 1821) im Zürcher Zoo. Preprint Zoo Zürich

Hartmann-Furter M. & Schiess-Meier M. 1994: Seile als Klettergelegenheiten für Nebelparder (Neofelis nebulosa Griffith 1821) im Zoo, Preprint Zoo Zürich

Leyhausen P. & Falkena M. 1966: Breeding the Brazilian Ocelot-Cat Leopardus tigrinus in Captivity, Intern. Zoo Yb. 6,176

Pfleiderer M 1990: Zum “Verteidigungsschlaf” von Carnivora im Zoo. Der Zool.Garten N.F. 60, 228

Pfleiderer M. 2001: Sleeping Cats in the Zoo, PAAZAB Ann.Conf.Proc., this issue

Rabinowitz A., Andau P. & Chai P.P.K.1897: The Clouded Leopard in Malaysian Borneo. Oryx 21, 107

Yamada J.K. & Durrant B.S. 1989: Reproductive parameters of Clouded Leopards (Neofelis nebulosa). Zoo Biol. 8, 223

Sleeping Cats in the Zoo

Mircea Pfleiderer, Inst. f. Zoologie, Leopold-Franzens-Universität Innsbruck, Austria


Whenever a visitor comes to a zoo, expecting some nice display of animal action, several of the animals are, to his disappointment, fast asleep. “Lazy cats”, the visitor will probably think, as quite often the subject lying around in this very offensively inactive way, will be some or other representative of the feline family.

Indeed, the Felidae, when at ease, usually spend the greatest part of the day resting, looking around while lying, grooming themselves or each other, or more or less fast asleep. It is a well-known fact that nearly all cat species are active mainly at dusk and dawn, and that some are predominantly or even strictly nocturnal. In addition, cats, like most other carnivores, tend to save energy in between hunts. Some felines have the reputation of being lazy in the extreme, as f.i. lions who have been reported to sleep and rest up to 20 hours a day. So it seems quite logical that zoo cats are not very inclined to show too much activity during visiting hours.

While observing Carnivora, and Felidae in particular, in European, American, and also South African Zoos, it soon became obvious to me that the animals sometimes sleep in circumstances in which one would expect them not to do so. I gained the impression that the animals refuse to let the stream of visitors disturb their rest, even if their only sleeping places are exposed such as to allow the visitors a good view of them. Nevertheless, most of the cats did not seem to be at ease at all, in spite of showing signs of being asleep.

Psychological investigations and experiments on human sleep reveal that sleeping includes many more aspects than a mere absence of activity, and that there certainly is more than just one cause for sleep. The list of ethological papers on the subject of sleeping, however, is surprisingly short, considering the large portion of life that animals spend somnolent.

EWER (1968) was one of the first to point out that the concept of sleep as phenomenon on its own with specific causal mechanisms is a valid one not only physiologically but also ethologically, as evidenced by the choice of certain sleeping places and postures. HASSENBERG (1965) and IMMELMANN (1962) make special reference to species-type sleeping behaviour.

There is a particular form of sleep – or rather half-sleep, or pseudo-sleep, or situation-forced sleep – that has been known for some time to occur in birds but few reports are present in the literature. SCHERZINGER (1970) described pseudo-sleeping of the Pygmy Owl in situations from which there is no escape. THALER et al. (1981) observed it in a Bald Ibis whose young continued to beg importunately for food after the supply was exhausted. Inasmuch as, apparently, a stress situation is responsible for this kind of sleep, it is certainly related to the subject I shall discuss in the following.

The Problem

My first investigations, done on Lynx lynx, Felis silvestris, Canis lupus and three mustelid species in the Alpenzoo (PFLEIDERER 1990), revealed an aspect of animal keeping which so far had received little attention and is connected with the problem certain groups of animals have while being displayed to the public: “Being visible”, by no means a problem per se, is changed instantly by the presence of visitors to mean “being seen”, even “being stared at”. This is not so much a feature of animals who cannot stare straight at one another because their eyes do not face forwards (f.i. ungulates, rodents). Staring as a means of intimidating a social competitor is not part of their behavioural pattern.

It is quite a different matter with the animal species whose eyes face forwards – mainly primates and carnivores. Eye contact and looking away are important components of their social behaviour. Staring means dominance or at least the attempt to dominate. Correspondingly, letting oneself be stared at gives a feeling of social inferiority. In the zoo, those animals have constantly to face eye contact with the visitors, thus feeling as if they have been “placed in pillory”. When there is little opportunity to hide, the only means of escaping the continuous social stress is to cut off contact as thoroughly as possible. The animal turns its back to the ogling public and closes its eyes tightly: “If I cannot see you, perhaps you cannot see me either.” This often leads to a very characteristic kind of sleep, or feigning sleep – sometimes it is hard to tell – which I called “Verteidigungsschlaf” – “Defensive Sleep”.

