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Management of Wild Cats In
Captivity
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Taken from the Symposium 4 Proceedings
31 March 1979
Edited by Jon Barzdo
CONTENTS
THE HOUSING AND BREEDING OF BIG CATS AT MARWELL ZOOLOGICAL PARK
JOHN M. KNOWLES, Director, Marwell Zoological Park
Housing Designs
Two basic considerations for housing felines at Marwell had to be human
rather than animal orientated - a principle I deplore, but one which never-
theless is forced upon zoo directors where necessity puts them in the unhappy
position of having to finance their operations from money received "at
the Gate". These considerations were: firstly the need for relatively
low cost: secondly, the need to blend buildings into an English, Park
landscape.
The second consideration fitted well with the first, in that both called
for simple structures.. Thus, ideas of features desirable in a city zoo, such
as completely indoor viewing facilities, moats, and a large housing complex
showing many cats in close proximity to each other, giving the visitor the
benefit of comparisons between species were eliminated. It was also felt that
a large cat complex was undesirable from a disease control aspect.
The third, and overwhelming consideration was that, within the limits of
the first two points, we had to give our animals accommodation that would
satisfy both their health and behavioural requirements.
The apparent conflicts of these considerations weighed heavily on my mind
during the planning stages of Marwell, but, like so many problems, the
realisation was easier than first appeared. Whilst our initial solutions may
not have universal appeal, they have nevertheless stood the test of time (or
at least seven years of time) and, were I to start afresh, I would not build
in a manner vastly different from that of our present structures. Nothing in
our approach is, or was, very novel and none of it will win a design award'.
The indoor accommodation is, frankly, based on the concept of a circus
"beast-wagon", for I have always been impressed by the physical
health of cats in this type of accommodation, which is essentially warm and
snug without being stuffy - although I dislike seeing animals confined solely
to such accommodation.
We therefore built firstly for the Siberian tigers a house, the
specifications of which are as follows:-
8 inside dens with wooden floors, 0.68 metres above ground level, each with a
weldmesh front with a width of 2.135 metres. The depth of these indoor dens
is 3.1 metres, and the roof height (which is varied because of the sloping
roof) averages 3.5 metres. In front of the dens is a service passage 1.4
metres wide and public viewing windows of armour-plated glass look across
this passage, enabling visitors to view the animals from outside, under the
cover of an overhanging roof.
For the outside runs there was scope to provide for the behavioural needs of
the cats. With the tigers I regard space as important, as during the 24 hour
period, contrary to some popular conceptions, they use the whole of their
"territory" and are by no means stereotyped in their movement
pattern.
Most important in any outside enclosure is the provision of the right type
of "furniture" for the needs of the inhabitants. For tigers, the
essential item is a pool. Our tigers have so much pleasure from water that it
is sad to see Panthera tigris sub-species without such a facility.
Not only are the pools used for swimming and cooling, but for a great deal
of play activity by various interacting individuals, sometimes including both
adults and juveniles in our family group, for we have always been able to
maintain one adult breeding male, two females and their progeny (i.e. two
litters) in one integrated unit.
One outside run for the tigers has a perimeter fence of 150 metres in
length and the other two have lengths of 143 and 131 metres respectively. All
the outside runs have concrete perimeter paths, as heavy animals like tigers
create mud in wet periods just as much as hoofed stock. The fences are 4.5
metres high with an overhang of one metre. These lost dimensions are probably
greater than they need to be, in that never once has a Siberian or Sumatran
tiger shown the least interest in climbing or jumping and, to some extent,
the fences detract visually from the appearance of the enclosures. However,
had we used moats instead of these fences, there is no doubt that the visual
appearance to the visitor would have been enhanced, but the tigers themselves
would have had less space.
Similar sized dens, though smaller in number, are used for leopard,
jaguar, cheetah and snow leopard. The outside pens, except that of the
cheetah, each have a roof, but here the similarity ends. Behavioural needs
are catered for with, for instance, ledges for the snow leopards, high enough
for them to overlook the visitors, and trees inside the runs for the
leopards, thus enabling them to assume their favourite resting position, with
legs draped on either side of branches, as well as facilitating climbing
activity.
I have already indicated that I regard space as important, but still more
is the quality of space. An area the size of a football field, with no trees or
suitable substitutes, would be as unattractive to leopards as a vast, bare
room would be to human beings.
The lynx are housed in a mini-version of the Rotterdam Kennel system, with
their outside run furnished with low logs and hazel coppice, as well as grass,
growing in the enclosure. Native hazel is the only vegetation other than
gross, which we have found consistently able to withstand the play and
claw-sharpening activities of cats.
Diet
Virtually all our feeding to felines is beef on the bone, with an added
supplement powder, but the snow leopards have an occasional rabbit, without
the liver, and the lynx an occasional chicken. Calf is sometimes fed, but
beef remains. the- preferred diet.
Routine vaccination for Feline Enteritis is done annually with Felocine.
Breeding
In 1970 we began with young animals of the species Siberian tiger, leopard,
jaguar, lynx and cheetah, and other species have been added to the collection
more recently.
To date, our total failure has been with cheetah, although we have tried
to copy the Whipsnade system, and have seen some mating pre-play, but have
observed no matings. Our only crumb of success was that a mate who went to
Whipsnade, having achieved nothing in six years at Marwell, fathered a litter
within four days of arrival there.
Similarly, snow leopards are yet to breed with us, but as our only male is
still young (four years) and has some personality problems which I believe
arise from its having been involved in behavioural studies, I am by no means
unhopeful for the future.
Siberian tigers
From the two females already referred to, thirteen (from Nimana) and eleven
(from Amoga) cubs have been born to date, and of this total of twenty-four,
eight did not survive to weaning. The first pair of tigers received into the
collection did not breed, both of these being hand-reared specimens, and our
first Marwell-born male, who was also hand-reared, has not bred to date. In
both cases the male, although copulating frequently, fails to penetrate the
female and I wonder how common this is with hand-reared tiger males. With
hand-reared leopard and jaguar males, breeding success has been achieved.
Jaguars
Nineteen jaguars have been born to date, to two females with one male, and
all but four survived. Only one hand~rearing has been necessary. The first
pair, although compatible, failed to breed until the male mated with a female
with whom he would live peaceably only when she was in oestrus. it seems that
the stimulus of the other female having a cub in an adjacent enclosure, induced
oestrus in the second female after two years of non-breeding, and she then
became a regular breeding female. This pair is now in Belfast Zoo, having
been replaced in our collection by a young black female received from Rotterdam,
and a male from Chester.
Leopards
Twenty-three born, of which four did not survive, and one animal was hand-
reared. The only remarkable incidents were two cases of milk fever in two
different nursing females at different times. In both cases rapid response
was achieved after the administration of Calcium Boro-glucanate via dart.
Lynx
Eighteen born, of which all but three survived. The original male was wild-
caught and the female captive-born in Rumania.
The first female and one of her daughters, together with the original male, were
excellent breeders, but the original pair have died and we now have a young
male from the Aspinall collection, born in 1978, so breeding is not expected
for some time.
Serval
From a captive-born pair received from a European living in Uganda,
nine cubs have been born to date, of which four did not survive.
Asiatic Lion
1.1 in the collection received from the East Berlin Zoo where they were bred
on 6.4.78, at which time the male was one year old and the female three years
old. Mating activity has been observed during 1979, but is probably not
likely to be successful because of the relative youth of the male, for at
least another year. Similar housing to that of the Siberian tigers is planned
to be built during 1979 for these animals.
GRAHAM LUCAS, Senior Keeper, Cats, Whipsnade
Park, Zoological Society of London.
Before looking at the present day housing of cats, one should look back at
earlier enclosures in order to appreciate how knowledge gained of the
requirements of the animals has catered attitudes to such housing.
The first recorded forms of housing for members of the cat family in Europe
were, perhaps, the enclosures used in Ancient Rome. As for back as the third
century B. C. , felines were housed in simple stone cells with barred doors,
although quite complicated systems of gates and passageways must have been
used when the animals were required for displays in the arena.
Over the centuries the housing changed little, Even in the Dark Ages,
princes, barons and the like kept cats as prestigious pets. The cats, mainly
leopards and lions, were housed in very cramped and insanitary quarters,
usually in the lowest part of a castle. Gradually, however, the animals came
to be kept at ground level. Special quarters were built for them and the
barred door evolved to become larger until it became the cage front.
In the 12th century, Henry I of England
developed the Royal Menagerie at Woodstock,
Oxfordshire, and among the animals housed there were lions, leopards and
lynx. How they were housed there is not known, but when they were moved to
the Tower of London,
in 1252, they were kept in what was to become known as the Lion
Tower. An engraving of the Lion
Tower, mode in about 1820, shows
the animals housed under conditions very similar to the early stone cells. At
this time there were, of course, other menageries in Europe,
for example in France,
Portugal, and
Italy. Others
were to follow, but the methods used for housing the large cots, which were
generally those kept, were normally of the bar-fronted-den type. However, an
interesting illustration of the zoo at Belvedere,
Austria, which was
founded in 1716, shows a lion in a grassy outdoor enclosure with sleeping
quarters at the rear.
