ZURICH ZOO: GIANT TORTOISE HUSBANDRY 135

REFERENCES ADLER, H. J. (1991a): Conservation program of Chrotogale owstoni Thomas, 1912: some first results. Mustelid and viverrid conservation. Newsletter of the IUCN M VSG 4 8. ADLER, H. J. (1991b): On the situation of the Dela- cour's langur (Trachypithecus 1: delacouri) in the north of Vietnam: proposal for a survey and conser- vation project. Primate Rep. No.31: 6-7. ADLER, H. J. & PETER, W. P. (1993): New conser- vation strategies for endangered primates in Vietnam: a memorandum of agreement and an open letter to the Ministry of Forestry. Primate Rep. No.37: 69-12. ADLER, H. J. PETER, W. P. (Unpublished): Cuc Phuong National Park conservation program, north Vietnam. Project proposal. Internal Report of the Zoological Society of Frankfurt, FrankfurdMain, 1991.

Zoologische Gesellschaft fur Populations-und Arten- schutz, Miinchen 8(2): 1-5. ADLER, H. J. WIRTH, R. (1994): Species conser- vation priorities in Vietnam and the potential role of zoos. In Creative conservation: interactive manage- ment of wild and captive animals: 495-501. Olney, P.J.S., Mace, G.M. & Feistner, A.T.C. (Eds). London: Chapman & Hall. GROOMBRIDGE, B. (Ed.) (1993): 1994 IUCN red list of threatened animals. Gland and Cambridge: IUCN. MACKINNON, J. (1985): Review of the protected area system and species conservation needs of Vietnam. Gland: WWF. RATAJSZCZAK, R., ADLER, H. J. & SMIELOWSKI, J. (1993): The Vietnamese sika Cervus nippon pseudaxis conservation project. Int. Zoo Yb. 32: 56-60. STUART, S. & WIRTH, R. (1990): Preliminary project proposal: conservation and recovery of threatened species in Vietnam. Gland: IUCN.

ADLER, H. J. & WIRTH, R. (1992): Das Erhaltung- szuchtprojekt fur den Vietnamsika. Mitteilungen Manuscript submitted 23 November 1994

Int. Zoo Yb. (1995) 34: 135-143 0 The Zoological Society of London

Management of Giant tortoises

at Zurich Zoological Gardens Geochelone elephantopus and Geochelone gigantea

M. CASARES', R. E. HONEGGERZ & A. RUBEL3 'Department of Zoo Animals & Exotic Pets, University of Zurich, 8057 Zurich, 2Curator of Herpetology and 3Director, Zurich Zoological Gardens, Ziirichbergstrasse 221, 8044 Zurich, Switzerland

At Zurich Zoological Gardens 3.1 Geochelone ele- phantopus and 2.4 G. gigantea are maintained in an enclosure comprised of a 400 m2 outdoor area, which contains a shelter with a heated floor to which the tortoises have constant access, and an adjacent 65 mz indoor house with a partially heated floor, where the temperature reaches 30°C. In 1984 the diet offered was adjusted to a lower protein, higher fibre com- position after one animal died from chronic kidney disease and visceral gout. No eggs have been pro- duced by G. gigantea while maintained at the Zoo because the 99 have failed to ovulate which may be an indicator of unfavourable environmental condi- tions, such as low ambient temperatures. Since 1989 six G. elephantopus have hatched at the Zoo. Details

'All reprint requests to Mr R. E. Honegger.

of the management and reproduction of the colony are discussed.

The giant tortoises of the genus Geoche- lone, Fitzinger 1835, are the largest living terrestrial chelonians. Originally wide- spread on many islands of the western Indian Ocean and the Galapagos archi- pelago of the Pacific Ocean, over the last two centuries human exploitation has ren- dered all endemic species of the Indian Ocean islands extinct except the Aldabra Atoll Geochelone gigantea (Stoddart &

136 THE DEVELOPING ZOO WORLD

Peake, 1979). Although this species appears to be secure at present, like all island species it is vulnerable to natural or man-made disasters because of its extremely limited range. The populations of the Galapagos tortoises Geochelone ele- phantopus have also been severely deci- mated and the 11 surviving races are still threatened by predation andlor competi- tion from introduced mammals (Mac- Farland et a l , 1974). The species is classified in Appendix I by CITES and Vulnerable by IUCN (Groombridge, 1993).

