The Behaviour in Captivity of the African Civet, Civettictis civetta (Schreber)

1. What they did

The African civet is the largest African viverrid (the family that includes genets and civets). They are solitary, nocturnal and shy, so very little was known about their behaviour in the wild. The authors watched two pairs in captivity over several years and described their whole behaviour repertoire.

• The animals. One pair, obtained as juveniles from Jersey Zoo, lived at the National Zoological Park in Washington, D.C. The other pair was captured from the wild in Ghana at about 12 days old and raised by hand at the University of Ghana, first in a laboratory and later in outdoor enclosures with grass and shrubs. The Ghana female then had five litters, which the authors also followed.
• What they recorded. Colour patterns, senses, gaits and postures, calls, grooming, toilet habits, scent marking, prey capture and eating, mating and reproduction, the development of the kittens, and how the parents behaved towards them.
• Tests. They offered live and dead prey of many kinds (insects, mice, rats, a hen, frogs, fish, snakes), including mice and rats with the skin removed, to see how the civets attacked and ate them. They recorded calls for spectrographic analysis and filmed flehmen.

2. What they found

Colour, senses and body language

• The irregular spotted coat is very good camouflage: a civet in a clump of grass can be passed within a metre or two without being seen.
• The white neck stripe may act as a “bite director”. Fights, even playful ones, are stylized “neck fencing”, with bites aimed at the side of the neck where they do little harm.
• Smell is extremely acute. Food that is not moving is found by smell, not sight, even when it is in plain view. Hearing is also sharp; sight seems less important for finding prey.
• Civets have few facial expressions; mood shows in body carriage. A frightened civet lowers the body and flattens the crest. In defensive threat they raise the long hair of the crest (up to 105 mm) and turn side-on, so they look much bigger.
• An appeasement posture (quickly lying down on the side) stopped attacks between the familiar male and female. Of 16 cases, only 4 showed the belly.

Calls

• Three threat calls (growl, cough-spit, scream), a kitten distress mew, a miau during mating, a possible female sex call, and a contact call that sounds like “ha-ha-ha” said with the lips closed.
• Young kittens always answered a contact call and moved towards the caller. The mother answered her kittens’ calls but waited for them to come to her, and used the call to summon them.

Toilet habits and scent marking

• Each animal used its own fixed spot every night to leave droppings, so a dung pile would build up. They liked to defaecate from a height: the male climbed part way up a sloping log, the female went to the top. Kittens chose their own sites close to their favourite sleeping place.
• Females and young urinated straight down from a slight squat. The adult male sprayed urine backwards between the hind legs onto shrubs, grass tufts and other upright objects; the Washington male often kicked backwards as he walked. On smelling fresh female urine the male showed flehmen (lifting the head with lips parted). The Washington male did so 32 times after sniffing the female’s urine.
• Civets mark with a large perfume gland between the anus and the genitals, squatting on flat surfaces or backing up with the tail raised against upright ones. Because of this, the height of a mark reflects the size of the animal: the Ghana male’s marks were about 5 cm higher than the female’s. Males marked far more often, and mostly on upright objects; females mostly marked flat surfaces. Young animals only squatted.
• A new object, or one carrying a stranger’s scent, was almost always marked by the next morning, but only after the civets had become used to it.

Hunting and eating

• The paws are not used to catch prey. Attacks build up from a quick “run-away bite”, to “bite-and-throw” (often with a leap in the air), to “bite-and-shake” so violent that it broke a 42 cm snake’s backbone in 6 places, and finally a killing bite. Once the prey was disabled, a carefully aimed bite to the head often finished it.
• Unfamiliar prey, large prey and prey that fought back were treated with great caution. A young female took 5 minutes to make her first bite on a crouching rat. Experienced adults killed rats without hesitation.
• Snakes were first bitten near the tail, and the civet leapt aside after each throw to avoid a strike. File snakes, which smell unpleasant, were attacked hesitantly and hardly eaten.
• Lacking sharp cutting teeth, civets hold food down with the forepaws and tear pieces off with the incisors. Mammals, snakes and large lizards were eaten from the head down, even when skinned. Frogs were opened from the belly and the insides eaten; toads were usually refused. With experience, civets learned to strip meat off the large wing and tail feathers rather than swallowing them.
• Small fish, frogs and water insects were caught in the water dish with a quick scooping bite.
• Before or while eating, civets often rubbed their chin, neck and shoulder on strong-smelling or slightly rotten animal food, and especially on the guts of fresh kills. Fruit was never rubbed. The male also rubbed on mosquito repellent and people’s arms and legs. The purpose of this rubbing remains unclear.
• Grass eaten in quantity passed through almost unchanged, with the animals’ own hair concentrated in it, which suggests it may stop hairballs forming.

Mating and breeding

• The female’s heats lasted from one to six nights. She signalled readiness by running past the male to make him chase her, then crouched in a cat-like mating posture. In the two matings seen in full, the male stayed mounted for about 40 seconds after the likely moment of intromission, and 70 seconds in all.
• The female was mature at about one year and had her first litter at about 14 months. Litters were of 2 or 3 kittens, and she could have two litters a year. Births in Ghana (captive and wild) were recorded in at least eight different months, so breeding is not strictly seasonal.
• Gestation was about 65 days for two litters but 79, 80 and 81 days for the three litters conceived after a miscarriage or the loss of a litter.

Kittens

• Kittens are born furred, with eyes open or opening within a few days. They began exploring outside the nest at 17 or 18 days, ate earth and first took raw meat at about 3 weeks, and froze or crept into cover when alarmed from 9 to 18 days old.
• Young kittens alarmed by a sudden noise sometimes climbed a fence post or the cage wire and froze there.
• First kills of live prey followed the same cautious sequence as adults; experience simply brought confidence.
• In litter 1, each kitten kept to its own teat (the male used the left teat 25 of 27 times; the female the right teat all 27 times), but this “teat ownership” was established later than in domestic cats.
• Kittens licked their mother’s mouth for a short time at 35 to 40 days old, just before she first shared food with them.
• The mother did not bring whole prey. She carried food towards the kittens, called them, left scraps, and later stopped eating once they arrived. No regurgitation was found. Kittens were weaned at 14 to 20 weeks.
• In captivity the father became tolerant: kittens rested against him and he licked them, behaviour the authors think unlikely in the wild.

3. What they concluded

• The general coat pattern is camouflage, while the white markings (neck stripe, ear spots, muzzle) may have social functions.
• Smell and hearing matter most for finding prey. Civets are cautious killers that disable prey by biting and shaking and only then risk a bite to the head.
• Because the mother’s help with solid food is so simple, kittens depend on a long period of nursing while they learn to catch prey themselves. Teat ownership may matter most late in nursing, when kittens have teeth and could hurt each other.
• Licking at the mother’s mouth may be a signal that the kittens now need solid food.
• The longer gestations after a lost litter suggest a possible facultative delayed implantation, but the authors stress this is not direct evidence and needs further study.
• Several civet behaviours (the cat-like mating posture, climbing to escape, the aimed head bite as a final blow, simple food sharing) may help explain how the more specialized behaviour of cats evolved, and suggest civets became ground-dwelling relatively recently.

6. Other technologies, skills or innovations

• Sound spectrograms of the kitten distress mew and contact call, giving durations and frequencies of each note.
• Measurements of nose height during flehmen taken from cine film.
• Simple tests with skinned and head-skinned mice and rats to see whether skin and fur guide the direction of eating.
• Searching droppings for mouse hair to check whether the mother regurgitated food for her kittens.