NEW DEVELOPMENTS IN THE ZOO WORLD 229

sent out a second questionnaire in an attempt to compile a report on the experi- ences of facilities that hold storks and we hope to make this information available to interested institutions.

ACKNOWLEDGEMENTS

We are grateful to Guy Farnell, Charles Luthin and other participants at the workshop who contributed substantially to the design of the questionnaire. The preparation and mailing of the questionnaire was supported by the Brehm Fund for International Bird Conservation and the Savannah River Ecology Laboratory. Labels for mailing of questionnaires sent to institutions on the ISIS address list were provided by ISIS. We wish to thank the many people and their institutions for responding to the questionnaire. We also wish to thank the many individuals and institutions, including Professor Wang Qishan, Kyoko Archibald and the Parc Zoologique de Saint Martin la Plaine, that provided us with additional information. Analysis of the

results was supported in part by the National Environmental Research Park Program of the Savannah River Ecology Laboratory. This programme was partly supported by the United States Department of Energy, Savannah River Operations, contract DE-AC0976SR00-819 with the University of Georgia, Institute of Ecology, Savannah River Ecology Laboratory.

REFERENCES COLLAR, N. J. & ANDREW, P. (1988): Birds to watch. (Tech. Publ. int. Coun. Bird Presv.) No. 8: 1-303. JOHNSON. R. E., COULTER, M. C., LUTHIN. C. S., KING, C. E. 6r VALENZUELA, A. J. (1987): Storks: status, conservation and captive breeding. Colon. Waterbirds 10: 23624 I . LUTHIN, C. S. (1987): Status of and conservation priorities for the world’s stork species. Colon. Waterbirds 1 0 181-202.

Manuscript submitted 28 March 1988

Int . 200 Yb. (1989) 2 8 229-234 OThe Zoological Society of London

Reproduction of the Lappet-faced vulture

at Tel Aviv University Research Zoo Torgos tracheliotus negevensis

H. MENDELSSOHN & U. MARDER Department of Zoology, Tel Aviv University, Ramat Aviv 69978, POB 39040, Tel Aviv. Israel

The Lappet-faced vulture Torgos trache- liotus is well known in South and East Africa where the subspecies T. t. tracheliotus occurs. The North African subspecies T. t . nubicus is less common and appears to have become endangered in recent years (Cramp & Simmons, 1980; Bruun, 1981). The third subspecies T. t. negevensis, identified by Bruun et al. (1981), was first observed in Palestine in 1938 (pers. obs.) and seen on several later occasions. What appears to be the same subspecies occurs in different localities of the Arabian peninsula. The

status of this population which is threa- tened has been summarised by Jennings & Fryer (1984) and the decrease of the population in Israel has been described (Bruun, 1981; Bruun et al., 1981; Leshem, 1983; Mendelssohn & Leshem, 1983a). By 1975 it had declined from an estimated 25 pairs in 1945 to about seven pairs and the establishment of a captive breeding group was discussed at the Nature Reserves Authority (NRA). There were the usual conflicts of opinion but eventually G. Ilani, a zoologist at the NRA, took the initiative and collected three nestlings of

230 NEW DEVELOPMENTS IN THE ZOO WORLD

approximately six weeks of age. They were transferred to the Canadian Centre for Ecological Zoology (CCEZ). the research zoo of Tel Aviv University, and housed in separate rearing cages which were visually divided from each other in order to prevent possible sibling imprinting. The cages were located in an aviar-y with a pair of adult Lappet-faced vultures so that the young birds were exposed to their own species and could develop normal species-specific behaviour (Mendelssohn & Marder, 1983).

For several years no more birds were collected but in 1981. when the wild popu- lation had decreased to three pairs (Leshem. 1983). the efforts to establish a captive breeding group were renewed and by 1985 five more birds had been acquired either as eggs which were artificially incu- bated or as nestlings which had been removed from the wild and reared at the CCEZ using techniques as described by Mendelssohn & Marder (1 983).

By 1985--1986 the captive group comprised seven birds; two hatched in 1975. one in 1981. three in 1982 and one in 1985, this last being too young at time of writing to show any breeding beha- viour. In addition there were two old birds. a crippled 3 and a human- imprinted :. The <, found in 1978 with an injured wing. had been paired in the wild and had reared several young. I t courted several birds in the captive group but because of its injury failed to mate successfully. The hatched in 1956 had become sexually imprinted on one of her keepers. She laid her first egg when eight years old and often produced replacement clutches. For several years she was separ- ated from her original keeper and paired with a ;. Although the two birds partici- pated in nest building and copulated often. no eggs were laid. When the j' died and the Y was returned to the charge of the tame keeper, however. she again began to lay eggs. This imprinted <, removed from her parents' nest at the age of about one month. was therefore stimu- lated to nest build and to copulate by the

presence of a 6 vulture but apparently the stimulus for oviposition was her human 'pair mate'.