In the 19th century came a great zoo boom and, with the boom, living
conditions for wild animals generally were greatly improved. Perhaps this was
because these newer zoos were public zoos, rather than the status symbol of
some potentate, and people were paying to see the animals. People who were
paying wanted to see the animals clearly, so cages became bigger, although
not necessarily better. It become the order of the day to house members of
the cat family under one roof and, for some reason, these exhibits became
known as Lion Houses. They consisted of a number of indoor cages, usually
linked to outside barred enclosures. Sometimes separate sleeping dens were
provided, which were also used by pregnant females to have their cubs, giving
three enclosures in all. Perhaps the best of these Lion Houses was that built
at Regent's Park, London, in 1876
which, because it was originally built on a grand scale, was still adequate
for housing cats when it was demolished nearly one hundred years later. Early
in the present century come what was to be one of the greatest innovations in
the housing of cats - the use of a moat instead of bars to enclose the
outside area. Before opening his zoo at Hamburg
in 1907, Carl Hagenbeck carried out experiments to find out how high big cots
could jump. By suspending pigeons in their cages, he discovered that lions
and tigers could leap at least 12 feet high. Their long jump abilities were
also tested. Enclosures were then designed with moats approximately 30 feet
wide and for the first time visitors were able to see big cats without bars
obstructing the view. Many other zoos soon copied these ideas, some with very
wide moats, but for some reason the new idea didn't seem to catch on in Britain,
for only Edinburgh, and more
recently London, have moated lion
enclosures.
The backgrounds to the Hamburg
mooted enclosures are, in the main, man- made cliffs and the floors either of
sand or concrete. It was not until Whipsnade
Park, Bedfordshire,
England, was opened in
1931 that lions, and later tigers, were seen in grassy enclosures with trees
and bushes growing in them. These naturalistic enclosures featured something
that, in its own way, was almost as revolutionary as Carl Hagenbeck's moats -
the complete lack of artificial heating in the sleeping dens. The old Lion
Houses were well heated as it was thought that animals from tropical
countries could not withstand the European climate, especially in winter.
Even at Hamburg, where
acclimatisation experiments had taken place, the animals had (and still have)
heated quarters for their use during the winter months. But at Whipsnade, the
animals were given sheltered, drought- proof dens which they could enter at
will and where, for most of the cats, no heating was (or is) provided.
Other British zoos, particularly Dudley, quickly followed the Whipsnade
lead. But many zoos now use chain-link fencing instead of bars to enclose the
animal area. (Chester Zoo is believed to have been the first to employ wire
mesh to enclose lions). By making the enclosure much bigger and laying a road
through it, Jirnmy Chipperfield opened the first European- drive-through big
cat enclosure at Longleat, Wiltshire, in 1966 and the safari park was born.
For the smaller cats, the lion House concept, with somewhat smaller cages,
was used fairly extensively and it was for these smaller felines that wire
mesh was first used. At a later date, in some small cat enclosures the mesh
was replaced with glass. Glass is now fairly common in houses for small cats,
but it has only recently been tried for lions and tigers.
So much for a look at the history and evolution of the housing of cats,
but what of today?
In Britain most of the small barred cages have been demolished, some being
replaced by more naturalistic enclosures, whilst others seem only to have
been replaced by wire mesh. For most lions and tigers today the typical
enclosure is a grassy area, with growing trees and bushes in some instances,
surrounded by a high wire-mesh fence, with sleeping dens attached, as, for
example, at Blackpool, Chessington and Marwell. Most other cats, because of
their leaping abilities, are normally housed in wire-mesh aviary-type
enclosures with adjoining dens. Because accommodation for these cats
generally has to be roofed, they tend to be much smaller than lion and tiger
exhibits. At Chester there is, however, an open-topped exhibit housing
serval. Calderpark Zoo, Glasgow, and Chester Zoo both have modern cat houses
for felines of leopard size and smaller. In both cases the indoor quarters
are not on show to the public, and face into a central service passageway.
Both houses also feature glass-fronted outdoor enclosures. Cheetahs at most
British zoos tend to be housed in similar enclosures to those for lions and
tigers but, as these animals tend not to jump very high, the enclosing fence
is generally much lower.
It appears that, in Britain at least, the Lion House is a thing of the
past. Zoo architects now seem to have a complex about cat-housing and so
instead of cat houses they design cat complexes. Two have been built during
the past few years, one at Bristol and one at London.
The Bristol complex consists of five hexagonal aviary-type enclosures in
two of which glass panels form part of oneside of the enclosure. The sleeping
dens are not on show to the public, a trend which has gained popularity of
late. The complex is for cats of puma-size and above, although the lions have
their own wire-rnesh enclosure in another part of the zoo.
The London complex - The Lion Terraces - consists of seven enclosures for
cats ranging in size from caracals to lions. With, the exception of the lion
area, the enclosures are of the wire-mesh aviary type, but instead of being
square or rectangular, as in most zoos, the animal areas are of a
free-flowing design. The lion enclosure is open-topped and moated, the moat
being water-filled. (The moat at Edinburgh is dry). London also employs
glass, not only in the three indoor dens on show to the public, but also in
most of the outside enclosures, where there are glass panels forming part of
the perimeter fence. In fact, at the lion area, the visitor can view the
lions over the water moat, through mesh or through glass. London's Lion
Terraces are some of the best landscaped enclosures for cats, not only in
Britain, but in Europe. Each enclosure was designed with one species in mind;
for example, the caracals have a desert habitat and the jaguars a forest or
jungle-like habitat.
The Lion House concept might well be dead in Britain, but it is still
going strong in the rest of Europe, although moated enclosures for lions and
tigers are common. Traditional Lion Houses con be found in many continen- tal
zoos. The one at the Menagerie du Jardin des Plantes, Paris, features the old
style barred enclosures, whilst more modern buildings, such as that at
Wuppertal, Germany, have outdoor enclosures.
Other zoos, such as those at West Berlin and Frankfurt in Germany and
Arnsterdam in Holland have moated enclosures attached to their carnivore
buildings. At Frankfurt and Amsterdam however, the houses themselves are
fairly old and the moated enclosures are recent additions. At West Berlin the
situation is reversed with a modern house attached to a pre-war moated
enclosure. The enclosure at Amsterdam is modernistic, whilst that at
Frankfurt is naturalistic, thus, they show quite well the two extremes of
this type of exhibit. One zoo, East Berlin, has moated enclosures for lions
and tigers both outside and indoors, although other cats are housed in more
typical enclosures.
I have concentrated mainly on the big cats, and the small cats have had
hardly a mention. This actually reflects the situation in British zoos and in
zoos worldwide, as very few zoos have many of the small cats on display.
Apart from pumas, serval and lynx, which all tend to be displayed in
aviary-type enclosures, it is not often that one finds exhibits of other
members of the genus. The cat houses at Chester and Glasgow zoos have already
been mentioned and, at present, these are the two main displays of small cats
in Britain. Continental zoos with good collections of the smaller cats
include Wuppertal and Rotterdam, where they are housed in quite conventional
accommodation. However, Rotterdam's house does have one innovation. The
sleeping dens are movable and can be used as transport crates.
These then are some of the ways cats are housed in Britain and Europe. It
would seem from just this brief look that the days of the barred cage are
numbered and there are, I think, two reasons for this. The first is cost -
iron bars are more expensive than chain-link netting, although they do last
longer -and the second is public opinion. Iron bars might not make a cage as
far as poetry is concerned, but to the general public they do. Of course, to
the animal it makes little difference whether it is confined behind bars,
netting, glass or a moat.
It is questionable whether moats are the ideal barrier for some of our
smaller urban zoos. For, to be effective barriers, moats have to be at least
twenty-five feet wide. There is no need, I feel, to go as far as the Parc
Zoologique, Paris, which has a forty feet wide moat. This wide area is of no
use as living space for the animal and, where space is at a premium, it
seriously decreases the area available for the cats.
One thing noticed with most water barriers, is that during the winter
months, when the water freezes over, the moat becomes useless as a barrier
and the enclosure is therefore useless as there is insufficient height from
water (ice) level to the top of the moat wall. Lions and tigers could just
walk over the ice and jump out of the enclosure. Some zoos can, of course,
solve this problem by lowering the water level, but at others, even if the
moat were to be emptied, there would still be insufficient height to act as
an adequate barrier. So, for northern European zoos at least, a great deal of
thought must be given to the design of the moat as it is little use having an
enclosure that is usable for only part of the year.
Glass is more expensive than wire mesh, but cheaper than a moat and is
very effective as it can give the enclosure a bar-less look, and should prove
useful where there is a shortage of space. However, care must be taken to
ensure that the glass is kept clean, otherwise this effect is spoilt,
References (used but not cited)
Fisher, James (1 966) : Zoos of the World, pp 41,50,166
Jennison, George (1928) : Noah's Cargo, p97
Joans, June (1969) . Zoo Without Bars, pp 37-39
D. G. ASHTON and D. M. JONES,
Veterinary Officer and Senior Veterinary Officer, Zoological Society of
London.
