B R E E D I N G 215

Breeding the Brown lemur

a t the Quebec Zoo Lemur macaco fulvus

JACQUES PRESCOTT Biologist, Jardin Zoologique de Quebec, 81 91 avenue du Zoo, Charlesbourg, Quebec, Canada GIG 4G4

The Brown lemur, native to Madagascar, is arboreal and feeds mostly on foliage, fruits and seeds. In its natural habitat it lives in small groups of 4-17 individuals (Sussman, 1977), each group defending a well defined territory and the animals communicating in the main by vocalisation and olfaction; visual communication is also well developed (Jolly, 1966). In June 1973 the Quebec Zoo acquired 1.2 adult Brown lemurs which have since successfully reared ten (3.7) young; a second generation birth (unfortunately not reared) has been recorded. This paper outlines in brief the special attention required to breed this species and discusses the possible causes of infant mortality (see Petter (1975) for a general review of breeding conditions for lemuroids).

HOUSING The lemurs are kept all year round inside the Carnivore and Primate House. They are housed in glass-fronted, two-section cages, each section being 2.4 m high with a floor area of 1.68 x 1.42 m; the compartments are separated by a sliding door, 37 x 37 cm. They are maintained at a constant temperature of 27°C.

In the wild Brown lemurs have been estimated to rest for 61% of daylight hours (Sussman, 1977), and considerable noctur- nal activity is reported (Harrington, 1975). At the zoo they are kept under a reversed light cycle: each cage section is illuminated from 0800-1800 hours by two 40 W red fluor- escent tubes and for the remainder of the night by two 40 W white fluorescent Vita-lite tubes. The latter reproduce 91% of the natural light spectrum. Preliminary obser- vations indicate that the activity level of the animals averages 58% under the red light and is much lower under the white.

The lemurs are fed twice a day with monkey chow biscuits, lettuce, carrots, celery, oranges, apples and currants. Cages are provided with running water and sloping branches and are washed and disinfected daily before feeding.

BREEDING Between 1973 and 1980 the original breeding group of one d and two op successfully reared 3.7 young out of 20 births from 17 different litters. Twins were four times recorded (Table l), one occasion (31 January 1979) constituting a full second generation birth. Here the three-year-old parents, born at the zoo on 26 and 27 October 1975, had been separated from the founder group and kept together, in similar accommodation, since 1976; the litter was unfortunately lost a day after birth.

Eight young survived less than six days and seven of them, weighed shortly after death (Table 2), were considerably lighter than the 78 g birth weight reported by Napier & Napier (1 967) for the species. In every case indicated by an in Table 1, the d parent succeeded in taking the neonate away from its mother and prevented nursing. Once, the hind legs and abdomen of a dead infant were missing, presumably eaten by one of the adults shortly after death. Another time the skull of a two-day-old young was broken and the brain apparently eaten, as the rest of the corpse seemed intact. On three subsequent occasions the neonates had been bitten on the head and muzzle, probably by the d.

The annual distribution of births is presented in Table 3. Of the first generation births, 35% (n=17) took place between March-July and 53% between September- December, suggesting a biannual breeding pattern.

216 BREEDING

INTERVAL BETWEEN BIRTHS LITTER

SIZE LONGEVITY MOTHER (daysf DATE OF BIRTH

First generation

19 Ju173 alive

1.0 0.1 21 alive

22 days 12 Sep 74 3 Oct 74

1 (pair 3) 0.1 26 Oct 75 27 Oct 75

alive alive

91 9 2

7 JuI 76 8 Nov 76

1 .o 1.1 124 stillborn

6 2 days+ g 6 days+

unknown

91

19

5

29 Mar 77 17 Apr 77

0.1 1 .o

alive 5 days+

QI

92

19 Oct 77 24 Oct 77

1 .o 1 .o

1 .o

1 .o 1 .o 4

2 days+ I day+

Qi

QI

6 Jun 78 stillborn 92

27 Jan 79 31 Jan 79

1 day 60 days

Q2

QI

30 Nov 79 1 Dec 79

Second generation 31 Jan 79

0.2 1.0 1 alive

alive Qz Q1

Q3 2.0 1 day+

+adult d directly responsible for death of young

Table 1. Breeding history of the Brown lemurs Lemur macaco fulvus at Quebec Zoo. The extent to which births were synchronised between the two s i n the founder group is shown in column 2.

In March 1977 Q~ gave birth 141 days after her last parturition and exactly 135 days after the death of her previous offspring. If the gestation period of the species is between 120-135 days, as reported by Napier & Napier (1967), this would indicate that she came into oestrus immediately after the death of her six-day-old infant. Further evidence of

a post-partum oestrus was obtained in February 1979, when the young zoo-born 0 was seen copulating only six days after the loss of her day-old twins.

Later in 1977 p1 again gave birth, only 209 days after her last, which means that this prolific Q probably came into oestrus while she was nursing a three-month-old offspring.

BREEDING 217

secluding themselves when they need to. Even if they were provided with boxes isolated from the main cage, the continued presence of an adult 8, as discussed in the following paragraphs, could still present a problem. Although it has not so far been practicable to enlarge our lemur accommo- dation, we have managed to separate the young as soon as they are fully independent of their mothers. The two present breeding groups consist, respectively, of the three original adults and the young zoo-born pair.

Consideration should also be given to the presence of the d and to the synchronisation of breeding amongst op in the group. In each of our colonies, the adult d is clearly dominant over the op. After each birth he stays in close proximity to the new mother and tries to take the young away from her. If he succeeds, he often continues to carry the infant about and prevents the Q from touching or from nursing it; after one or two days it eventually dies of inanition. Some- times the 8 may bite the neonate to death.

