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The results of this study demonstrated that in ponies in which KLEIN, L. (1978) Proceedings of the 24th Annual Convention of the American
anaesthesia was induced with xylazine, ketamine and diazepam, Association of Equine Practitioners. p 89
LEE, Y-H., CLARKE, K. W. & ALIBHAI, H. 1. K. (1993) Jourtnal of Veterinary
both the co and the intramuscular blood flow were better main- Anaesthesia 20, 41
tained during anaesthesia with isoflurane than with halothane. LEE, Y-H., CLARKE, K. W. & ALIBHAI, H. 1. K. (1998) American Jouirnal of
However, the differences were small, and only became significant Veterinary Research 59, 1463
with the change from isoflurane to halothane anaesthesia. The LINDSAY, W. A., PASCOE, P. J., McDONELL, W. N. & BURGESS, M. L. F.
(1985) A,nerican Journal of Veterinarv Research 46, 686
change from halothane to isoflurane anaesthesia did not signifi- LINDSAY, W. A., ROBINSON, G. M., BRUNSON, D. B. & MAJORS, L. J. (1989)
cantly improve cardiovascular function, but did prevent further American Journial of Veterinary Research 50, 404
deterioration. The study also showed that under conditions in MATTHEWS, N. S., HARTSFIELD, S. M., CORNICK, J. L., JACOBSON, J. D. &
which the intracompartmental pressure within the muscles might WILLIAMS, J. D. ( 1992) Cornell Veterinarian 82, 21
MONTEIRO, A. A., SVENSSON, H., BORNMYR, S., ARBORELIUS, M. &
be presumed to have been low, the changes in intramuscular blood KOPP, S. (1989) Archives of Oral Biology 34, 779
flow were unrelated to changes in MAP, but followed the changes PAVLIN, E. G. & SU, J. Y. (1994) Anaesthesia. 4th edn. Ed R. D. Miller. New
in cardiac output. York, Churchill Livingstone. p 125
RICHEY, M. T., HOLLAND, M. S., McGRATH, C., DODMAN, N. H., MAR-
Acknowledgements. - This project was supported by the Horserace SHALL, D. B., COURT, M. H., NORMAN, W. M. & SEELER, D. C. (1990)
Veterinary Surgers 19, 392
Betting Levy Board. The authors would like to thank Vickers SERTEYN, D., COPPENS, P., MOTTART, E., MICHELET, S., MICHEELS, J.,
Medical UK for the loan of the Datex Capnomac Ultima, Kontron PHILIPPART, C. & LAMY, M., (1987) Veterinary Record 122, 324
UK for the loan of the halothane monitor and Moor Instruments UK SIGURDSON, G. H., BANIC, A., WHEATLEY, A. M. & METTLER, D. (1994)
for the loan of the laser Doppler flowmeter. Y-H. L. was in receipt British Journal of Anaesthesia 73, 826
STEFFEY, E. P. & HOWLAND, D. Jr. (1980) American Journal of Veteriniary
of a Scholarship from the Korean Ministry of Education and UK Research 41, 821
Overseas Research Studentship Scheme. H. I. K. A. was in receipt STEFFEY, E. P., HOWLAND, D. Jr., GIRI, S. & EGER, E. 1. (1977) Attleric-an
of a Fellowship from the Danish International Aid Agency. Expert Journal of Veterinary Researc h 38, 1037
technical assistance was provided by Gillian Hughes. STEFFEY, E. P., KELLY, A. B. & WOLINER, M. J. (1987a) American Jouirnal of
Veterinary Research 48, 952
STEFFEY, E. P., HODGSON, D. S., DUNLOP, C. I., MILLER, M. F., WOLINER,
M. J., HEATH, R. B. & GRANDY, J. (1987b) Journal of Veterinary
References Pharnacology and Therapeutics 10, 290
CLARKE, K. W. & GERRING, E. E. L. (1990) Proceedings of the 35th Annual STEFFEY, E. P. & PASCOE, P. J. (1994) Journal of Veterinary Anaesthesia 21, 43
Convention of the American Association of Equine Practitioners. p 629 STILL, J., SERTEYN, D. & VAN DER MERWE, C. A. (1996) Journial of the Solth
DUNLOP, C. I., STEFFEY, E. P., MILLER, M. F. & WOLINER, M. J. (1987) African Veterinary Associationi 67, 199