Introduction
The 36 species of non-domestic cats
are anatomically and probably physio - logically very similar to the domestic
cat. They all have 38 chromosomes, except the small South American species,
which have 36. They are basically solitary animals, only the lion being truly
gregarious. They appear to be susceptible to the some infectious diseases as
the domestic cat, but many of the veterinary problems encountered in
captivity ore related to management factors such as nutrition, housing and
abnormal behaviour.
Nutrition in Captivity
The cats are the most carnivorous of the carnivores. There is evidence from
the domestic cat that the adaptation to a purely carnivorous diet is so for
advanced that some of the metabolic pathways found in other carnivores are
absent in cats. Artificial diets designed for the domestic cat are often not
well accepted; however, in the USA commercial "exotic cat" diets
are used successfully. In Britain and Europe most cats are fed on meat or
whole car- cases. Whereas whole carcases are a natural, complete and balanced
diet, meat, and selected parts of carcases are not. (While carcases of some
artificially fed laboratory animals may be deficient in certain minerals, so
that it may be advisable to supplement these (P. leyhausen, pers. comm.)).
Severe disease has been caused, particularly in young growing animals, by the
feeding of unsupplemented parts of carcases. Meat (i.e. muscle tissue) is the
most commonly fed and this has a very low calcium to phosphorous ratio.
(1:20, the optimum is 2:1), together with low levels of Vitamins A and D.
Nutritional bone disease which results in greenstick fractures and
deformities can occur on such a diet. Supplementation of meat with calcium,
or the feeding of carcase parts containing a high proportion of bone, may
correct or improve the Ca/P ratio, but does not correct the vitamin
deficiencies. The Vitamin A requirement of the domestic cat has been shown to
be 1,000 - 2,000 i.u. per day (equivalent to 250-500 iu/kg, or approximately
80-160 ug/kg retinol per day (Scott et al, 1967). Experimental work to
determine the requirement in non-domestic species was carried out by J.M.
Hime (Regent's Park Hospital) and the evidence suggests that approximately
350 iu/kg per day (100 ug/kg) is adequate.
Vitamin D2 or D3 requirements are
approximately 5% of this, i.e. 17 iu/kg per day. The requirements for other
vitamins and minerals are unknown. Deficiencies of Vitamin E (especially on
diets high in fat, e.g. meat heavily supplemented with cod liver oil) have
been reported. Meat tends to be low in iodine and copper and it is probably
advisable when feeding meat to include these nutrients in the supplement
used.
Excessive levels of calcium in the
diet may interfere with the absorption of Mn and Zn, and deficiencies of
these two elements have been, reported (Wallach, 1970). Milk is low in iron
and iron deficiency anaemias may occur in suckling animals (Theobold, 1978).
Diets high in offal, particularly liver, have produced Vitamin A poisoning in
domestic cats and should be avoided. Adverse reactions of tigers particularly
to horse meat have been reported (Jones, unpublished observations; Theobald
op.cit..)
Fresh meat is subject to bacterial
overgrowth, and decomposition even when refrigerated at 40C. It should not
therefore be stored for long periods before being fed, unless deep frozen at
temperatures below -150C. Once removed from the refrigerator it should be
consumed within 24 hours.
Preservatives used in the
manufacture of certain semi-moist domestic animal foods may be harmful to
cats in view of their limited ability to detoxify and metabolise certain
compounds.
Quantities fed should range from
180g to 6.5kg depending on species, activity and size. It is important that
the quantity fed is adjusted to the condition of the animal. There is never
any excuse for obesity and animals should not be thin because of inadequate
calorie intake.
Housing
This subject has been dealt with in other papers. However, certain veterinary
aspects are listed below:-
1. Adequate area should be provided for exercise, and as a guide the
formula 1 square metre/kg body weight can be used as a minimum
requirement. Less than half this should be considered inhumane. The
exercise area should be surfaced with a relatively nonabrasive material
to prevent the development of sore pads in individuals which
persistently pace. For many species, the provision of items of interest
in the exercise area for, climbing, jumping etc., is as important as the
dimensions of the enclosure.
2. The house/den should be dry, drought proof, and have a well insulated
lying area.
3 . There should be proper drainage of both house and exercise area, so
that excreta can be removed easily.
4 . Provision should be made for easy observation of all parts of the
house and exercise area.
5 . There should be a restraint facility strategically sited so that the
animals are accustomed to passing through it routinely.
6. "Escape routes" and "introduction pens" will be
necessary for the successful breeding of many species with minimum risk
of injury.
7. There should be some provision for hospitalisation and the isolation
of sick individuals from other members of the group.
8. Attempts should be made to prevent access of domestic cats, which may
carry infectious disease.
The Dangerous Wild Animals Act
applies to all non-domestic cats kept in private collections, and stipulates
that in addition to proper provision for the animals' welfare, there should
be good security against escape.
Handling
The smaller species and all young animals can be caught in nets or handled
with thick gloves. Although the larger species can be immobilised with a f
lying syringe, it is preferable, especially where treatment with a course of
injections is required, to have a crushing facility built into the
accommodation. This should be sited so that the animals are used to passing
through it every day, and so designed that the animal is unable to tell
whether or not it is about to be caught. Smaller species con be sedated or
immobilised with the alphaxalone/alphadolone mixture "Saffon"
(Glaxo), either intramuscularly, or intravenously if sufficiently restrained;
the larger species with Ketamine (Vetalar, Parke Davis), preferably mixed
with Diazepam (Valium, Roche), to counteract the convulsive tendencies of the
former. Wiesner (1977) reports the use of a mixture of Ketamine and Xylazine
(Rompun, Boyer) which allows a lower than normal dose rate of both drugs, and
a much smaller volume injection than with the Ketamine/Valium mixture. Trials
with this mixture in varying proportions at Whipsnade and London suggest that
at certain dose rates it is preferable to phencyclidine (Sernylan) in that
recovery times are faster, relaxation is better, and there is less risk of
convulsive episodes. Once immobilised, conventional gaseous and intravenous
anaesthetics con be used.
Breeding
Approximate gestation periods are 100 days for the larger species, 90 days
for the intermediate sized, and 65 days for the smallest. The recorded range
for the family is 55 days in the case of the African wild cat to 115 days for
the African lion. Most are seasonally polyoestrus, and by extra- polation
from the domestic cat are thought to be induced ovulators. litter size ranges
from 1 to 6, usually 2 in the smaller species and 3-4 in the larger. Most are
relatively easy to breed, but the practice of keeping pairs. of animals
together for long periods often inhibits mating activity. In the wild males
and females of the non-gregarious species in breeding condition usually come
together only to mate. The initial courtship and post-coital periods can be
dangerous for both sexes, and in captivity males and females of unknown
compatibility should be introduced gradually before being enclosed together.
There is a tendency in some species, after mating, for the female to attack
the male and drive him off; in such cases a suitable escape route should be
provided. Where accommodation and management practices are designed to allow
for such behaviour, the risk of serious injury is much reduced.
It is important to minimise
disturbance in the vicinity of a female with cubs. Unaccustomed noises,
people, and activity can result in neglect of the litter, or cannibalism.
Hand-Rearing
There appears to be a wide variation in the milk composition of different
species, and although a standard milk substitute based on cows' milk can be
successful, it is probably better to use one of the commercially produced
formulations which approximate to the natural composition. Table 1 shows the milk
compositions of two non-domestic species (Ben Shaul, 1962) compared with
those of domestic cat and cow. These figures may be based only on one or two
samples at an unknown stage of lactation.
Most species will begin to take
solids at about 3 weeks, and may be weaned by 10-12 weeks.
Contraception
Breeding has been so successful in some species (particularly lions) that
various methods are employed to limit reproduction. Vasectomy of breeding
males, hormonal implants, injections and tablets, separation of oestrus
females, and neutering of both males and females have been used. All have
some drawbacks and the method of choice will depend on the circumstances.
Infectious Diseases
Viral Diseases
Feline Panleucopenia
Also known as feline infectious enteritis, a highly infectious fatal disease
which is still relatively common in domestic cats. The symptoms include
marked depression, loss of appetite, and vomiting. Diarrhoea may develop
after several days. The number of circulating white blood cells are pro- foundly
depressed (hence the disease's name). Infection in pregnant females con
result in young being born with brain damage which results in abnormality of
movement. The virus responsible is very resistant and may survive in infected
premises for several months. Recovered animals may continue to be infectious
to other cats for long periods, There is some evidence that the South
American species are less susceptible to this disease. Infection frequently
occurs when animals are being transported and stress undoubtedly makes
animals more susceptible. Animals should there- fore be vaccinated before
transportation or similar stress, not during or after .
Two types of vaccine are available,
attenuated live virus and killed virus preparations. The former stimulates a
stronger and longer lasting immunity with one injection, than the latter,
which requires two. Although live virus vaccines have been used in several
large species of cat with no ill effects, there have been reports of illness
in the smaller Felidae, including the European wild cat (Felis sylvestris)
after the use of attenuated live virus vaccine (Scott, 1979). Pregnant
females of all species should be vaccinated with a killed vaccine.