The future of giant tortoises in their natural range remains unpredictable and there are relatively few reports of breeding successes outside their native islands: Gal- apagos giant tortoises have bred at Hon- olulu (Throp, 1975), San Diego (Bacon, 1980), Seffner (Noegel & Moss, 1989), Brownsville (Hairston & Burchfield, 1989) and Zurich (Honegger & Rubel, 1991); Aldabra giant tortoises have bred at Sydney (Peters & Finnie, 1979), Jackson- ville (Collins, 1985) and Stanford (Stearns, 1988) (see also Table 1).

After the first hatching of two Gala- pagos tortoises in Zurich in 1989, addi- tional G. elephantopus hatched in 1992 (one) and 1993 (three) [in 1995 (20)]. However, this initial success must be eval- uated cautiously.

The giant tortoise colony at the Zurich Zoological Gardens comprises 3.1 G. elephantopus ssp and 2.4 G. gigantea. The breeding G. elephantopus was obtained in 1946 from the Galapagos islands (exact origin unknown but prob- ably Santa Cruz) and was identified as G. e. porteri on the basis of her morpho- logical characteristics. The $$ were obtained in 1962 at which time they weighed 0.65, 2.35 and 7.45 kg. Two were collected in Northern Isabela and were believed to be G. e. becki. Both of these tortoises exhibited a degree of saddle- backed shape although they were not true saddlebacks. The identification of the third 8 remains unclear; although it is pre-

sumed that it was collected in Santa Cruz, and is therefore believed to be a G. e. porteri, its shape is unusual. How- ever, this might be related to either a high protein diet or a lack of ultraviolet light as a juvenile (S. Thomson, pers. comm.). All the Aldabra tortoises G. gigantea were originally obtained from Aldabra.

ENCLOSURES The exhibit comprises an inside and an outside enclosure connected with gates. Both enclosures have a 3.2 m2 cement-bot- tomed pool containing fresh water c. 30 cm deep.

The 400m2 outdoor enclosure is planted with shrubs and grass and is sur- rounded by a 1 m high cement boundary. Various logs and mounds are provided to encourage the animals to climb. There is a block-house shelter 5 x 2.5 x 1 m high with a heated concrete floor, to which the tortoises have constant access. Electric heat lamps are fixed to the ceiling and covered with a metal grille to protect the animals from the bulbs. Although accli- matized adults are fairly cold resistant and can tolerate temperatures near to 0°C (Collins, 1985; Stearns, 1988), when day- time temperatures drop below 15°C their movements become extremely slow and it is necessary to move them into the heated shelter at night.

In autumn, when the weather becomes colder, the indoor enclosure is opened and the tortoises either move in of their own accord or they may need to be lured in with food. These 65 m2 winter quarters have a partially heated floor where the temperature reaches 30°C. A soil substrate area of 6 m2 and 0.5 m deep is provided as a nesting site. This substrate forms an adhesive but workable mud when wet, as recommended by MacFarland et al. (1974). Infra-red and ultraviolet lamps were installed in 1988 to increase light and temperature locally. The lamps are sus- pended at a height of 2.5 m and are angled downwards towards an area where a small amount of grass, leaves or lettuce is

ZURICH ZOO: GIANT TORTOISE HUSBANDRY 137

Geochrlone gigantea IUCN (R) 1984 Jacksonville USA 3 1989 Kuwait 6(5) 1990 Atlanta USA 4 1992 Seffner USA 19

Geochelone elephantopus IUCN (v) 1985 Brownsville USA 6; Philadelphia USA (1); San Diego USA 21 1986 Brownsville USA 1 1987 Philadelphia USA 0.1; Phoenix USA 4; San Diego USA 4; Seffner USA 19(17) 1988 Phoenix USA 1 1989 Brownsville USA 3; Seffner USA 10( 1); Zurich Switz 2 1990 Oklahoma USA 5; Seffner USA 10* 1991 Brownsville USA 4; Oklahoma USA 2; Seffner USA 23 1992 Oklahoma USA 38(2); Phoenix USA 1; Seffner USA 94*; Zurich Switz 1 1993 Ahmedabad Ind 3; Oklahoma USA 23; Seffner USA 63; Zurich Switz 3