American Jouirnal of Veterinary Research 48, 1250 TAYLOR, P. M. (1991) Journal of Veterinary Anaesthesia 18, 8
GASTHUYS, F., MESSEMAN, C. & DE MOOR, A. (1994) Journal of Veterinarn TAYLOR, P. M. & YOUNG, S. S. (1990) Journial of VeterinaryAnaesthesia 17, 35
Anaesthesia 21, 60 TEVIK, A. (1983) Nordisk VeterinaerMedicinz 35, 175
GEIGY SCIENTIFIC TABLES (1994) 8th edn. Vol 5. Basle, Ciba Geigy. p 40 TRIM, C. M., ADAMS, J. G., COWGILL, L. M. & WARD, S. L. (1989) Equinle
GRANDY, J. L., STEFFEY, E. P., HODGSON, D. S. & WOLINER, M. J. (1987) Veterinary Journal Suppl 7, 84
American Journal of Veterinary Research 48, 192 TRIM, C. M. & WAN, P. Y. (I 990) Journal of Veterinary Anaesthesia 17, 45
HARVEY, R. C., GLEED, R. D., MATTHEW, N. S., TYNER, C. L., ERB, H. N. & WHITE, N. A. & SUAREZ, A. (1986) American Journal of Veterinary Reseairtch 47, 2257
SHORT, C. E. (1987) Veterinary Surgery 16, 184 WHITEHAIR, K. J., STEFFEY, E. P., WILLITS, N. H. & WOLINER, M. J. (1993)
JOHNSTON, G. M., TAYLOR, P. M., HOLMES, M. A. & WOOD, J. L. N. (1995) American Journial of Veterinary Research 54, 1693
Equine Veterinary Journal 27, 193 YOUNG, S. S. & TAYLOR, P. M. (1993) Equine Veterinary Journal 25, 147
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TABLE 1: Summary of survival of the 33 hedgehogs released in the ficient of variation of the bodyweight of three wild hedgehogs
three studies reviewed. All the animals released in Devon and Jersey
were overwintered juveniles with little or no previous experience of liv- weighed there each night for at least a week, was 6.6 per cent,
ing in the wild slightly more than for six released captives (4.5 per cent). Wide
daily fluctuations are natural and it is unwise to read much into
Alive after individual weight measurements, it is the trend over time which is
Hedgehog 4-5 weeks 6-7 weeks Cause of death significant. Being overweight at release is not necessarily bad for
Suffolk-210 Probably Unknown the animals. Extra fat reserves might help support released hedge-
Suffolk-215 Unknown Unknown hogs while they adjust to finding natural food or if they have to
Suffolk-229
cope with unexpectedly cold or dry conditions when such food is
Incomplete recovery scarce. They will also benefit from substantial fat reserves in
from treatment for
respiratory infection advance of hibernation.
Suffolk-249 Yes Yes During the Devon study, the nights became very wet and cold,
with temperatures around 0°C, posing an additional challenge to
Suffolk-274 Yes Yes Killed by car survival, because invertebrates are less active in the cold. Heavy
Suffolk-298 Yes Yes Probably drowned rain also threatens the ability of homoeotherms to maintain body
Suffolk-315 Probably Unknown temperature, especially if dry nesting sites cannot be located. It
Suffolk-350 Probably Unknown was for this reason that the animals had not been released even
Devon-211 Yes No Killed by badger earlier in the year. Some had sustained small bite wounds in cap-
Devon-221
tivity (Sainsbury and others 1996), but in every case these healed
Yes Yes quickly after release and several of the affected animals survived
Devon-228 Yes Yes until the end of the study. There is no evidence that these minor
Devon-244 Probably Unknown wounds represented a significant hazard, although it would have
Devon-256 Incomplete recovery been better had the animals not received them. For this reason, it
from treatment seems wise to cage the animals individually, particularly males in
euthanased after 17 the breeding season, before they are released.