Vaccination of young animals should be left until 3 months of age when there
is little danger of infection. If vaccination prior to this is necessary,
further vaccination at 12-16 weeks must be given in case maternal antibodies
circulating in the youngster have inactivated the initial dose. Povey and
Davis (1974) suggest that the use of increased doses of killed vaccine in
large cats is of doubtful value.
Feline Respiratory Diseases:
Two main viruses are responsible for upper respiratory tract diseases in the
cat (sometimes called "cat 'flu"), feline rhinotralheitis virus and
a cal- civirus. Symptoms include sneezing, nasal discharge, conjunctivitis,
mouth ulceration, a raised temperature, depression and loss of appetite. In
severe cases the loss of appetite can eventually result in death. As with
feline panleucopenia, domestic cats are probably the main source of
infectection, but the carrier state almost certainly exists in non-domestic
species too. Vaccines are now available, although their safety and
effectiveness for many of the non-domestic species is not proven.
Feline Viral Lymphosarcoma and
feline infectious peritonitis
These are two other viral diseases of the domestic cat which can infect some,
and perhaps all, non-domestic species.
Cowpox
There have been four outbreaks of this disease in non-domestic cats, two in
Britain. Symptoms vary from an acute pneumonia, with death in 2-3 days, to a
more prolonged disease with numerous ulcerated nodules in the skin and
ulceration in the mouth. Some of the animals with skin lesions recover. The
disease is infectious, but the source of the infection in Britain is unknown.
Vaccination with smallpox vaccine is of doubtful value. The disease con be
transmitted to humans. It is essential to isolate suspicious and infected
cases immediately to try to limit spread to other animals.
Rabies:
This disease is unlikely to appear in captive non-domestic felines in Britain
because of quarantine restrictions.
Bacterial Diseases
Salmonella:
Salmonella infection can be acquired from contaminated raw meat. The symptoms
range from mild diarrhoea, to severe gastroenteritis with generalised
infection. Some infected animals may appear unaffected. The disease can be
transmitted to humans.
Tuberculosis
Tuberculosis infections are usually
acquired from infected food. Wild birds are a potential source of Mycobacterium
avium The low incidence of bovine tuberculosis in Britain, and the feeding of
meat from carcases that have been properly inspected has made tubercular
infections in cats an unusual occurrence. However, the disease can take
several months or even years to develop, and animals imported from abroad may
be affected with similar or closely related bacteria. Symptoms are usually
marked weight loss. The disease can be transmitted to humans.
Feline Infectious Anaemia:
This is caused by an organism called Eperythryzoan felis, that is probably
transmitted by blood sucking insects. The smaller Felids are susceptible, and
symptoms are usually preceded by stress.
Parasitic Diseases
Nematodes:
Most non-domestic cats carry Toxascaris common roundworm with a direct life
cycle, whose eggs are very resistant and difficult to eliminate From the
environment. Heavy infestation can cause loss of condition, poor growth rate
and, in severe cases, death due to intestinal obstruction.
Young animals particularly should
have regular parasitic egg counts carried out on faeces and receive treatment
with antihelmintics if necessary. A variety of effective compounds is
available for administering in food. The more recently introduced
preparations, Mebendazole and Fenbendazole, appear to be more effective and
possibly safer, than Piperazine.
Cestodes:
Tapeworm are less frequently found in non-domestic cats. Most tapeworms have
an indirect life cycle, with an intermediate host such as fleas, or other
animals. Meat infected with tapeworm cysts can be a source of infection. Some
species of tapeworm can infect man. The segments of some species can be seen
with the naked eye in the faeces. Relatively safe drugs are now available for
the treatment of tapeworm burdens.
Ectoparasites
Otodectes:
Ear mites can cause severe irritation and discomfort, and may result in rup-
tured ear drums and chronic bacterial infection. Affected animals and those
in contact with them should be treated regularly with an ear preparation to
kill the mites.
Mange Mites:
Sarcoptic and Notoedric mange mites occasionally affect non-domestic cats.
Signs of severe irritation and hair loss are seen. Treatment with acaricidial
shampoos are usually effective, and in-contact animals should also be treated
to prevent further spread.
Protozoal Diseases:
Toxoplasma gondii is a ubiquitous
protozoan parasite that affects many species, including man. Relatively
recently the domestic cat was shown to be a host in which sexual reproduction
of the parasite occurs, without obvious disease. The infective cysts are
passed out in the faeces. The relationship of this parasite with the
non-domestic felines has not been studied in detail, but some species appear
to be susceptible to a generalised infection and become severely ill, showing
signs of fever, depression, loss of appetite and central nervous system
damage including blindness. Care should be taken to prevent domestic cats
coming into contact with bedding, food or water, used for non- domestic
species.
Other Diseases
Injuries:
Probably the most common problemn in captive Felidae. Good management and
housing can reduce the incidence.
Periodontal Diseases:
Common in older animals especially those receiving insufficient hard food. The
accumulation of dental calculus damages and inflames the gum and allows the
tooth root to become infected. This can result in excessive salivation- and
an inability to eat. Removal of affected teeth which have become loose is
necessary.
Ingrowing Toenails:
Another problem which affects older animals, it is often not detected until
the animal is obviously lame.
Gastro-intestinal Upsets:
Symptoms of diarrhoea and vomiting can be caused by the infectious diseases
already mentioned, and by some poisons, but more frequently is a result of
feeding contaminated or decomposed food, or of a sudden change in diet.
Anything which causes malfunctioning of the animal's digestive system may
result in abnormal growth of the bacteria normally present in the small intestines
and this can cause an enteritis and diarrhoea. In some instances toxins can
be produced which make the animal severely ill. Treatment will depend on
veterinary advice, but initially at least food should be witheld. Prevention
of gastro-intestinal disease from these causes demands good quality fresh
food, and avoidance of sudden dietary change.
Nervous System Diseases:
A number of diseases of the nervous system have been described including
convulsions in young tigers (Jones and Ashton, unpublished observations),
star gazing or or labyrinthritis (Theobold, op.cit.), ataxia and
proprioceptive defects associated with skull deformities in lions (J. M.
Hime, 1972 and 1974). Although various theories for some of these conditions
have been put forward, including hypoglycaemia, Vitamin A deficiency, and
nutritional bone disease, the precise aetiologies are not known.
Toxoplasma should be included in the
differential diagnoses,
Poisoning:
Cats are particularly sensitive to many poisonous substances, including the
phenol-containing disinfectants. Any disinfectants used in accommodation for
the Felidae should be thoroughly rinsed away and, if possible, those
containing phenols should be avoided.
Barbiturate poisoning of large cats
has occurred in a number of collections. Meat from animals destroyed with
these drugs has been responsible in some cases. In others malicious feeding
appears to have occurred.
Hairballs:
Accumulations of hair in the stomach particularly in periods of heavy coat-
shedding have been implicated in loss of appetite and weight loss. Treatment
with liquid paraffin in the food or water is usually effective.
Hair Loss:
The cause of patchy hair loss is frequently difficult to determine.
Nutritional deficiencies, persistent licking due to boredom or hormonal
disturbances may be responsible. The fungal infections known as ringworm may
also cause loss or breaking of hair, together with dry scabbiness of the
skin. Ringworm can affect humans.
Conclusion
With the existing knowledge of the nutritional and behavioural requirements
of the Felidae, and the judicious use of vaccines, antihelminthic drugs and
quarantining of new animals, disease in captive felines can be reduced to
very low levels.
References
Ben Shaul, D.M. (1962). The composition of the milk of wild animals. lnt.Zoo
Ybk. 4 333.
Hime, J.M. 1972. Scientific Report
Zoolog.Soc.Lond.,J.Zool.Lond. 166 509.
Hime, J.M. 1974. Scientific Report
Zoolog.Soc.Lond.,J.Zooi.Lond. 173 43.
Povey, R. C. and Dovis, E. V. (1
974). Panleukopenia and respiratory virus infections in wild felids.
Ann.Procs.Symp. Felines, Seattle, Washington, 1974.
Scott, P. P. et al (1967) .
Nutritive requirements for carnivore in Conalty M. L. (ed.) Husbandry of Lab.
Animals. London Acad. Press, 1967.
Scott, W.A. (1979) : Use of vaccines
in exotic species. Vet Rec 104 199
Theobald (1978) in Zoo and Wild
Animal Medicine,Ed. M.E. Fowler. Publ. Saunders, p.650.
Wallach, J.D. (1970) : Nutritional
diseases of exotic animals. J.Am.Vet.Med.Assoc. 157 5.
Wiesner, H. (1 977) : Zur
Narksosepraxis mit dem Blasrohurgewehr Kleintierpraxis 22 327.
R. P. LAWRENCE, Head Warden, West Midlands Safari Park
To the average member of public entering a safari park the immediate and
overriding impression is one of extreme spaciousness and almost unlimited
freedom of movement for the animals. While this impression can be justi- fied
for many of the species kept, where predators are concerned - particularly
wild cats - this apparent freedom of movement represents little more than the
paper thin surface of a carefully nurtured image. Visitors are allowed to see
what they wish to see. Certainly, however, the freedom of movement is a
complete myth.