As the two feline species of the Alpenzoo showed the most pronounced inclination to this kind of sleep, and I even observed the phenomenon with domestic cats in certain conditions, it seemed worthwhile to do further investigations on the felid family.

Species and Methods

Observations were carried out over a total of 17 years in 35 Zoos (5 South African , 15 German, 4 Austrian, 3 Polish, 2 Swiss, 1 Hungarian, 1 Czech, 1 Spanish, 3 North American) on 35 of the 41 species of the Felidae, in 11 of the 13 genera, according to the system of Leyhausen (1979) and Pocock, and Leyhausen (1999, posth.),

Time of day, weather, air temperature and volume of visitors were noted, as well as details of the animals’ behaviour. As the nature of the resting place (f.i. mould, tree stump, hollow trunk, branch) also may have an influence on the sleeping posture, it was included in the notes and sketches. Where possible, I took photographs of the enclosures and the animals. As far as possible, all personal contact with the animals was avoided (such as visiting certain specimens throughout too long an observation period, because in some cases, the animals began to recognise me and behaved atypically), so that the observer resembled a zoo visitor.

These investigations were complemented by observations on 5 Feline species (Acinonyx jubatus, Panthera leo, Profelis caracal, Felis libyca and Felis nigripes) in the wild (Southern African national parks, nature reserves and farms), on numerous tame specimens (Acinonyx jubatus, Panthera pardus, Panthera leo, Neofelis tigris, Profelis concolor, Profelis caracal, Felis nigripes) and by observations on domestic cats.

The main observational information needed for the present purpose is very simple: “If a cat sleeps, how does it sleep?”


The various sleeping and resting positions may be divided into two clearly different basic types; these occur, however, in all possible transitional forms, which may sometimes flow into one another.

a)Recuperative sleep, relaxing

Recuperative sleep occurs when there is a genuine need for sleep. This is the normal, and, in favourable conditions, exclusively observable type, which felids in particular need several times a day. Especially the large representatives of the Felidae which are provided with generous amounts of food but only at long, irregular intervals, sleep long and frequently to save energy between active phases (hunting, strife with conspecifics, sexual activity). In the Zoo and in the wild the lions are the most persistent sleepers. In the Etosha and the Kruger National Parks I was able to observe all forms of purely relaxed rest and sleep behaviour with them, but never any sign of stress. This also applied to the Caracals and the Blackfooted Cats I found sleeping in the veld; only they felt more easily disturbed by the click of the camera, and, in most cases, bolted.

Zoo felines also can display recuperative sleep, but this is almost always restricted to occasions when, on account of the weather, the time of day or season, the stream of visitors – the main source of stress – is greatly reduced.

Characteristics of recuperative sleep

  • Face: “Smooth” expression – no skin folds on the forehead and around the bridge of the nose. The eyes are lightly closed, there is hardly any noticeable movement of the eyelids.

  • Position of ears: Symmetrical, slightly turned outwards, not pricked. No movement of ears in order to follow noises.

  • Body posture: There is a distinction between resting and sleeping positions. In the resting posture, the animal usually lies on its stomach, less often on its side. The head is held upright. When the animal is relaxed, the neck is more or less stretched, the chin held forwards, not pulled in. The limbs and often the tail are stretched out freely. The more arboreal felids, as f.i. Panthera pardus and Neofelis nebulosa, often display their habit of letting their extremities hang down from the tree branches they are lying on. The animals need not actually sleep, but may show some – not very intensive – activity, such as yawning, washing, looking around or straight ahead, blinking, stretching one paw etc.

In the actual sleeping postures, muscle relaxation is still more complete. The position of the body may vary greatly, the primary aim being comfort. In nearly all cases, a certain openness of posture is recognisable. Frequently the particularly sensitive and vulnerable parts such as the nape, the throat or the belly are exposed, the back stretched. The extremities are free and often stretched also. Without doubt the clearest example of a relaxed sleeping position is lying on the back; this occurred in all felids I could observe at being totally at ease, but only the lion shows this sleeping posture as a typical routine.

b)Defensive sleep, resting under tension

Characteristic of this is that the animal looks uneasy. It is often unclear whether it is really sleeping, or only trying to or feigning sleep. This basic category must also include the exhausted sleep which follows considerable or long-lasting tension and which after a while can gradually merge into recuperative sleep. The sleep or doze of sick animals also belongs in this category.

Characteristics of defensive sleep

  • Face: The eyes are tightly closed, often in a noticeably cramped manner, which usually results in the formation of skin folds, especially at the inner corners of the eyes, sometimes extending over the bridge of the nose.

  • Position of the ears: All forms of threatening and defensive positions occur, and occasionally that of attentiveness (see LEYHAUSEN 1979, Fig 79). The intensity is much slighter than in a waking state and is dependent on how irritating or menacing the animal finds the situation to be.