days A regular change of nest site is normal in wild hedgehogs, espe-
Devon-270 Yes Yes cially among males (Reeve and Morris 1985). The released hedge-
Devon-288 Yes No Killed by badger hogs repeatedly found their way back to the nests they had built and
Devon-298 Yes No Killed by badger most re-used a previous nest at least once, indicating an ability to
Devon-312 Yes Yes orientate in a new habitat. Similar results were obtained in a study of
three hedgehogs released in Yorkshire (Morris and others 1993b).
Devon-324 Yes No Killed on road after 4
weeks Although none of the release studies showed a consistent pat-
Devon-332
tern suggestive of 'homing', several of the hedgehogs disappeared
No - - Killed on road after 5 from the study areas, having travelled considerable distances. In
days
Devon-353 Yes
Jersey, at least one moved over 5 km, and four of the Suffolk ani-
Unknown Unknown Radio lost after 3 mals were last seen over 2 km from the release point. Although
weeks
these animals were 'lost' from the study, their dispersal suggests
Jersey-21 0 Yes
that they had sufficient stamina to travel significant distances and
Jersey-220 Yes were thus probably healthy.
Jersey-227 Unknown In Cheshire, 22 rehabilitated hedgehogs were released into a
Jersey-242 Yes wood and radio tracked, but all of them departed to nearby vil-.
Jersey-255 Yes
lages within two weeks (Key 1997). A similar result was obtained
Jersey-282
when hedgehogs were released into a woodland in Surrey (N. J.
Yes
Reeve, personal communication). Dispersal may be a response to
Jersey-287 Yes being moved to an unfamiliar type of habitat, suggesting that
Jersey-298 Unknown hedgehogs should be released in the habitat to which they were
Jersey-31 1 Yes accustomed (Morris and others 1993a). However, in Jersey there
Jersey-324 Unknown
was no statistically significant difference between the behaviour
Jersey-331
of animals released close to where they had first been captured
Yes
and of those unfamiliar with the release site (Morris 1997). All the
Jersey-339 Yes animals in that study were juveniles, and were perhaps not old
Jersey-351 Yes enough to have become fixed in their behaviour towards their
home site before being taken into captivity.
Thirty of the 33 hedgehogs released were still alive after three
studies, there was frequent courtship behaviour between wild and weeks, long enough for them to have starved if they had been
released hedgehogs, often protracted, at the expense of foraging unable to obtain natural food. A high survival rate is particularly
behaviour. In one case (in Devon) it is believed that a successful significant because 25 of them were juveniles with little or no pre-
pregnancy resulted (Sainsbury and others 1996). vious experience of life in the wild. If they could be fully reinstat-
ed, then adults should also be capable of survival.
Although several hedgehogs died in the two months after they
Discussion were released, this is to be expected in a species whose average
life expectancy is about two years. Moreover, but for the care the
A consistent feature of these studies was weight loss, and the animals received in captivity, they would all have died much ear-
large percentage of bodyweight lost by some of the animals may lier. Over half not only survived, but some may also have bred.
suggest that they-were unable to find sufficient food and were Premature deaths were due largely to accidents (drowning, or
starving. However, the significant (P<0-001) relationship between being killed by traffic or badgers) unrelated to their health. After
bodyweight at release and maximum weight loss, indicates that an experimental release of 30 wild hedgehogs with no history of
these animals were shedding excess weight put on in captivity. ill health or veterinary care in to Wytham Wood in Oxfordshire,
For this species, weight loss should be interpreted carefully. 11 were killed by badgers or traffic within 10 weeks (Doncaster
Substantial changes are quite normal in hibernators like the 1992). Rehabilitated animals will inevitably be exposed to such
hedgehog (Kristoffersson and Suomalainen 1964) and even from dangers.