Well fed lions can spend up to 80 per cent of their time sleeping, and a
pride might occupy only thirty square yards of, say, a five acre reserve. The
rest of the area invariably double-fenced is enormous expense, goes largely
unused. When the lions are active their movements are a source of constant
concern. Thirty lions ranging over five acres can enter any number of
dangerous situations, for it is physically impossible to watch them all, all
of the time. Tigers, more individualistic in behaviour, roam about seeking a
solitude which is totally denied them.
Fig. 1 is a sketch Plan featuring one of our former four cat reserves and
is typical of what one might find in a safari park, in that the briefest
glimpse will confirm the unfortunate and widely held belief that safari
parks, of which West Midland was standard, were thrown together with little
or no forethought or planning. In particular, it shatters the fallacy of
free- ranging captive felines.
For the tiger in this reserve the areas north of the line AA and south of
the line BB were definitely 'no go' areas, Cats in these areas were either
too close to the gates, and the rather exposed gatemen, or could approach
them unobserved by either the patrolling warden, or the gatemen them- selves.
The shaded area to the right represents a depression in the ground where
tigers were not visible from the road. While there is, of course, a strong
argument for giving one's animals a certain privacy, the object of display is
somewhat defeated if the animals can never be seen. The area to the left of
the road had also to be regarded as out of bounds because, in the event of a
security scare - such as tyre biting, open car windows or doors, visitors out
of their vehicle, or a tiger sitting on a car roof causing the windows to pop
out - the patrol vehicle was rarely able to cross the stream of traffic to
react to the situation. All we are left with is a fifty-yard stretch of
ground between the house and the pool. To con- tain the inevitably irritable
and highly strung tigers within this area required two staff vehicles, one at
each end. The tigers usually become so weary of being chased each time they
moved from an easily-viewed position that they would retire to the roof of
the house for the day, further negating any claims of their freedom of
movement
Anything more unsatisfactory, more stressful to the cats and more futile
in terms of manpower is difficult to imagine. Two gatemen, two drivers and
two expensive vehicles would be inextricably committed to providing what at
best is only minimal security for a handful of tigers; four staff unable to
contribute in any way to the welfare of their charges, such was the
inflexibility of the system. A wastefulness repeated in each cat reserve.
For all the public's professed love of seeing animals roaming at will,
there can be little doubt that a close look behind the scenes would have
caused considerable unease. For the poor standards of which safari parks were
so frequently accused proved to be the rule rather than the exception in our
case. Take the pool for instance, that people love to see the tigers
splashing about in. It is seldom appreciated that, in the absence of running
water, this was filled by surface run off from the road, together with
spilled oil, petrol, battery acid, diesel and all the other effluent produced
by the modern motor vehicle, as well as the cats' excreta and urine. This
mess usually constituted the only drinking water.
Feeding was usually carried out by the dangerous, albeit spectacular,
method of driving into the reserve with the meat contained in a metal cage
mounted behind the vehicle; a tedious daily process requiring, in our
situation, up to four vehicles and seven staff for the undertaking; a
ridiculously complex arrangement for a purely routine function.
Cleaning out the sub-standard housing was, of course, impossible with the cats
in residence, so they were turned out. The cleaning process was highly
dangerous because they lurked in the proximity. Returning the cats after the
cleaning-out operation, or at the end of the day, involved a further highly
stressful exercise; a round-up and chase by three or four vehicles. Clearly a
ludicrous and utterly chaotic situation prevailed beneath the apparently
innocuous care-free, spacious ideal visible to the public.
The solution was more easily found than put into practice. The entire complex
of colt reserves was unmanageable and required dismantling.. But what form
would the new image take? The ineptitude in the park's design was not
restricted to the cat reserves; they typified the mismanagement of most of
our species. The further the soul searching progressed, the more apparent it
became that the only solution was a complete and fundamental restructuring of
the entire park. Accordingly, almost three years ago we undertook the
complete rebuilding of the park. The existing eight, and similarly overmanned
reserves were streamlined into just three geographical groupings, i.e.
African, Eurasian and American. In centre of each, surrounded by a suitable
viewing area, new compounds were constructed to contain the indigenous
predator of each respective continent. In Fig. 2 we see an example; the new
lion compound. As is also the case with the tigers and wolves, the animals
can be viewed from a distance of feet but, in complete departure from the
traditional safari concept, through a fence There was initial public
scepticism that, with all due deference to the zoo world, the animals were
all penned-up, as in a traditional zoo. There was undoubtedly a loss of
'thrill value' without the close encounters with predators, that had hitherto
been possible. This drawback from the public standpoint was dwarfed in
significance by the overwhelming improvements now possible in the management
and welfare of the animals. Although, for instance, the area to which the
cats were now introduced was significantly reduced, they enjoy considerably
greater freedom. They have access to every square inch of the reserve without
any form of stressful hindrance.
Indeed, the freedom of the compound is total, for no vehicle ever needs to
enter the area. To house the lions at night, for instance, nothing more
harmful than a run around the perimeter fence and a little colourful language
is required. Any reluctance to enter the house can be countered by
occasionally feeding at night. Otherwise the cats are fed at 8.30am promptly
in the small compound in front of their house before being released into the
reserve at 10.00am, when the park opens. The coast is then clear for the
bones to be picked up and for cleaning out.
Measuring 45ft x 15ft, the new housing can be regarded as adequate rather
than elaborate, but nevertheless, a substantial advance on previous
accommodation, which was rather primitive. It is divided into three equal
sections; two are large pens, whilst the third is sub-sectioned into two
cubbing or handling pens. This proves satisfactory for the mere 12-15 lions
now kept; a stark contrast to the extreme of 80-90 previously kept.
Lionesses in oestrus are segregated to preclude breeding and to keep
numbers down, Electricity, running water, ventilation and provision for heating
is built into the housing; basic essentials maybe, but previously considered
to be luxuries.
Having done so much to reduce the stress factor at every stage of the
daily routine, the sex ratio of the lion pride (currently 6.9 of various
ages) seems immaterial, for little or no in-fighting occurs. The previous
unwieldy group of tigers (8.3) has been reduced to 3.3 with similar results.
From the public standpoint the new informality has become - highly popular.
Not only is the security greatly enhanced by the fence between the car and
the cat, to prevent mishaps, but open windows and sunshine rooves can be
actively encouraged, to increase viewing comfort. After all, sitting in a car
for a solid hour with the windows tightly shut is hardly my idea of a pleasant
summer's day outing.
It may seem ironic, after little more than a decade, that, in order to
achieve the potential that safari parks undoubtedly have, the wheel has
turned almost full circle. One can only feel, albeit reluctantly, that for
cats, other predators and large mammals, such as elephants and rhinos, the
free ranging spacious ideals are not only unworkable, but totally
unrealistic.
PROF. DR. PAUL LEYHAUSEN
Max - Planck -Institut Fur Verhaltensphysiologie
ladies and gentlemen, I apologise very much for my attire, but I had to
come to you in all my dirt because the world's most experienced airline
managed to lose my suitcase. That also means that I lost my notes and my
slides. So, for the time being at least, I hope you will excuse my talking
'off the cuff'.
If I had only to answer the question posed as the title of my lecture, I
could finish very quickly because I could say just one word - 'No'. I am
convinced that captive breeding will not be the answer to species survival
and much less so in carnivores, especially the felids, than perhaps in other
species, such as ungulates. The reasons for this are manifold.
Wildlife conservation does not mean the preservation of any species in its
present state; that could be achieved quite successfully by modern museums in
which it would stay unchanged for ever. In the final analysis, what
conservation is really about is the preservation of the process of evolution.
Evolution is not something which, at any moment, has reached or will ever
reach its final goal; it is an ongoing action and will remain so. However, it
is not possible to create the necessary conditions to preserve the all-
embracing process of evolution within -the limits set by any captive breeding
population. Furthermore, a species is not a monolithic entity, it encompasses
a diversity within a basic uniformity. Variance within a species, and
especially within its genetic make-up, is one of the principal requirements
for its survival; for, variation provides a species with the ability to adapt
to new and changing conditions.
I think the first person to discuss the fact that within any captive
breeding colony you inevitably have only a selected part of the true
gene-pool (the total pool of alleles of each gene of the species), was Dr
Spurway, the wife of the late Professor Haldane. She has shown, convincingly,
that captive populations of the size a zoo can manage cannot have the allele
diversity of a wild population. In principle we face the some kind of problem
when a wild population goes down in numbers to a very low level and then
recovers. A typical example, for instance, is the Northern elephant seal
which at one time, some 15 or 20 years ago, was down to 40 specimens and now has
a population which is so large that, at the CITES Conference at San Jose
(1979), it was transferred from Appendix I (the most endangered species) to
Appendix II (species which need rnonitoring in trade but which, within
limits, can well be utilized). Yet the intrinsic genetic diversity of that
large population is still no greater than it was when there were only 40
animals left.
Thus, within a relatively short period of time, a species can increase
drastically in numbers, but in that short time the genetic diversity and
concomitant evolutionary adaptability are not recreated. In principle this
large population is therefore no better off than, say, the 40 were with
respect to changes in environmental and ecological conditions.