G. e. elephantopus (= vicina) 1987 Brownsville USA 6 1989 Brownsville USA 2; Seffner USA 27(5) 1990 Seffner USA 7(1)* 1992 Hamilton Berm 1 l(3)

G. e. guntheri 1989 Seffner USA 34(3) 1990 Seffner USA 8

G. e. microphyes 1990 Brownsville USA 1

G. e. porteri 1990 Seffner USA 2*

* An asterisk after the name of the institution concerned denotes that one or more of the animals born there was bred in the second or subsequent captive generation. Table 1. Collections that have bred Geochelone elephantopus and G. gigantea as listed in the Yearbook Volumes 26-34.

offered twice daily. The lamps are switched on for 30 minutes while the animals are fed and the temperature at the top of the carapace can reach 32°C.

Because of the cold natural climate of Switzerland, the tortoises are maintained indoors for approximately six months from October to May. On warm days they have access to a small outside area (40 m2). During this period the indoor air temperature ranges from 24 to 28°C. Temperature directly on the heated floor area, frequently occupied by the animals, is 32°C.

Positioned on the roof of the winter quarters are ten 0.48 x 0.12 m windows.

However, indoor and outdoor measure- ments of light intensity, carried out regu- larly between 1300 and 1400 hours from April to September 1994, demonstrated a considerable loss of light intensity inside the house. On sunny days, while outdoor light intensity reached 120 000 lux, the indoor readings did not exceed 70 000 lux (42% loss); when the sky was overcast, 13 000-60 000 lux, indoor light intensity ranged from 4000 to 12 000 lux (69-80% loss).

FEEDING After the death of an active and appar- ently healthy G. gigantea (carapace length

138 THE DEVELOPLNG ZOO WORLD

80 cm) on 20 December 1983, from severe chronic interstitial and glomerular nephritis and severe visceral gout, we altered the diet offered to the tortoises based on observations by R.E.H. in 1962, 196311964 and 1972 in the field, and the information by Swingland & Coe (1978). The new diet was of improved quality and reduced quantity. Green corn and minced meat was no longer offered and we added CarniconB and more Ca-grit (Table 2).

Up to 900/0 of the outside yard is planted with grass which is an important feeding base during the summer. After overwintering indoors, the tortoises are conditioned to grazing with a mixture of grass and hay in early May. Food is never offered in one feeding trough but is ran- domly scattered around the enclosure which serves to keep the tortoises active and in public view.

DOMINANCE BEHAVIOUR AND COURTSHIP The Galapagos tortoise $6 are frequently involved in confrontations with each other, especially between either JlO or $20 and 830. During these agonistic interactions, the largest tortoise ($30) typically adopts an imposing posture with its neck and legs fully extended (neck extend), opens its mouth and moves towards its opponent without completing

the bite (gape, bite attempt). Although dominance should be determined by the height reached in the vertical neck exten- sion (Schafer & Krekorian, 1983), the smaller J& never adopt the neck extend but usually bite 630’s head, neck or fore- limbs without showing any other behav- ioural patterns. As a result $30 withdraws his neck and head into the carapace (retract) and moves away from his oppo- nent (retreat). The apparently superior status of the smaller G. e. becki is con- sistent with the observations of behav- ioural differences reported between the more aggressive saddle-backed (e.g. $10 and $20) and the dome-shelled (J30) races of G. elephantopus (Schafer & Kre- korian, 1983).

The limited agonistic behaviour observed in G. gigantea may be related to the high population density in its natural habitat (Grubb, 1971). Occasionally $50 has been observed to interfere in the mating activity of $40 by pushing him.