This Q laid her first egg at the age of eight years and another imprinted 9 laid her first egg at the age of ten years, so it would appear that T. t . negevensis reaches sexual maturity at a fairly late age. The captive-bred 9 mentioned below, however, has bred successfully at six years of age. Preliminary courtship may begin at the age of two or three years and pair formation of adult T. t . negcvenszs has been described (Mendelssohn & Leshem, 1983b).

The seven young birds, together with the two older ones, were housed in an aviary provided with perches, a 2 x 2 m nesting platform placed 2 m above the ground and an open shelter extending the whole length of the aviary and affording protection from the rain. In their natural habitat in Israel annual precipitation is c.50 mm. In Tel Aviv it is 500 mm and far less solar radiation is experienced in winter. The birds, however, make little use of the shelter.

In November 1986 a 6 hatched in 1981 and a 2 hatched in 1982 formed a pair bond and in December they began to carry branches to the nesting platform. An unpaired 0 hatched in 1975 tried to take possession of the platform and she and the 3 of the pair often fought over the right to build on it. The young 9 of the pair was less often involved in the aggres- sion. On the morning of 30December 1986 the 5 was found dead and partly dismembered, the throat lacerated, trachea and oesophagus torn, and breast muscles of the right side lacerated and partially eaten. The aggressive 5' was removed to a separate aviary where, when she was 12 years old, she built a flimsy nest on the ground, ignoring the elevated nesting platform and laid her first egg and a replacement egg a month later.

Within two days the young C formed a new pair bond with a 3 hatched in 1982. The pair showed a large amount of display behaviour towards each other,

NEW DEVELOPMENTS IN THE ZOO WORLD 23 1

with the head-stretch-and-turn display (Mendelssohn & Leshem, 1983b), bill- touching and some allopreening, and they built a nest on the platform.

Copulations began in midJanuary 1987. At first the inexperienced 6 attempted to mate with perches near to the 0 but after a few days coition with the $2 was achieved. At the end of January the birds began to take turns to incubate in an empty nest, the five-year-old 0 not having produced an egg. After about a week both birds were found incubating stainless steel food dishes, turned upside-down on the ground, each bird incubating and defending its own dish. When the keepers turned the dishes the right way up the birds deliberately up-ended them within a few minutes and continued to incubate. When a dish was placed in their nest the pair took turns to incubate it for about three weeks after which the incubation and other aspects of reproductive beha- viour ceased although the pair bond was maintained.

In autumn 1987 the pair was moved to an aviary of its own provided with perches and two nesting platforms. At the beginning of January 1988 both birds began to build a nest on one of the plat- forms and on 21 January the first copula- tion was observed on the nest. Nest building activities became more intense and the pair copulated several times daily during which they both uttered high- pitched vocalisations.

Both birds of the pair had been hand- reared using the method described by Mendelssohn & Marder (1983) and their reproductive behaviour developed normally. They were not shy and could be observed at a distance of 3 m from an observation platform erected outside the wire netting of the aviary; a hide was not necessary.

The nest had a base of large branches, thinner branches on the inside and a lining of dry grass and wool. The Q did not make moulding movements to shape the nestcup like the Griffon vulture 0 Gyps fulvus (Mendelssohn & Leshem,

1983b) but this could have been due to a difference in the type of nesting material.

It was easy to recognise the sexes. The 0 was larger and had many creamy-white feathers on her back and shoulders. In three other young 99 these conspicuous feathers have been seen to appear in the second year, reach their maximum development in the fourth and fifth year, and decrease in number thereafter. Ten- year-old 99 are almost entirely dark brown. The breeding $2 is the fourth to show the feature. Since the feathers appear to reach their maximum develop- ment at the age of pair formation, it is possible that they aid in sex identification. Immature 88 never revealed this charac- ter. Cramp & Simmons (1980) state that ‘mantle and back sometimes variably intermixed with buff or off-white feathers’ but they do not relate the description to subspecies, age or sex.

On 12 February 1988 the 0 remained on the nest for the whole day arranging nesting materials and sometimes squatting on her tarsometatarsi but not lying down. In the morning of I3 February an egg was laid and incubated by the 0. The 8 took over the incubation the following day but we removed the egg for artificial incuba- tion in order to stimulate a replacement clutch. It was found to be infertile.

In the following weeks the pair con- tinued to nest build and copulate but at a lower intensity than previously. On 9 March a second egg was laid and incu- bated by the 9 and then the d as before. The interval between the first and second egg was shorter than has been observed in the Griffon vulture in which the period between loss of egg or chick and the replacement clutch is 25-32 days (Mendelssohn & Leshem, 1983b).