That also applies, of course, to practically all captive breeding
enterprises, with the notable exception of the large landowners, in Texas and
elsewhere, who have successfully bred large herds of antelopes on their
ranches. it will take many generations after the recovery of a population for
mutations eventually to recreate adequate genetic diversity.
With true captive breeding we have another problem. Not only do we start
from a very narrow section of the basal genetic make-up of the species, but
we are also, consciously or unconsciously, selecting. We are, for instance,
selecting breeding groups of harem animals that flourish and raise their
young successfully under captive conditions. Thus, the characteristics which
we select are those useful for survival under these captive conditions. They
may not be at all useful if, for example, we want to reintroduce animals from
our captive colony into their original habitat; and although this is very
important, we can very rarely do it successfully. Some Arabian rulers have
now set aside large tracts of land for reserves and natural parks on the
Arabian peninsula, and we will see how the animals which have been bred in
the United States will manage in these reserves. It is hoped, but it is not
at all certain, that they will breed well.
The problem is slightly different with respect to the animals we are
talking about here. Nobody is able to keep large breeding colonies of some
cat species in the way many antelopes are kept in Texas. To put it in a nut
shell, you may easily have 5,000 deer on, say, 50 sq. kms., but for 5,000
lynxes it would have to be 25,000 sq. kms. Thus, when we establish successful
breeding colonies of cats, we will immediately be selecting for social
compatibility and tolerance of a kind which is quite unlike the norm in wild
cat populations. And here again, on reintroduction of such cats into some
habitat which we assume to be adequate for them to live, even if they settle
down successfully and breed they will never again be the same animals they
were before captive breeding.
I said in the discussion that large predator populations may,, perhaps,
make do with a lot less genetic diversity than, for instance, ungulates. The
degree of homozygosity in predators seems to be greater than it is in
herbivores. Here I should explain a little about how animals manage
themselves How is the natural gene flow within the population regulated?
Among other factors which I will not discuss here, social behaviour, social
organisation and sexual behaviour seem to be of importance. The first in regulating
population density, the second in governing the mobility of the individual
within that population, and the third in determining productivity. The more
densely packed the population is, the higher the probability that different
animals will meet and mate at random . The same applies to motility. The more
mobile the individual is, the greater the probability of its meeting a non-
related strangers, and the less inbreeding will occur in that population.
Thirdly, reproductive rate, and survival rate of the young are factors which
increase or decrease kin mating. Now, with respect to these factors, the
population has to strike a balance. On one side of the scales there is the
necessity of preventing too much inbreeding, because inbreeding will, of
course, increase homozygosity and decrease genetic variation - genetic
diversity within the population and to prevent this, we need some mechanism
in the population, which prevents too many closely related animals from
mating. This is called an incest barrier. Also, in order for natural
selection to be effective, it must be possible to weigh the advantage of one
combination of alleles against another. So on the other side of the scales
there must be a mechanism to ensure that, in some parts of the population,
certain alleles are more frequent than in others, so that you have a
diversity of allele combinations within that population, with differential
valence.
This is usually described nowadays under the heading of kin selection or
kinship, or the not very technical title of 'the selfish gene'. There are two
factors favouring this: shortening intervals between reproductive cycles, and
increasing litter size and survival. Density and motility antagonize each
other to some extent. The higher the density, the less motility is needed for
random mixing, and vice versa. Motility, both within a population and between
populations, determines the degree of diversity in allele combination within
that population and over the whole range of the species. Hence populations of
low density (and carnivore populations are very low in density compared with
herbivore populations) need extremely high motility in order to maintain a
common gene pool throughout the entire habitat. A population of, say, 2,000
tigers needs a minimum area of between 20,000 and 60,000 sq. kms. Tigers
sometimes travel as much as 40 or 60 kms in one night, but they very rarely
keep that up without change of direction, Thus gene flow from one extreme of
such a large area to the other is slow and takes many generations. Therefore,
local sub-populations differ more or less in their genetic make-up. If such a
sub- population is cut off from the rest, then this will lead to inbreeding.
Most extant tiger populations are, by now, cut off from each other by human
settlement and cultivations. Most of these isolated pockets comprise no more
than from one to three or four dozen animals, and continuous inbreeding poses
a serious threat.
How important gene flow and gene exchange are is demonstrated beautifully
by the social organisation of lions. In this species, a pride is formed by a
group of females who recognise each other but do not necessarily stay
together all the time. They occupy a particular area, the boundaries of which
remain relatively stable over long periods of time. This breeding group of
females will not tolerate strange females. The pride members which they lose
from disease, accident or old age are replaced exclusively by females bred in
the group. Surplus females hove to move out to find males. Young males likewise
have to leave because the pride males will not tolerate them within the
boundaries. These surplus females and males form more or less vagrant groups
on the fringe of the settled territories, and it is on these animals that the
population draws when the territorial animals suffer unexpected and severe
losses. So this is a kind of reserve part of the population. They are
breeding, but the survival rate of the young is low, much lower than in the
established prides. The pride is a very sedentary affair and the question is
how do these animals prevent severe inbreeding? They do this by changing
their males. The males usually live in a kind of brotherhood group of 2 - 6
which takes over a pride. Their main task is to defend their territory
against other males and females, and thus secure the quiet and undisturbed
atmosphere necessary for successful breeding and rearing of young. This puts
them under considerable stress and strain, and on average they cannot bear
this for more than two or three years in succession. The breeding life of the
adult female is somewhere around 10-12 years. Thus the average female may
have up to 5 different groups of males to breed with during her lifetime.
This quick turnover of pride males quite evidently prevents inbreeding. As you
know, it is claimed that this is a model case of kin selection because in
some cases when the new males take over they kill some of the cubs sired by
the previous males, and also because, due to the upheaval during the
take-over, the females are disturbed and tend to neglect their cubs whose
survival is therefore limited. Biologists have tried to interpret this in
terms of kin- selection; the new males (or, rather, their genes) supposedly
have an interest in eradicating the genes of their predecessors, and in
replacing them as soon as possible with their own. However, those who make
this kind of assertion seem to forget that the new mates have to leave in
their turn, after 2 to 3 years' time on average. Then what was gained
initially will be cancelled out by the new change. On balance, there is no
potential advantage at all, but the effect is to ensure genetic change within
a large and sedentary population.
To sum up: if we entertain any ideas of trying to save a species by
captive breeding, our first mental exercise must be to devise some means of
providing for sufficient change of animals to ensure a genetic diversity in
different social groups.
I can only say again that, with medium-sized and certainly with large
predators, this will be an extremely costly and difficult operation which may
work for a relatively limited period of time, but certainly not for long. For
that reason, although I do not say it must not happen, I would never advise
it while there still remains a chance to preserve a species in parts of its
natural range. Of course, there are desperate cases were captive breeding
offers the only chance at all of keeping a species alive, if only for a short
time, when there is no chance of releasing it in the wild again. Our first
duty, however, is to support any organisation which fights to preserve the
continued existence of a sufficiently large, natural and varied habitat for
these animals to exist where nature originally designed them.
R. J. P. 0' GRADY, M.A.
Director/secretary, The Zoological Society of Glasgow & West Scotland
I have been interested in genetics in a practical way for many years, and
in fact started reading about the subject whilst still a schoolboy, but I
would be the last to describe myself as an expert.
It was with some surprise that I found on starting zoo work, in 1972, that
considerable confusion existed in the minds of many people, with regard to
colour inheritance in cats, primarily with black leopards but closely
followed by the white tigers at Bristol Zoo.
As one who travelled half the length of Britain as a fifteen year old,
specifically to see the first pair of white tigers at Bristol Zoo, I was
probably more interested in them than in black leopards. However as we were
more likely to be able to acquire black leopards at Calderpark than white
tigers, it seemed logical to concentrate on them. Seven years later, in spite
of numerous hints and downright direct approaches, we still haven't been able
to winkle a white tiger out of Bristol Zoo, so I was probably right.
My attention was drawn to black leopards in 1972 because, at the time, we
had a trio of spotted leopards in our cat house. One pair, every so often in
the past had produced a dead black cub amongst their normal spotted litters.
Soon after I arrived the female of this pair died of an ectopic pregnancy. It
was the attempts to explain why her black cubs were being produced, and that
it was not by 'Act of God' chances or first mutations, that started me off on
the present line of enquiry.
Black leopards in the wild state are well known, as is the production of
black cubs from two spotted parents, or where one parent is black and one
spotted. The occurrence of black and spotted cubs in the same litter had also
been frequently documented. It seemed to me fairly obvious, although at this
stage I had not done any reading on it, that the observations just described
immediately suggested that in the black leopard we were dealing with a
straight forward recessive factor; and so it turned out.
If you cast your minds back to the situation in British zoos in 1972, the
picture was quite different from today's. Breeding results in zoos were only
a fraction as good as now. This can be attributed to changed standards of
awareness, new and better diets, and much higher expectations on the part of
nearly everybody. There was no ABWAK and no Cat Survival Trust, and there
were very few small cats. Good young pairs of black leopards were almost
impossible to obtain without importing them, and then at very high cost.