Courtship can take place throughout the year but it reaches a peak in August and September. Noegel & Moss (1989) reported that 9 G. elephantopus come into oestrus before the nesting season begins, evidenced by a swelling of the cloaca and a discharge of a gelatinous substance. In contrast, gelatinous discharges are occa- sionally observed all year round at Zurich

FOOD 1970-1983 ‘Al VOL. FOOD 1984PRESENT ‘ 4 VOL.

Lettuce Bananas Tomatoes Apples, pears Minced meat Ca-grit Green corn Grass

15 20 30 30

5 + S

S’

Kale, red cabbage Carrots Vegetables (various) Carob Shredded horn, shrimpmeal Ca-grit CarniconB Grass, fresh leaves or dried leaves, chopped hay

15 45 30

5 3 1.5 0.5 S’

W

’Food items offered three times per week. Table 2. Volume of food offered to the Giant tortoises Geochelune spp at Zurich Zoological Park. s = summer, w =winter.

ZURICH ZOO: GIANT TORTOISE HUSBANDRY 139

and do not correlate to the reproductive cycle. The sexes are housed together con- tinuously and most mating attempts recorded were unsuccessful. In 1992 the 9 G. elephantopus was removed from the group for two months because of an injury to its carapace. On reintroduction to the group on 31 August a successful copulation was immediately observed. It has been indicated that separation of the sexes may be an important factor in stim- ulating breeding in the giant tortoises (Throp, 1975; Hairston & Burchfield, 1989; Noegel & Moss, 1989) but Mac- Farland et al. (1974) found that it was not necessary to separate G. elephantopus in order to achieve successful mating.

Bacon (1980) suggested that social hier- archy may influence courtship and mating patterns in G. elephantopus. However, the Galapagos 0 at Zurich has an obvious preference for $30 which appears to be low on the social scale. The 0 usually responds to the mating attempts or imposing postures of other 68 by biting them or walking away. A possible explanation is that the smaller $10 and 320 lack the physical strength necessary to overpower and dominate the $? (Hair- ston & Burchfield, 1989). Moreover, Nib- lick et al. (1994) reported that 8 size may be an important factor in choice in the Desert tortoise Gophevus agassizii. The mating activity of G. gigantea appears to be promiscuous.

There is minimal interaction between the Galapagos and the Aldabra giant tor- toises. Sometimes the aggressive 610 G. elephantopus was observed to mount the largest $2 G. gigantea, even though the penis was not erect and the tail was not pushed under the 9's carapace. This 9 has occasionally had aggressive behaviour directed at it from the 0 G. elephantopus, and has also been observed attempting to mount the smallest G. gigantea.

REPRODUCTION The gonadal cycles and morphology of wild Aldabra giant tortoises have been

investigated (Bourn, 1977; Swingland & Coe, 1978) but information relating to the reproductive biology of the Galapagos tortoises remains extremely limited, mainly because their threatened status restricts the use of techniques for studying reproductive patterns. Non-invasive methods, such as ultrasound scanning, have recently been applied to endangered chelonian species for evaluation of ovarian function (Robeck et al., 1990; Kuchling & Bradshaw, 1993; Rostal et al., 1994). In order to monitor the ovarian activity of the giant tortoises at Zurich, the animals were routinely examined using ultrasound scanning over two years, from late 1992 to 1994. A detailed discussion of these findings are the subject of another paper. In contrast to the seasonal folli- cular cyclicity of wild populations of Aldabra tortoises (Swingland & Coe, 1978), both large vitellogenic and atretic ovarian follicles were found all year round in the captive animals. The four G. gigantea at Zurich (one arrived in 1955, another in 198 1 and two in 1984) have not laid eggs because they have failed to ovu- late, although follicles of preovulatory size were frequently observed. Ovulation is sensitive to unfavourable environmental factors (Kuchling & Bradshaw, 1993) and the failure of our animals to ovulate may suggest sub-optimal captive conditions. Furthermore, the nesting season of cap- tive G. gigantea in the Northern Hemi- sphere begins in October/November (Collins, 1985; Stearns, 1988) when various factors like decreasing photope- riod, especially as tortoises are moved into the winter quarters, and temperature might have a negative effect on ovulation. In fact, extensive ovarian atresia was observed in G. gigantea between Sep- tember and November. Otherwise, atresia was more abundant in G. elephantopus following the nesting season, which is a general feature of post-breeding reptilian ovaries (Licht, 1984), suggesting that folli- cular atresia may temporarily suspend breeding activity when the environmental

140 THE DEVELOPING ZOO WORLD

conditions are not appropriate for the optimum survival of offspring.