As shown in Table 1, fully synchronised births obtained in 1973 and 1975 led to the successful rearing of all offspring. In these cases, we observed that the mothers would often nurse alien young and even exchange infants during nursing bouts, thus generally improving milk intake. They would also combine forces in defending the young against the d. Poorly synchronised births, on the other hand, each time resulted in the death of at least one of the neonates. Interestingly, p1 succeeded in 1977 and 1979 in raising her infant, despite the fact that the births were not synchronised with those of Q? On these occasions she was highly aggressive both towards the other 0 and towards the d and thereby succeeded in keeping them at a safe distance from the offspring. When p2 gave birth three weeks after 9, in 1977 and four days before her in 1979, her lack of aggression was in marked contrast - so much so that neither of her young survived longer than five days. A high level of aggression was clearly of breeding advantage to Q ~ .

In the younger pair, it was also absence of aggression in the 0 which led to the death of her firstborn, as she could not prevent the d

I)ATF,OF AGE WEIGHT

BIRTH SEX (days) ( !4

8 Nov 76* d 2 473 9 6 57.4

17Apr77 d 5 5 3.4 19Oct77 d 2 57.9 27Jan79 d 1 48.5 31Jan79* d 1 663 (62.9t)

d 1 58.3 mean 55.6

*twins tweighed shortly after birth

Table 2. Weight at death of infant lemurs; day 0 = birthday.

~

NO. OF LITTERS

MONTH ISTGEN. 2NDGEN.

January 2 1

April 1 June 1 July 3 September 1

March 1

October 5 November 2 December 1 - -

Total 17 1

Table 3. Annual distribution of births within the two breeding groups.

Even though nursing was still observed at that age, the young was in fact quite independent of its mother.

DISCUSSION There are several factors that might account for the rather small proportion of young lemurs that have so far been reared. The first is cage size. According to Petter (1975), it is possible to breed lemurs in relatively small cages. In the accommodation available at Quebec Zoo, however, parturient PQ are in constant close contact with other members of the group and have no opportunity of

21 8 B R E E D I N G

from pursuing her and trying to touch the twin infants.

To resolve this problem, we once tried isolating a mother and young shortly after parturition. However, the p soon became apathetic and refused to feed until she was reintroduced to the group. On another occasion we isolated the d, with no better results. Our latest manipulation was performed shortly after the birth series at the end of 1979 (Table 1) where we removed the

and a juvenile 9, segregating them from the two mothers by a wiremesh partition. This move has led to the so far successful rearing of all three offspring. Even so, after eight months of separation, we are still apprehen- sive about reintroducing the d to the family group.

According to Jolly (1966), breeding in lemuroids is governed by pronounced social or environmental synchronising mechanisms. The synchronisation of oestrus and partur- ition in pp of the same group may be caused by the social and olfactory stimulation gained from mutual grooming and anogenital scent- marking on objects and conspecifics. The hypothesis is favoured by the fact that the frequency of these behaviours increases noticeably during the breeding season (Harrington, 1974, 1975). Synchronisation does occur in captivity and has obvious bearing on the survival of young; whenever it is out of phase, the parturient pp will have great difficulty in defending their offspring against an adult d.

As synchronisation of breeding is appar- ently related to scent-marking, it is likely that daily disinfection of the floor and walls of the

lemur cages reduces the synchronising effect of the marking odours. Reduction in the frequency of disinfection during the breeding season should thus improve the synchronis- ation of oestrus and, as a result, breeding success within the colony. Owing to the demands of hygiene in the present instal- lation, it has not yet been possible to put this hypothesis to the test.

ACKNOWLLOGEMENTS The author is gratefully indebted to Mr Robert Morin, primate keeper, for his assistance in gathering data and his helpful observations. Thanks are also due to Dr Robert Patenaude, who kindly reviewed the paper, and to Miss Jeannine GagnC, who typed the manuscript.

PRODUCT MENTIONE[> IN T H E T E X T Vita-lire: fluorescent tube manufactured by Durn-Test Electric I.td, Montreal, Quehec, Canada.

REFERENCES HARRINGTON. J. (1974): Olfactory communication in Lemur fulvus. In Prosimian biology: 331-346. Martin, R. D., Doyle, G. A. & Walker, A. C. (Ids.) . landon: Duckworth. HARUNGTON, J. (1975): Field observations of social behavior of Lemur fulvus fuluus E. Geoffroy 1812. In Lemur biology: 259-279. Tattersall, 1. & Sussman, R. W. (Eds.). London: Plenum Press. JOLLY, A. (1966): Lemur behavior. Chicago: University of Chicago Press. N A P I ~ R , J . R & NAPIER, P. H. (1967): A hondbook of living primates. London & New York Academic Press. PF.TTER. J. J. (1975): Breeding of Malagasy lemurs in captivity. In Breeding rndangered species rn rapttviry: 187-202. Martin, R. D. (Ed.). London & New York: Academic Press. SUSSMAN. R w (1977): Feeding behaviour of Lemur catto and Lcmur fulvus. In Primate ecology: 1-36. Clutton- Brock, T. H. (Ed.). London & New York Academic Press.

Manuscript submitted 23 April 1979

Breeding the Malayan giant squirrel

at London Zoo Ratufa bicolor

R . B. WILLIS Overseer of Mammals, The Zoological Society of London, Regent’s Park, London, NWI 4R Y, Great Britain

There are four species of the genus Ratufa, best known in zoos is R. indica, the Indian the giant squirrels of South-east Asia. The giant squirrel or Malabar squirrel, which has