Dublin and Howletts were breeding them, and Colchester and Jimmy Chipperfield
were producing the odd one or two, and that was the lot. Spotted leopards, on
the other hand, were plentiful and rapidly approaching the situation which,
we find today, of over production.
Then, in a chance conversation with Terry Murphy at Dublin Zoo, he
mentioned to me that he had a litter of 3 male and 1 female black leopards.
As the males were starting to fight he had separated them but was having difficulty
disposing of them as single males. He agreed to give us one, at a listed
value of £700, the favour to be returned at a later date.
We immediately set about acquiring three, unrelated, spotted females, in
the hopes that one at least might be carrying black as a recessive. We knew
that for years Bristol Zoo bad been in the habit of matinq its spotted
female, Jenny, to the male of its pair of blacks. All the cubs from this very
productive female had been spotted and were distributed throughout British
zoos. I was sure our original pair had originated from this pair, and I just
hoped we could pick up one or two other females descended in the same way.
Of course, the arrival in a small zoo of four leopards, more or less
simultaneously, aroused a lot of comment. It was a considerable relief in
1975, when the first cubs were born, to find the entire litter was black. The
first litters from both of the other females were entirely spotted. So, after
another litter or two from each, we disposed of these two females.
During this time I had been asked to give an explanation many times. I am
indebted to Dr Cohen, of the Genetics Department of Glasgow University, for-
drawing my attention to some published work by Ray Robinson, the existence of
which I had not suspected. Until then I had not believed that any systematic
work had been done, I had been working on the assumption that the confusion
existed because of misleading remarks like, 'work at Hanover Zoo has shown
that the black colour is due to a dominant gene, probably homologous with
dominant black in other forms' (Searle, 1968), or like that of Crandall
(1964) that in his experience he had never known of a black cub being born to
spotted parents.
An East German publication by Puschmann (1975) states that, 'Two black
parents usually produce black cubs'. And Henderson, in his book 'Circus
Doctor' published in 1951, states that a black pair may produce spotted
young. Eaton (1977) says that this was the only reference to the phenomenon
he was able to discover in the literature.
Robinson cleared the matter up conclusively. He contacted 128 zoos known
to be breeding leopards and received returns from 72 describing their
breeding experiences with the spotted and black forms. He was able to
demonstrate quite conclusively that the black form of the leopard con be
shown as an autosomal recessive to the spotted. In the interests of brevity,
I have, with assistance, shown this in a diagram (see Appendix).
Robinson also found that the average litter size from black females is
smaller than that from spotted. He has hypothesised on the reasons for this
and discussed the possible significance with regard to the wild distribution
of black leopards in parts of South-East Asia . From my own observations and
those of our Curator, Lutz Kuschinski, we strongly suspect that inbreeding
may have a great deal to do with it.
Many black leopards in Britain are known to be more aggressive and highly
strung than spotteds, many of which are almost ridiculously tame. I am
ashamed to say that our original male exhibits pronounced stereotyped
behaviour patterns such as exaggerated pacing and star gazing, and this seems
to have been inherited to a lesser degree by one of his black daughters that
we have kept. None of our other leopards ever did this, and they were kept in
identical conditions. I feel sure that a lack of understandinq about the
genetic possibilities of introducing fresh blood has been a major factor in
encouraging the persistence of breeding black to black, often in brother and
sister pairs.
I hope the accompanying Appendix will go some way to encouraging people to
branch out in a systematic fashion.
Of course, it may have relevance in other directions. Melanism has been
recorded in many species of exotic cats, for example: jaguar, puma, serval,
lynx, Temminck's golden cat, tiger, Geoffroy's cat and probably others. In
these species it is likely to be inherited in the same way as in leopards.
Whilst a black tiger is not everybody's cup of tea, (1 once heard the
white tigers at Bristol Zoo described as an abomination of nature) there is
no doubt that black jaguars, for example, are. In Britain at the moment there
is only one black jaguar, and that is a female recently imported by Marwell.
The prices quoted on the continent and in the U.S.A. are ridiculous, at
£8,000 or more a pair (although single males cost less than half).
At the same time it is becoming extremely difficult to find good homes for
spotted jaguar cubs in this country,
One could say, well so what? What's the point of breeding melanistic
specimens? Surely we should be trying to encourage the breeding of more
endangered species ? I have considerable sympathy with this point of view.
When a melanistic form occurs in nature, as part of the wild population, then
there seems to be every justification for trying to breed it. Never- theless
when one sees the great scientific and popular interest aroused by the white
tigers at Bristol Zoo, and the excitement amongst cat keepers when referring
to Marwells black jaguar, it's hard not to feel envious.
Anyway, what would the people present here do if they had bred a lion, as
we have done, which, instead of being normally-coloured, was born with a
pitch black patch extending the length of the inside front leg and across the
chest? Melanism in lions is unknown. The only reference I can find is in June
Kay's book 'Okavango' (p.68), which describes a hearsay account of a black
lioness in the African bush.
In conclusion; inbreeding amongst black leopards in captivity is so well
known that a zoo Director said to me recently 'Ah, but my pair was wild
caught'. If the specimens in captivity are to maintain the population into
the foreseeable future without the undesirable necessity for reinforcements
from the wild state, and this paper helps in any way in maintaining a healthy
population, then that's all one can ask.
References
Crandall, L., 1964, The management of Wild Mammals in Captivity. University
of Chicago Press.
Eaton, R. L., 1977, Reproductive Biology of the Leopard. Zoo. Garten N F
Jena 47 (5) S.329-351 .
Henderson, 1951, Circus Doctor.
Kay, J., -, Okavango,
Puschmann, W., 1975, Wildtiere in Menschenhand, p.321, Saugetiere Berlin.
Robinson, R., 1970, Inheritance of the black form of the Leopard (Panthera
pardus)
Robinson, R., 1970, The breeding of black and spotted leopards, J. Bombay
Nat. Hist. Soc. 66 (3); 423-429.
Robinson, R., 1978, Colour Inheritance in small livestock. Fur &
Feather Publication.
Searle, A.G., 1968, Comparative genetics of Coat Colour in Mammals, p.148,
Logos Press.
Ulner, F.A., 1941, Melanism in the Felidae, with special reference to. the
Genus Lynx J. Mammal. 22: 285-288.
This paper by Terry Moore, Director and Vice Chairman of the Cat Survival
Trust, is unfortunately not available, but is here summarised.
The Author -notes man's rapid depletion of wildlife in context of the
first cat- like mammals having appeared 70-80 million years ago. He laments
that zoos have achieved little with the smaller cats. larger cats have been
those kept largely because of popularity with public and ease of location and
capture in the wild. larger size and easier capture also facilitated over-
exploitation of big cots. Smaller cats stood better chance of surviving in the
wild - but not now, because of massive habitat destruction and killing for
skins.
Author believes evolution is now controlled by human activities. Wildlife
adapts to human or becomes extinct. Species becoming endangered could be captive
bred and later introduced to reserve areas or wild, where possible. This is a
problem we created for ourselves by lifestyle, especially materialism. In
particular, author notes urbanisation, spread of agriculture, destruction of
forests, exploitation of finite mineral resources. Author believes biggest
threat to man is his destruction of the environment; disapproves of farming
endangered species for fur coats except as last resort, but remarks on
benefits for reducing pressure on wild populations.
Author notes importance of maintaining strong breeding stock, and argues
for selecting breeders without congenital diseases, and for avoiding
inbreeding. He remarks on the lack of knowledge of minimum gene pools. He
argues for concentrating on breeding endangered cots as a priority - to
produce a surplus for reintroduction to wild or reserves. Problem is little
known about status of small cats, and research needed as a priority. The
author argues against cross-breeding species or sub-species, but for rationalisation
of classification. He also argues for education to environmental
appreciation, through zoos, societies, schools. He notes the need for
studbooks for all cat species and for protective legislation.
The author finally mentions possible environmental consequences of
destroying cat populations (e.g. rodent overpopulation), and notes the
aesthetic value of wildlife - i.e. priceless.
EDITOR
A HALF- CENTURY OF HOME BRED CHEETAHS
V.J.A. MANTON, MRCVS
Curator, Whipsnade Zoo Park, Zooloqical Society of London
The cheetah (Acinonyx iubatus), in man ways a unique non cat-like species
of Felidae, has been kept in captivity for so long that the title of my paper
might lead the reader to suspect that I was about to review the last fifty
years experience of keeping them. Although at times I feel that I have
existed in the zoo world for more than fifty years, I am deliqhted to reveal
that the half-century refers to the number of cheetahs bred at Whipsnade, and
only to that.
As far back as1972, I first attempted to obtain inforrnation from all the
Cheetah breeding units in the world. My aim was to summarise and compare the
conditions in each, in the hope of isolatinq the main factors necessary, as a
prerequisite, before breeding was attempted. Surprisingly enough, knowing all
I know today, some seven years later, nothing has led me away from the
conclusion I reached then and still hold now, that -and I quote from the
report of the Jersey Conference on Breeding Endangered Species in Captivity -
"the most important factor of all is to keep separating and re-
introducing sexually mature animals which should otherwise be kept in
isolation from one another." I think it is important to remember that
that was written in 1972. In 1974 Dr Randall Eaton published his tome on the
cheetah, 'The Biology, Ecology and Behaviour of a non domestic species'. He
has organised three international symposia on the world's cats, and was very
surprised to learn at the Seattle Symposium in that some year, that
"Whipsnade has bred cheetah from parents themselves captive bred".