Little is known about the natural sper- matogenic cycle of giant tortoises and detailed information on the environ- mental cues underlying their seasonality is lacking. Temperature appears to represent an important timing cue for breeding cycles in reptiles and has been linked to the seasonal gonadal cycles of other che- lonian species (Licht, 1984). In mature J G. gigantea testes weight increases from September to October (the onset of the hot and wet season) and reaches a peak in February, one to two months before the mating peak, which occurs in MarchlApril (Swingland & Coe, 1978). Because peak .enlargement of the testes occurs at the final stages of testis recrudescence, the spermatogenic cycle of G. gigantea may therefore develop mainly during the hot season, when temperatures reach a mean maximum of 31°C from November to April (Bourn, 1977). Most copulation of wild G. eleplzantopus also occurs during March and April (Hendrickson, 1966). However, the hot and wet season in the Galapagos Islands lasts from January to the end of April and average daily maxima do not reach 30°C until February (MacFarland et al., 1974). If the testicular cycle of G. elephantopus followed a pat- tern similar to that of G. gigantea, the testes growth would take place at least partially during the preceding dry and cooler season. Thus, testis recrudescence in G. elephantopus may require lower tem- peratures than it does in G. gigantea. The spermatogenic cycle of 3 tortoises main- tained at sub-optimal temperatures can be seriously disrupted (Kuchling et al., 1981) and this may be associated with the poor success in captive breeding of G. gigantea compared with G. elephantopus, because at most zoological collections it is not usual to have natural temperatures of 30°C over several months. However, the possibility of influence of environmental factors other than temperature (e.g. pho-

toperiod) or endogenous components, cannot be ruled out.

In order to check 3 fertility, moist secretions around the cloaca1 orifice of either 6 or 9 were collected after court- ships and copulations and these were examined with a microscope for the pres- ence of spermatozoa. Spermatozoan mor- phology has been described previously (Olsen, 1976; Platz et al., 1980). While sperm of apparently normal morphology were observed in G. elephantopus 330, samples of G. gigantea exhibited a high percentage of sperm abnormalities, including abnormal heads and middle portions, cytoplasmic droplets and strongly coiled tails.

NESTING AND INCUBATION The nesting season of G. elephantopus occurs from October to February, although recently ovulated eggs were detected by ultrasound as early as August. Several days before egg laying, the 9 spent much time wandering round and sniffing the nesting area. The nesting and laying process occurred overnight and has only been observed once. On 30 November 1992, the 0 was first observed excavating the nest with her hind feet at 1530 hours. Ten eggs were laid between 1625 and 1642 hours at intervals of one-to-four minutes and the nest was finally covered with soil at 2045 hours. Occasionally the $2 would lay an egg on the surface without excavating a nest. Oxytocin has never been used to stimulate oviposition.

Some of the eggs of the three clutches laid in the 199311994 season had a thick shells and were heavier (over 150 g) than the others, suggesting that they had originated from different ovulations. Geo- chelone elephantopus is capable of retaining shelled eggs until the next breeding season (Noegel & Moss, 1989). Up to four clutches were produced during a nesting season with a highly variable inter-clutch interval, from 43 to 96 days (mean 61). Clutch size ranged from five to 15 eggs (mean 8.6).

ZURICH ZOO- GIANT TORTOISE HUSBANDRY 141

Staff carefully excavated the nesting area every two-to-three days throughout the breeding season. Eggs were removed from the nest, marked with a small pencil cross on the top and placed in plastic boxes with a 15 crn layer of moistened vermiculite in which the eggs were par- tially buried. Incubation temperatures were 30-32°C with 65% relative humidity. The top of the incubator was removed daily for a few minutes in order to allow a replenishment of fresh air. Hatchlings started to emerge after 121-161 days (mean 139.5) and escaped from the egg three to five days later.