The pair shared the incubation. The incubating bird crouched very low in the nest, as described by Jennings & Fryer (1984), and was almost completely hidden by the high rim. By contrast, incubating Griffon vultures sit rather high in the nest. Occasionally the incubating bird would stand up, look at the egg and sometimes

232 NEW DEVELOPMENTS I N T H E ZOO WORLD

Plate I . Lappet-faced vulture T o r p s rruchehtus negrwnsis pair at Tel Aviv University Research Zoo with 20-day-old chick. The is on the left of the photograph. A . Shooh.

loosen the lining of the nestcup with half- open bill in a way similar to storks.

The egg pipped on 2 May and hatched on 3 May giving an incubation period o f 55 days. confirming the previous esti- mated record in Cramp & Simmons (1980) and one day longer than the maxi- mum estimate in Brown Pt al. ( 1982). The nestling was brooded but feeding did not begin until 4May when the dismem- bered and consumed a rat except for the skin. stomach and intestine. She then drank some water. flew to the nest and fed the chick. by regurgitation. from bill tip to bill tip. This proved to be the general pattern of feeding during which she produced copious amounts of saliva which were given with the food or some- times drops were passed to the chick. For the next 1 1 days only the 't: was seen to feed the nestling. She accepted only rats and guinea pigs to feed to the nestling and both parents ate the remnants. At this stage neither adult ate other meat. During this period the 3 undertook the larger

part of brooding and shading of the chick as required.

On 15 May the skin of a rat was seen on the rim of the nest and the following morning the 9 was seen carrying a rat to the nest in her bill. Standing in the nest she dismembered the rat and passed pieces to the nestling. The pieces were quite large and. as far as could be seen from a distance of 3m, contained bone and skin. The young bird had difficulty in swallowing the piece with the tail attached but when the 0 tried to pull the tail back, the chick appeared to be stimulated to greater effort and succeeded in swallowing the piece whole. Like some other raptors, the Lappet-faced vulture thus shows two methods of feeding its young; regurgi- tation and direct feeding. In contrast Griffon vultures appear to feed only by regurgitation except for meat containing bones which is carried in the bill to the nestling (Mendelssohn & Leshem, 1983b).

fed the young for the first time when it was 12 days old and from then on

The

N E W DEVELOPMENTS IN T H E ZOO WOKLD 233

both parents fed the chick by regurgi- tation; only in a few cases were the rats taken to the nest and dismembered. When the nestling was two weeks old the parents began to take meat again and apparently fed the chick regurgitated meat. When the chick was 12-14 days old the ambient temperatures rose to 40°C and it received only a few meals.

On the morning of 25May, we observed for the first time, the parents feeding the nestling by regurgitation before they were fed, and it appears there- fore that the food must have been stored in their stomachs from the day before. This behaviour is shown regularly by Griffon vultures when feeding small chicks (Mendelssohn & Leshem, 1983b).

At one month the chick seemed to be better developed than hand-reared chicks of the same age (Mendelssohn & Marder, 1983). It was now fed on regurgitated meat but rats were taken to the nest and fed to it. During the first two weeks only one parent was on the nest, for most of the time, brooding or shading the chick as required. Later both parents were often observed standing on the nest, one of them shading, the nest being exposed to the sun until noon each day, and both frequently looking at the chick with lowered heads. Unlike the Griffon vultures which preen small chicks often (Mendelssohn & Leshem, 1983b), they rarely preened their offspring and the young bird was seen to preen itself for the first time at the age of 17 days.

On day 34 the young vulture was left alone for the first time in the afternoon when there was no need for shading. In the mornings it was, however, continually shaded until it was fully feathered at the end of July. From then on it was left alone, exposed to the sun, and was constantly panting. Even when the young was still being shaded by its parents it panted frequently and apparently the parents’ saliva given with the food could not compensate for the loss of body water because when the chick was one month old the parents had already begun to give

it water from the crop. Later it was given water daily, more frequently than has been seen in Griffon vultures.

The development of the young bird has been described (Mendelssohn & Marder, 1983). The 1988 young left the nest on 1 September at 131 days old, compared with the 125-135 days recorded in Brown ef al. (1982). For six days it sat on the ground and on the fourth day one of the parents brought it a rat. After eight days it began to fly and for several days it frequented the nest. Although the parents often perched close together after the young was fledged, it showed no tendency to remain close to them. Antagonistic behaviour was directed towards them one month after it had left the nest. In nature the young Lappet-faced vulture stays with its parents for several months (Cramp & Simmons, 1980; Brown et al., 1982; Mendelssohn & Leshem, 1983b).