The threatening cat keeps its ears pricked and turned outwards, so that the back side of the ears is visible from the front; in most felid species the back side of the ears shows special markings to enhance the gesture (Examples: Black and white in the Lynx, Neofelis and some of the Panthera species, black in Panthera leo and Profelis caracal, bright orange in Felis libyca).

Disturbed cats turn their ears sidewards to backwards and flatten them against the head when really frightened.

Pricked ears, turned to the front, indicate that the attention of the animal is aroused.

During sleep, the ears are frequently moved and directed towards a source of sound. The eyes, however, are seldom opened, but rather squeezed, even more tightly shut if the disturbance is particularly intense.

A special form of displayed nervousness and uncertainty shown by a constant flickering of the ears I have observed, up to now, only in Profelis caracal; nevertheless, it also is a sign for defensive sleep in this species.

  • Fur: under extreme stress, the fur of the tail, and sometimes even of the lower back, stands on end.

  • Body posture: The head seldom lies; mostly is held in a somewhat cramped, semi-upright position. Vulnerable parts of the body (nape, throat, stomach) are protected by pulling in the chin and the neck and curving the back to a crouched or rolled-up posture. In addition, the extremities are tucked in or held close to the body. If the tail is not drawn under the body the tip often moves restlessly.

    Observations on adult Cheetahs (Acinonyx jubatus) reveal that this long-limbed sprinter is quite reluctant to fold his legs under its body, so it always rests in a stretched-out posture. Still, a possible feeling of uneasiness is easily recognisable by the pulled-in head and the chin drawn towards the breast. The tell-tale ear positions are obtained more markedly as in most other defensive-resting felids.

A typical and rather striking sign of tension and uneasiness in a felid is a cramped closed crouch on its belly, with the legs folded underneath and the tail drawn close to the body. If the facial expression still remains relaxed and the head points towards the source of stimulus, the animal might only demonstrate a feeling of uncertainty. If the head is turned away, the cat certainly and deliberately refuses to face a disturbing or unnerving situation. Sometimes the “cut-off” (CHANCE 1962) is so complete that there is little or no reaction to stimuli from the surroundings. In such cases, the stress and the suffering is considerable though unrecognisable for the untrained observer. When, in addition, the ears are turned outwards and sidewards and stand away from the head, and parts of the fur are bristled, especially at the back of the head, the cat very probable is severely ill, and/or in pain.

The body posture of exhausted animals may, surprisingly, correspond to the more relaxed lateral type with the extremities extended; in almost all cases, however, the face, the position of the ears, and the occasional tail movements as well, reveal moderate to strong tension and unease. Breathing is usually accelerated and the pupils may be dilated.

c) Special forms of sleep

Sometimes, a tightly closed sleeping posture may be deceptive. A good example is the “frost sleep” of Felis silvestris, which occurs when it is very cold but, occasionally, for other reasons as well, and which TEMBROCK (1978) has also mentioned, claiming that during sleep other behaviour patterns also become activated, f.i. regulation of thermal loss by change of position.

The “cold weather sleep” is easily identified as it can only be observed in late autumn and winter at sub-zero temperatures, and, well, sometimes also when the weather is not so cold after all. The cats adopt an extremely closed posture. They cower in as round a shape as possible with the extremities tucked in and belly downwards, or, at lower intensity, on their side. A typical characteristic, which does not occur in the other types of sleep, is to protect the forehead by laying it on the ground.

Despite this extremely closed posture, the animals give the impression of being at ease. They do not react to disturbances and show no sign of readiness for defence or aggression. Occasionally, when it is very cold, a component of uneasiness enters the picture, indicating that extreme cold might be a stress factor per se for the animal.

Besides in Felis silvestris, I have seen frost sleep (in a lower-intensity but nevertheless very typical form that was totally free of any sign of tension) also in my own two Felis libyca.

Neofelis nebulosa also often can be seen sleeping curled up in a tight ball – mostly during their prolonged sleeping phase around midnight. These very arboreal cats sometimes seem to prefer flat troughs on loose ground to tree branches, but very rarely during visiting hours, as they clearly feel safer when staying high up in a tree, well camouflaged and thus practically invisible amongst the leaves. Anyway, when I observed a Clouded Leopard resting or sleeping on the mulch-ground in a rolled-up position, its facial expression looked relaxed. The whole set-up (rainy late-autumn weather with temperatures just over 0°, open doors between outside and inside enclosure, floor-heating under the mulch) during the 2 weeks of my observations led me to the conclusion that this species also shows something similar to the Wild Cat’s “frost sleep”, although its really striking feature, the turned-down forehead, is amiss. FURTER (1991) who observed some Clouded Leopards over prolonged time, denied that the closed-up sleeping posture had anything to do with stress situations, but remarked that the cats never obtained any sort of open, stretched-out resting postures when lying on the ground during visiting hours – only at night. At present, it remains a question whether or not this type of sleep contains a component of defensive sleep. If strongly stressed, however, the Clouded Leopards very readily adopt the closed crouch on the belly.