Because several eggs were broken during laying or staff searching activities, it was difficult to estimate the fertility of the clutches but it appears to be much lower than in the Galapagos Islands, either in the wild or at the Charles Darwin Station (MacFarland et al., 1974). For example, from a clutch of ten eggs laid in November 1992, three hatchlings emerged successfully and the other seven eggs were infertile. Throp (1975) noted that fertility could be high in some clutches but totally absent in others.

CARE OF HATCHLINGS The hatchlings remain in their plastic boxes in the incubator for several days until the yolk sac is completely absorbed. They are then moved to a floor-heated ter- rarium, measuring 120 x 40 x 30 cm high, with a gravel substrate. Only pebbles over 5 mm in diameter are used as a substrate in order to avoid ingestion, which may in turn cause impaction. Two 40 W black- light lamps and a 60 W heat lamp (12:12 hours 1ight:dark) are placed 25 cm above the surface of the terrarium. On warm days juveniles are placed in a secure out- door enclosure planted with grass to allow grazing and to expose them to natural solar radiation.

The young are weighed and measured once a month to monitor growth (Table 3). In the first year their weight could increase to between 350 and 650 g.

~~

MONTH WEIGHT (9) MEAN

ID 1 ID 2 ID 3 ID4

1

1 2 3 4 5 6 7 8 9 .o 11 12 13 14 15 16 17 18 19 20

78 83

106 126 138 175 224 271 320 335 429 529 580 665 876 927

1073 1103 1278 1465

85 I13 144 177 225 266 280 305 321 366 463 538 620 144 808

1024 1144 1305 1524 1 a49

81 102 126 147 179 235 267 300 334 395 497 546 638 704 705 91 1 929

1079 1198 1342

76 94

123 154 191 242 298 329 362 410 497 554 650 754 778 955

1016 1162 1338 1515

80 98

124.75 151 183.25 2295 267.25 301.25 334.25 376.5 471.5 541.75 622 716.75 791.75 954.25

1040.5 1162.25 1334.5 1542.75

Table 3. Growth of four Galapagos tortoises G. elephantopus hatched at Zurich Zoological Gardens.

The tortoise born in June 1992 weighed 2125 g by the end of its second year.

The first two young born at Zurich in May 1992 died suddenly after 14 and 15 months. One showed chronic inter- stitial pneumonia and the other showed slight interstitial nephritis and thymitis. In addition, both animals developed signs of metabolic bone disease. Generalized bone metabolism disorders (rickets combined with osteodystrophia fibrosa) were also recorded in four G. gigantea which arrived at Zurich in 1964 at the estimated age of six months and which were euthanased at between ten and 12 years (Hauser et al., 1977). It is noteworthy that the three young G. elephantopus obtained in 1962 and maintained on the same regimes as the G. gigantea obtained in 1964, have shown no symptoms of metabolic bone disease.

The young tortoises were originally offered an adult diet made with smaller pieces of food but this was changed to a

142 THE DEVELOPING ZOO WORLD

lower protein, higher fibre diet and the volume was reduced. The vegetables vary depending on season between chopped parsley, basil, sage, celery, fennel, cab- bage, red cabbage, dandelion, vetch and mulberry Morus spp leaves; the sliced car- rots are dusted with crushed sepia or egg- shells. Three times per week each tortoise receives about one tablespoon of curds mixed into the chopped vegetables. Hay and other dried leaves and water were available ad libitum.

ACKNOWLEDGEMENTS

This study was supported in part by the Swiss National Science Foundation, Project No. 31- 29979.90. We particularly appreciate the collabora- tion of Scott Thomson, University of Melbourne, who identified the Galapagos tortoises to subspecies level and made constructive observations. We also thank Professor Dr E. lsenbiigel and Professor Dr K. Zerobin for their co-operation, and the members of the tortoise keeping staff at Zurich Zoo for their interest and help.