On 15November 1988 the 9 again began to carry branches to the nest starting the next reproductive cycle.

ACKNOWLEDGEMENTS

Our sincere thanks are extended to Mr Ben-Adaret and Mr Bartuv, to keepers at the CCEZ who provided some of the observations and to Ms N . Paz who edited and typed the manuscript. This study was supported by the Israel Cohen Chair of Environmental Zoology.

REFERENCES BROWN, L. H., URBAN, E. K. & NEWMAN, K. (1982): The birds ofAfrica 1. London, New York: Academic Press. BRUUN, B. (1981): The lappet-faced vulture in the Middle East. Sandgrouse 2 91-95. BRUUN (BRUNN-SIC), B., MENDELSSOHN, H. BULL, J. (1981): A new subspecies of lappet-faced vulture Torgos tracheliolirs from the Negev Desert, Israel. Bull. Br. O m . Club 101: 244-247. CRAMP, S. & SIMMONS, K. E. L. (1980): Hundbook o/ the birds of Europe, the Middle East and North A.fricu: the birds of the western Paleurctic 2. Oxford: Oxford University Press. JENNINGS, M. C. & FRYER, R. N. (1984): Birds of Saudi Arabia. The occurrence of the lappet-faced vulture Torgos /rarheliorus (J. F. Forster) in the Arabian Peninsula, with new breeding records from Saudi Arabia. Faun. Saudi Arab. 6 534545. LESHEM, Y . (1983): The rapid population decline of Israel’s lappet-faced vulture Torgos trarheliotus negevensis. Int. Zoo Yb. 2 3 41-46,

234 N E W I)EVELOPMENTS IN THE ZOO WORLD

MEN~XLSSOHS. H. & LESHEM. Y. (I983a): Status and conservation of vultures in Israel. In C'ulture hiolugj, und mtmagemc~nt: 86--98. Wilbur. S. R. & Jackson. J. A. (Eds). Los Angeles: University of California Press. MENDI.LSSOHN. H. B LESHEM. Y . (1983b): Obsenations on reproduction and growth of Old World vultures. In I.'ulture hiologx and r,iaiiugrniutir:

214-231. Wilbur, S. R. & Jackson. J . A. (Eds). Los Angeles: University of California Press. MENDELSSOHN, H. & MARDER, U. (1983): Hand- rearing Israel's lappet-faced vulture Torgox rrachcliotus negevensis for future captive breeding. In!. Zoo Yh. 2 3 47-51,

Manuscript submitted 23 January 1989

[ The Zoological Society of London

Breeding the Himalayan griffon

at the Paris Menagerie G j . p himaIuj*ensi.s

MARSHA A. SCHLEE MusPwn Nutionul d'Hisroire Nururelle, Menagerie rlu Jurdin des Pluntes, 57 rue Cuvier, 75231 Paris Cedes 05, Frtince

The Himalayan griffon Gj-ps hiniuluj,ensis is a resident of the high mountain ranges of Cmtral Asia from Afghanistan to Bhutan, including the Pamirs. Turkestan and Tibet (Brown & Amadon. 1968). I t is the largest of the griffon vultures and, with ~1 weight comparable to that of the Andean Condor, is one of the heaviest flying birds in the world (Houston. 1983). It feeds entirely on carrion, usually ungu- late carcasses. and nests on cliff ledges in small colonies of four to six pairs (Whistler. 1963).

Several successful breedings of the Himalayan griffon have been achieved at the East Berlin Zoo (Minnemann. 1984). To d d e , however, we have found no publication dealing with this species' breeding behaviour in captivity and few details are available on nesting in the wild (Brown & Amadon, 1968).

The breeding pair at the Paris Mena- gerie had already reared a chick, in 1982 when both birds were c.17 years old, at the Reserve de la Haute-Touche. In June 1983 they were transferred to Paris follouing completion of the new raptor

exhibit. The pair started breeding the following year and have hatched a chick every year since then, successfully rearing the young in 1984, 1985, 1987 and 1988. Most of the observations reported here concern the 1986 and 1987 breeding seasons. when one chick was reared successfully and the other died a t 72 days of age, with some additional information based on the 1985 and 1989 chicks.

HOUSING A N D DIET The griffons are housed outside through- out the year and are not sheltered in bad weather or periods of low temperatures. Their enclosure measures 7.6 x 8.7 x 5.8 m high; an artificial rock cliff runs up the back and a nest cave, which is never used, is located to one side at about 2.6 m from the ground. The enclosure is provided with a pond, a tree and logs placed across the face of the cliff for perching. A nest site consisting of a wiremesh platform (c. 1 x 1.3 m) covered with birchwood branches and straw, and bordered with logs was installed on the ledge above the