Prior to that no one across the other side of the Atlantic had thought to ask
the Society how far we had got, although at the preceding symposium in 1973,
at Oregon, it was reported second-hand that "more recent births at Whipsnade
in 1971 and 1972 are believed to have involved captive born young of the
first births". This, as you know, is untrue in any case, but it does
confirm what previous speakers have said on the importance of keeping records
and passing on information. You should have seen the faces of our American
colleagues at that symposium when I was able to list the litters that had
been born at Whipsnade. Figure 1 shows the situation now. This is reproduced
by kind permission of the editors of 'Wildlife' where it first appeared in
the December 1978 issue. For those of you who are not toxophilists, I should
explain that the "Whipsnade Strikes Gold" refers to the bull target
in archery for which you score 50. This total is made up of 20 males, 27
females and 5 unsexed animals in 16 litters.
Not only have we bred from the original pair "Juanita" and
"Jack" but we have now bred from a total of 5 males and 4 females,
During this symposium we have heard a lot about the introduction of new stock
into captivity to give genetic variability, and the ability of captive born
stock to acclimatise readily. I must admit straightaway that three of the
four breeding females were themselves born at Whipsnade. Only one Whipsnade
born male has been involved directly in the breeding, the others arrived
either straight from the wild or via another zoo. You have already heard of
Mr. Knowles' chagrin when he admitted that one of his animals, imported from
South-west Africa (Namibia) in 1970 or 1972, was sent from Marwell to
Whipsnade where, possibly due to the stimulus of a new habitat and a fresh
female, it very soon achieved a mating, followed by a litter. The fifth
breeding male is interesting because he had been away from the collection for
some time and returned from Chester to mate with a female fairly soon after
introduction.
Mr. Ashton earlier mentioned housing and so did Mr. Knowles. The original
breeding house at Whipsnade, described in Volume 10 of the International Zoo
Yearbook, is a "dog kennel" with double-wood walls in which a number
of species has been kept, including, if I am reliably informed, Giant Panda
in the war, and in which the first three litters to Juanita were born. We
made it absolutely private. There is a keeper door at the back (i.e. there is
no access to one side) and on the front we added a "porch" so that
one cannot see round into the cubbing den. This is further protected by a
wall built across the house, so that even from the rear door it is not
possible to see into the private breeding quarters. We also put in a false
ceiling so that the animals self-generated heat should not be dissipated
through the roof. Indeed, when the first litter was born we had to drill a
hole through the roof to make sure that Juanita was still alive and healthy.
And indeed she was, as was shown by the arrival of her September 1967 litter.
We then built a brand new cheetah house (described further in Volume 14 of
the International Zoo Yearbook). There are basically four pens circling the
house, and inside are four dens along the front and two dens at the back with
sliding doors in between. It is a brick house covered with timber to make it
fit in with the woods in the background. Litters have been born in here
within both sound and siqht of other animals. We have also bred cheetahs in
the converted chicken house at the rear. Here we removed the egg-laying box
from the side but apart from that it is a standard chicken house as can be
bought on the agricultural market today. It is very successful for chickens
and we adapted it only very slightly for breeding cheetahs. This involved
putting a barrier of straw bales three-quarters of the way across the front
of the house so the animal had to go around the wall into a private den. The
side door was blocked with bales so that in the back was a little nest area.
(This is portrayed on p. 128 of International Zoo Yearbook Volume 14). In
here built up a terrific "fug" during the winter, which prevented
anyone who was mucking out from taking deep breaths unless, of course, they
were impervious to high levels of ammonia. Nevertheless, this has been
successful for the natural rearing of cheetah.
Originally I looked into as many factors involved in breeding as I could
and during my investigations it was interesting to note the relationships
between the dates of introductions or re-introductions of sexually mature
animals and the dates of subsequent birth, and therefore to calculate the
time after introduction that mating took place, Figure 2 shows the relevant
dates. We know some of the parturition dates accurately and we know some of
the mating dates accurately so we could say the pregnancy was so long. Other
pregnancies are "guesstimates" and combined with a possible
breakdown of the period between introduction and birth as is shown in the
last eleven litters. We are now able to select when sexually mature males are
introduced to our females coming into oestrous. We have heard from Mr Ashton
about stress and disease, and we have heard from others about the question of
interference and I would admit that we, of course, have not had 100% success
in rearing. Janica had a youngster in May 1974. We were not particularly
concerned about her nineteen days later but perhaps some three days after
that, when we saw the youngster, we should have appreciated that there was
not much in the way of meat on its hindquarters. Although the dam had reared
successfully before, we should have realised that she was being disturbed by
the movement of serval in the next cage. For, although she was a full sister
to Jiffie, who has successfully reared five - the first F2 captive generation
- Janica was unable to withstand the pressures of other animals close to her
pen.
So, in summary Mr. Chairman, you might ask "Where are they now?"
because, after all, it is no good breeding if you are not able to disperse
these animals around the world. Of the first litter, these animals are all
dead now. Jason, in fact, was humanely killed only in July 1978, having
suffered some deformity of his pelvis probably for many years; perhaps this
was caused soon after birth, during the period of stress when he and his
sisters were moved from their breeding den during an electrical breakdown.
The female Jasmine was also the subject of euthanasia, because we wanted to
look at her central nervous system as there appeared to be some difficulties
with her locomotion, and Jacaranda died in an epileptic type fit. The two
parents are also now dead. Jack, who had been easy to handle and was even led
on a chain outside his paddock, died as one of the cowpox victims. And
Juanita, who had always retained her wild behaviour and could never be
handled, suffered an anaesthetic death at Regent's Park Zoo hospital, here,
in 1977. Frank is at Jersey; I hope, still. Grace, died at Regent's Park
where she was in the collection up until 1969. Janica is still with us, as is
Jiffie although Jo-jo died in Regent's Park in 1972. Jonquil is at the
hospital in Regent's Park, at the present time undergoing observation on her
sexual physiology, in other words, observations via laproscope to see if
ovulation can be induced by hormonal therapy rather than just by rnatinq.
Jaino is at Montpelier in exchange for Raoul, and Juba is in Paris. Adorn,
the first of our youngsters born to a captive bred female and who I do not
regard as second generation, because his grandfather in fact sired him, died
in 1978 with kidney failure. The May 1974 male died at 28 days of age. Pueblo
is still at Whipsnade, as is the male from Chester, and Eve. Of the females
in the 1973 litter, one is at Melbourne, the other at Chester. Of the
September 1974 litter, one male is at Dublin, a male and a female at Glasgow,
the second female died of injuries three days after moving, and we never
found the carcase of the fifth animal, which is why it was not sexed, but we
did know there were five animals in the litter. Of the May 1975 litter, two
males are in Gelsenkirchen, a male and female are in Edinburgh, and we did
not recover the carcase of the fifth animal which died about eleven days
later so that we were unable to determine its sex. Of the March 1976 animals,
Alexander is now in London, and the female is in Dubai. The other three
animals disappeared and it is interesting that, on checking, they died or
were killed about the some time as we were moving lynx from behind the cheetah
enclosure to Regent's Park to help fill the new Cat Terraces there. The dam
was the same animal Jonica, who in fact was disturbed when she had her little
male cub in May 1974, next to the Servals. Alexandra is at London and the
male and female of the June 1976 litter are at Amsterdam, the other male
having also gone to Dubai. Of the April 1977 litter, the male is in Taronga
with one female, one female is with Count Herberstein in Austria and the
other female is a mate for Frank in Jersey. Rocky and Rima are still with us,
Rhona is at Marwell as an exchange for the loan of their male which sired the
September 1977 litter, Unfortunately, this male did not survive long. After
mating, he died in September 1977 of a tuberculosis infection. Of the 16th January
1978 litter, the male went to Chester in September 1978, replacing the male
that was returned to us. The rest of this litter and the whole of the August
1978 litter are still with us; and the only animal I have not mentioned is
the little female who died on the 18th January this year (1979).
She was sired by the "Chester male" and was the first of the
animals we have seen that has been deformed. This deformity included the
presence of only one kidney in the carcase and a complete absence of a soft
palate on the dorsal aspect of the mouth. Remember, the "Chester
male", as I call him, was born in October 1973. His mate on this
occasion was his mother Jiffie. We have also had a sibling mating with Pueblo
and Eve, with normal healthy offspring.
So to summarise quickly, of the 53 animals born at Whipsnade, 39 are, to
the best of our knowledge, still alive today from 16 litters born at
Whipsnade. These have been bred from only three animals directly imported
from the wild. I think, Mr. Chairrnan, this is some indication of the
progress we have made at Whipsnade since the first step in 1967, which we
hope will be of some help to other collections in maintaining and breeding
this species.
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