day to day, the weights of wild hedgehogs, studied simultaneously It has been argued that hedgehogs should not be released into
with the released animals in Suffolk, varied noticeably. The coef- areas frequented by badgers because of the likelihood of preda-
636 The Veterinary Record, December 5, 1998
tion. However, the most recent published distribution map mulation of tartar on the teeth. Claims that hedgehogs had con-
(Arnold 1993) shows that badgers are present throughout Wales tracted lungworm while in care as a result of being fed on a diet of
and most of southern England. They occur in all but two of the 10 dog food instead of earthworms were particularly misleading. In
km grid squares between Land's End and central Wiltshire. fact, earthworms are a natural source of lungworm, Capillaria eri-
Avoiding them, at least in the south west, is not a realistic option. nacei, common in wild hedgehogs (Romashov 1980).
Where badgers are scarce, it is often because the habitat is unsuit- These studies have clearly demonstrated that released hedge-
able, and it is frequently inadequate for hedgehogs too. hogs, even inexperienced juveniles, coped well with life in the
Similarly, avoiding traffic is unrealistic. Hedgehogs can travel wild, found food, nested normally and integrated well with resi-
up to 3 km in a night (Morris 1988), and few suitable areas in the dent wild hedgehogs, often in spite of bad weather and other
lowlands are more than 3 km from a road. The road deaths in adverse factors. The rehabilitation of hedgehogs is therefore a jus-
these studies occurred on tiny lanes, used by very few vehicles per tifiable use of resources to the extent that full integration into the
night. In all cases, wild hedgehogs were already present at the wild is an achievable aim. There is no evidence that welfare is sig-
release sites, irrespective of badgers or road traffic. nificantly at risk, except from accidents to which all hedgehogs
Drowning is a common cause of death in hedgehogs, not are exposed. However, captive hedgehogs should not be encour-
because they cannot swim, but usually because they fall into water aged to become tame and unwary of potential dangers. Substantial
and cannot climb steep smooth surfaces to escape. Such mishaps weight loss is to be expected after they are released, but this
are likely to occur irrespective of the animal's health or previous appears to be a natural readjustment to weight gain in captivity.
life in captivity. Nevertheless, despite all these difficulties, sur-
vival among rehabilitated hedgehogs was very high, compared, Acknowledgements. - The author thanks Hugh Warwick and
for example, with the minimal success of oiled seabirds after they Louisa Gammidge for field assistance in Devon, despite inclement
have been cleaned and released (Schmidt 1997). weather. Susan Sharafi and Kathy Meakin assisted in Suffolk; Dru
A probable risk factor is that the released hedgehogs were Burdon, Kim Matthews, Susan Sharafi and Ruth Temple in
accustomed to being handled. Many had been captive almost since Jersey. Mary Morris provided valuable assistance with data analy-
birth and were quite tame. They were insufficiently wary of sis, and Tony Sainsbury kindly read a draft of this paper. The con-
'attack' and several barely bothered to roll up when caught for tinued support of the British Hedgehog Preservation Society is
weighing each night. Tameness may increase the danger posed by gratefully acknowledged, as is help provided by 'Pro-Igel' of
badgers and road traffic. It would therefore seem wise to discour- Switzerland. The author is especially grateful to the RSPCA for
age tameness in animals intended for release. support, and to Pete and Jean Swanston for allowing access to
The assertion that rehabilitated animals should be released in their farm.