REFERENCES BACON, J. P. (1980): Some observations on the cap- tive management of Galapagos tortoises. In Repro- ductive biology and diseases of captive reptiles: 97-113. Murphy, J. B. & Collins, J. T. (Eds). Society for the Study of Amphibians and Reptiles, Contri- butions to Herpetology No. 1, USA. BOURN, D. (1977): Reproductive study of giant tor- toises on Aldabra. J. Zool., Lond. 182: 27-38. COLLINS, D. E. (1985): Captive breeding and management of the Aldabra tortoise, Geochelone gi- gantea. Proceedings of 8th annual reptile symposium on captive propagation and husbandry: 16-84. Hahn, R. A. (Co-ord.). Thurmont, MD: Reptile Sym- posium Zoological Consortium. GROOMBRIDGE, B. (Ed.) (1993): 1994 IUCN red list of threatened animals. Gland and Cambridge: IUCN. GRUBB, P. (1971): The growth, ecology and popu- lation structure of giant tortoises on Aldabra. Phil. Trans. Roy. Soc. Lond. B. 260 321-312. HAIRSTON, C. & BURCHFIELD, P. M. (1989): Manage- ment and reproduction of the Galapagos tortoise Geochelone elephantopus at the Gladys Porter Zoo. Int. Zoo Yb. 28: 70-17. HAUSER, B., METTLER, F. & HONEGGER, R. E. (1977): Knochenstoffwechselstorungen bei Seychellen- Riesenschildkroten (Testudo [Geochelone] gigan- tea). Proc. 19th Int. Symp. on Diseases of Zoo and Wild Animals, I977 1 9 121-125. HENDRICKSON, J. D. (1966): The Galapagos tor- toises, Geochelone Fitzinger 1835 (Testudo Linnaeus 1758 in part). In Proceedings of the symposium on

Galapagos international scientijic project: 252-257. Bowman, R. I. [Ed.). Berkley, Los Angeles: Univer- sity of California Press. HONEGGER, R. E. & RUBEL, A. (1991): Aufzucht und Erkrankungen der ersten in Europa nachgezogenen Galapagos-Schildkroten (Geochelone elephantopus). Proc. 4th Int. Coll. Pathol. Med. Rept. Amph., 1991 4 225-243. KUCHLING, G. BRADSHAW, S. D. (1993): Ovarian cycle and egg production of the western swamp tor- toise Pseudemydura umbrina (Testudines: Chelidae) in the wild and in captivity. J. Zool., Lond. 229 4054 19. KUCHLING, G., SKOLEK-WINNISCH, R. & BAMBERG, E. (1981): Histochemical and biochemical investi- gation on the annual cycle of testis, epididymis, and plasma testosterone of the tortoise Testudo hermanni hermanni Gmelin. Gen. Comp. Endocrinol. 44: 194-201. LICHT, P. (1984): Reptiles. In Marschall's physiology of reproduction. 1. Reproductive cycles of vertebrates: 206282. Lamming, G. E. (Ed.). Edinburgh: Chur- chill Livingstone. MACFARLAND, C. G., VILLA, J. & TORO, B. (1914): The Galapagos giant tortoise (Geochelone elephant- opus). 11. Conservation methods. Bid. Conserv. 6 198-212. NIBLICK, H. A., ROSTAL, D. C. & CLASSEN, T. (1994): Role of male-male interactions and female choice in the mating system of the desert tortoise, Gopherus agassizii. Herpetol. Monogr. 8: 124-132. NOEGEL, R. P. & Moss, G. A. (1989): Breeding the Galapagos tortoise Geochelone elephantopus at the Life Fellowship Bird Sanctuary, Seffner. Int. Zoo

OLSEN, J. H. (1976): Notes on Galapagos tortoise reproduction. Zoonooz 49(4): 13. PETERS, U. W. & FINNIE, E. P. (1979): First breeding of the Aldabra tortoise Geochelone gignntea at Sydney Zoo. Int. Zoo Yb. 19 53-55. PLATZ, C. C., JR, MENGDEN, G., QUINN, H., WOOD, F. & WOOD, J. (1980): Semen collection, evaluation and freezing in the green sea turtle, Galapagos tor- toise, and red-eared pond turtle. A. Proc. Am. Assoc. Zoo Vet. 1980 47-54. ROBECK, T. R., ROSTAL, D. C., BURCHFIELD, P. M., OWENS, D. W. & KRAEMER, D. C. (1990): Ultra- sound imaging of reproductive organs and eggs in Galapagos tortoises, Geochelone elephantopus spp. Zoo Biol. 9: 349-359. ROSTAL, D. C., LANCE, V. A., GRUMBLES, J. S. &