On seeing a sleeping feline it is often assumed that “the animal is sleeping, therefore it must feel well”; “well” in the sense of “relaxed”, “reassured”, or “safe”. In fact, however, defensive sleep is only an attempt by the animals to evade the considerable social stress which the visitors cause by merely looking at them. Further causes, as shouting and screaming, wild romping (school classes) can only intensify this problem, as stress factors have a particularly strong effect in combination. Above all, it is a fact that internal stress (wrong set-up of enclosure, space problems, disturbance by other animals) can be strongly intensified by the addition of external stress. As the mere quantum, the quality and the combination of different stress factors has an influence on the inclination of an animal for defensive sleep, as well as the animal’s individual character itself, it is not easy – and, by the way, not particularly rewarding – to quantify observations by, say, counting the volume of visitors and correlating this with the number of hours an animal spends in defensive sleep.

It is much more important to recognise behavioural disturbances whenever they occur, and to react to it. Defensive sleeping and its further problems (as f.i. apathy, agonistic and faulty reproductive behaviour, lowered resistance to all kinds of diseases), might not be eliminated but would at least be alleviated by the provision of freely accessible hiding places. The objection that, then, the cats would simply disappear from the scene all day proves not to be valid; after only a short period of adaptation the animal will come forward, driven by its invincible curiosity. The advantage will be on both sides: Instead of an immobile, shapeless bundle of fur the visitor sees a feline, beautiful in its movements, delightful when it plays with a toy – and possibly putting the stressed spectator at ease indulging in a restful sleep.

Other possible ways of protecting zoo animals from stress caused by the public would be to ensure that they are not so aware of being stared at. Big glass panes, as already used nowadays, should be fitted at an angle so that the reflections hamper the vision more for the cat inside than for the visitor outside. The visitor should as far as possible be able to watch the animals from a darkened place and thus not be so obvious to them. For looking into hiding-places, or in indoor cages, even one-way glass panes could be used.

One of the worst set-ups even in modern zoos is an enclosure accessible for visitors from two opposite sides. To avoid defensive sleep, and other problems as well, the enclosure then should be enormous.

Space, in general, certainly is the best means to make a feline feel safe and at home; sufficient depth of the enclosure gives the animal a feeling of comfort, knowing it can get away from the uncanny visitors’ side whenever it likes to. Alas, space is, for various reasons, not always easily obtainable. Good enclosure fittings, however, will more than make up for a possible lack of space. Cover should be the most important feature in any cat enclosure. A skilfully furnished enclosure will give the animal a perfect sense of protection without obstructing the visitor’s view too much. Beside the fact that the cat will be more inclined to come forward and display its normal activities, its appearance in a well structured “landscape” enclosure is by far more delightful to watch.


I should like to express my sincere gratitude to the late Prof. Paul Leyhausen for the encouragement to expand my investigations on the “Verteidigungsschlaf”, Barbara Leyhausen-Tonkin and Mary Collett for their unselfish help, and all the zoo-institutions, directors, curators and staff for their hospitality, their interest in this project and their co-operation.


Chance M.R.A. 1962: An interpretation of some ontogenistic postures: the role of “cut-off” acts as postures. Symp.Zool.Soc.London 8, 71

Ewer R.F. 1968: Ethology of Mammals. London

Furter, M. 1991 : Aktivität, Gehegenutzung und Verhalten von Nebelpardern Neofelis nebulosa (Griffith 1821) im Zürcher Zoo. Thesis, unpublished.

Hassenberg L. 1965: Ruhe und Schlaf bei Säugetieren. Neue Brehm Bücherei Bd. 338, Wittenberg Lutherstadt

Immelmann K. 1962: Beobachtungen über Schlafrhythmus und Schlafverhalten an drei afrikanischen Straußen. D.Zool.Garten NF 26, 215

Leyhausen P.1979: Cat Behavior.

Pfleiderer M. 1990: Zum “Verteidigungsschlaf” von Carnivora im Zoo. D.Zool.Garten NF 60, 228

Scherzinger W. 1970: Zum Aktionssystem des Sperlingskauzes (Glaucidium passerinum L.). Zoologica 41,1

Tembrock G. 1978: Verhaltensbiologie. Stuttgart

Thaler E., Ettel E. & Jakob S. 1981: Zur Sozialstruktur des Waldrapps, Geronticus eremita – Beobachtungen an der Brutkolonie des Alpenzoo Innsbruck. J. Orn. 122,109

Karoo Cat Research
cards4bliss - Karten zu Glück, Freiheit und Wohlstand