urban areas because they will be safer there remains unsubstanti-
ated. It may be true in respect of predation by badgers (although
badgers are common in some suburban areas), but avoiding one References
hazard may simply increase exposure to another. In the studies ARNOLD, H. R. (1993) Atlas of Mammals in Bntain. London, HMSO. p 145
BRIGHT, P. W. & MORRIS, P. A. (1994) Journal of Applied Ecology 31, 699
reviewed here, as many hedgehogs were lost to motor traffic as to BROCKIE, R. E. (1964) Nature 202, 1355
predators, despite the relative quietness of the country lanes in the DONCASTER, C. P. (1992) Proceedings of the Royal Society, London, B 249, 113
three study areas. The lanes were also narrow, slowing the few KEY, J. (1997) The Rehabilitator 24, 1
vehicles present after dark. In urban areas, traffic will be heavier, KRISTOFFERSSON, R. & SUOMALAINEN, P. (1964) Annales Academiae
Scientiarum Fennicae A, IV Biology 79, 1
and persist late into the night, almost certainly increasing the risk MORRIS, P. A. (1984) Journal of Zoology, London 203, 291
of accidental death. MORRIS, P. A. (1988) Journal of Zoology, London 214, 433
Studies of dormice (Muscardinus avellanarius) highlight the MORRIS, P. A. (1997) Animal Welfare 6, 317
benefit of 'soft release' methods involving use of pre-release MORRIS, P. A., MEAKIN, K. & SHARAFI, S. (1993a) Animal Welfare 2, 53
MORRIS, P. A., MUNN, S. & CRAIG WOOD, S. (1993b) Field Studies 8, 89
cages in which the animals can become accustomed to their new MORRIS, P. A. & WARWICK, H. (1994) Animal Welfare 3, 163
environment before release (Bright and Morris 1994). Some reha- REEVE, N. J. & MORRIS, P. A. (1985) Mammalia 49, 187
bilitators also take considerable trouble to accustom hedgehogs to ROMASHOV, B. V. (1980) Helminthologia 17, 181
the release site before they are freed, and provide supplementary SAINSBURY, A. W., CUNNINGHAM, A, A., MORRIS, P, A., KIRKWOOD, J. K.
& MACGREGOR, S. K. (1996) Veterinary Record 138, 61
food afterwards. There was no evidence that the provision of food SCHMIDT, K. (1997) New Scientist, May 3, p 41
or acclimatisation to the release site through the use of pre-release
cages had any beneficial effect. A similar result was obtained in
Yorkshire (Morris and others 1993b) where none of the three
released hedgehogs returned to feed at a bowl of dog food put out
for them for several nights, despite their continuing weight losses.
Nevertheless, soft release techniques which reduce the abruptness
of the transition from captivity to the wild are unlikely to be harm-
ful.
One of the Devon hedgehogs had defective teeth, which led to
Abstract
assertions that it should not have been released without 'effective Autonomic influences on cardiac
veterinary treatment'. However, dental anomalies (including miss-
ing teeth) were found in 39 of 77 wild E europaeus (Brockie rhythms in dogs
1964) and the hedgehog released in Devon showed no evidence of
progressive weight loss that might have been expected if it had THE autonomic nervous system has a fundamental influence on the
been hampered by dental deficiencies. It lost a maximum of 21 generation of normal and abnormal cardiac rhythms. For example,
per cent of its weight at release, then regained much of it, and the sinus arrhythmia and wandering pacemaker features of the
maintained its weight stable for three weeks at 81 to 90 per cent of canine electrocardiogram can be explained by the influence of
the initial weight, until it was killed by a badger. parasympathetic tone. Perturbations which enhance the sympathetic
The popularity of hedeghogs ensures that relevant press stories nervous system can also potentiate arrhythmias, or counteract
are widely read, and misleading reports and comments may have antiarrhythmic action. Disorders of the innervation to the heart may
significant negative effects. Articles in the press following the cause some life-threatening arrhythmias. This review considers the
publication of the paper by Sainsbury and others (1996) caused interactions between the autonomic nervous system and cardiac
confusion among animal carers and dismay concerning the validi- rhythms in normal dogs, and the specific arrhythmias observed in
ty of hedgehog rehabilitation. Animal carers became concerned boxers and German shepherd dogs, in each case relating the electro-
that dog food may be a dangerously inappropriate diet for hedge- physiological investigations to clinical practice.
hogs because press reporting emphasised a comment by Sainsbury
and others (1996) that eating soft dog food may lead to an accu- MOISE, N. S. (1998) Journal of Small Animal Practice 39, 460