ALBERTS, A. C. (1994): Seasonal reproductive cycle of the desert tortoise (Gopherus agassizii) in the Eastern Mojave Desert. Herpetol. Monogr. 8: 72-82. SCHAFER, S. F. & KREKORIAN, C . 0. (1983): Ago- nistic behavior of the Galapagos tortoise, Geochelone elephantopus, with emphasis on its relationship to saddle-backed shell shape. Herpetologica 39 448456. STEARNS, B. C. (1988): Captive husbandry and prop- agation of the Aldabra giant tortoise Geochelone gi-

Yb. 28: 18-83.

GSCC SAVI’S PIPISTRELLE BREEDING 143

gunteu at the Institute for Herpetological Research. Int. ZOO Yb. 21: 98-103. STODDART, D. R. PEAKE, J. F. (1979): Historical records of Indian Ocean giant tortoise populations. Phil. Trans. Roy. Soc. Lond. B. 286 147-161. SWINGLAND, I. R. & COE, M. (1978): The natural regulation of giant tortoise populations on Aldabra Atoll. Reproduction. J. Zool., Lond. 186 285-309.

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THROP, J. L. C. (1975): Note on the management and reproduction of the Galapagos tortoise at the Honolulu Zoo. In Breeding endangered species in captivity: 3942. Martin, R. D. (Ed.). London: Academic Press.

Manuscript submitted 9 November 1994

0 The Zoological Society of London

Rearing and first reproduction of the Savi‘s pipistrelle

at Group of Study and Conservation Chiroptera, Florence

PipistreIIus savii

G. DONDINI & S . VERGARI Group of Study and Conservation Chiroptera (GSCC), WWF Tuscany, 50100 Florence, Italy

In July 1993 a young Savi’s pipistrelle Pipistrehs savii 9 was taken into the Group of Study and Con- servation Chiroptera, Tuscany. In September 1993 she was observed mating with a S in the colony and two months later the 9 went into three-and-a-half months’ hibernation. In July 1994 she gave birth to 1.1 young. While the 9 was nursing the normal diet for adults, Tenebrio molitor larvae raised on protein- enriched bran, was supplemented with milk for human babies to avoid the possibility of calcium deficiency which could have an effect on ossification in the neonates. The young had completed their growth in four weeks and were capable of flight by 3540 days.

The Savi’s pipistrelle Pipistrellus savii, a member of the Vespertilionidae, is a Med- iterranean and mid-Asian bat, with a range from the Iberian Peninsula to north-eastern China and Japan, via the Caucasus and Mongolia, and on to Iran, Afghanistan and northern India in the south-east (Lanza & Finotello, 1985). Little information is available on the status of the species which is considered threatened in Europe (Stebbings & Grif- fith, 1986). Savi’s pipistrelle can be found

in coastal areas and mountainous areas with altitudes over 2500 m. Roosting sites during the summer are usually holes in buildings, caves and, occasionally, trees; during winter the bats hibernate in deep crevices, roof tiles or abandoned chim- neys. Pipistrellus savii occasionally mi- grates and can travel as far as 250 km from its original site (Schober & Grimm- berger, 1993).

In Italy Savi’s pipistrelles have been recorded in every region except Basilicata (Vernier, 1987) but there is no precise information on the frequency of the species; over time it has been classified as frequent (Lanza, 1959) and, almost 30 years later, less common, at least in the northern regions (Vernier, 1987).

The species is small, insectivorous and hibernates. An adult’s forearm measures between 31 and 38 mm long, with a maximum wing-span of 230mm and a weight ranging between 4 and log. The ear is broad and rounded, with a short

Address for correspondence: Via Vittoria 39, 51026 Maresca (PT), Italy.