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Published Papers: Studies on Released Rehabilitated Hedgehogs
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The Veterinary Record

Wildpro Reference Code: J3.138.w2

Health and welfare of rehabilitated juvenile hedgehogs

(Erinaceus europaeus) before and after release into the wild

A. W. Sainsbury, A. A. Cunningham, J. K. Kirkwood, S. K. Macgregor, P. A. Morris

Materials and Methods
Results
Discussion
Acknowledgements
References

Thirteen wild caught juvenile hedgehogs were treated and overwintered in a rehabilitation centre and 12 were released into the wild and monitored by radiotelemetry. Clinical examinations were carried out before they were released and twice afterwards, and any hedgehogs found dead were examined post mortem. The health of the animals was generally good but dental disease, obesity and minor injuries were common. One hedgehog died before it was released and had cardiovascular and respiratory lesions, and one was euthanased 28 days after its release and had verminous enteritis and parasitic bronchitis. Three hedgehogs were killed by badgers, two were killed by road traffic and four were known to have survived when the study ended.

An increasing number of free-living wild mammals and birds are being taken into captivity for the treatment of illness or injury, with the intention of returning them to the wild. Considerable resources are invested by individuals and welfare organisations in this activity. Many authors have drawn attention to the potential welfare and disease hazards (Harris 1989, Lewis and Stocker 1993). However, few studies have been undertaken, either in the United Kingdom or elsewhere, on the fate of mammals or birds after their release, and the effect it may have on their health and other aspects of welfare (Kirkwood 1993). The authors know of no detailed investigations which have been carried out on the effect that these releases may have on the resident population of the same species, either through aggressive interactions, competition for resources or by influencing their disease status.

The European hedgehog (Erinaceus europaeus) is probably the most frequently rehabilitated mammalian species in the UK (British Wildlife Rehabilitation Council 1993). Morris and others (1993a, b) investigated the survival of released adult hedgehogs, which had had previous experience of the wild, and found that the animals managed to feed and apparently behave normally; Morris and others (1993b) found that of four hedgehogs released one had died within three weeks and, in a separate study, eight hedgehogs were released of which three died and four were unaccounted for by 12 weeks after their release (Morris and others 1993a). However, in both studies, the deaths were not necessarily linked to the release, for example, road traffic killed some of the hedgehogs but many wild hedgehogs die from this cause every year. The annual mortality of wild adult hedgehogs is approximately 30 per cent (P. A. M., unpublished observations). Neither of the above studies investigated in detail the health of the hedgehogs before or after their release, and the causes of most of the deaths were not investigated post mortem.

A large percentage of the hedgehogs 'rescued' by rehabilitation centres are juveniles, sometimes nestlings, which are taken into captivity in the autumn. These may have been abandoned by their dam or may have been judged to have insufficient fat resources to survive hibernation. Morris (1984) showed that hedgehogs which weigh less than 450 g in the autumn are unlikely to survive the winter. Whatever their origin, many of these young hedgehogs have had little experience of life in the wild when they are released in the following spring.

This report describes part of an investigation into the fate of juvenile hedgehogs released into the wild after having been overwintered in a rehabilitation centre. Morris and Warwick (1994) have reported the findings on the behaviour, body mass and survival of these hedgehogs, and this report describes how the health of the hedgehogs was affected by their release, the causes of deaths and whether the hedgehogs harboured any infectious agents which might affect the welfare of the resident population. It includes the results of clinical examinations carried out before and on two occasions after the release of the hedgehogs, and the results of post mortem examinations of the hedgehogs which did not survive.

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Materials and methods

Thirteen hedgehogs, seven males and six females, had been overwintered at the Royal Society for the Prevention of Cruelty to Animals Wildlife Hospital at West Hatch, Somerset. They had weighed less than 350 g when taken into captivity the previous autumn. They were anaesthetised on April 2, 1993 for a clinical examination and fitted with radiotransmitters (Morris and Warwick 1994) so that they could be tracked.

The animals were released at Helecombe Farm, near Clayhanger in Devon (NGR ST 0224) where previous uncontrolled releases had taken place. The date of release varied for each animal (Morris and Warwick 1994). Six of the hedgehogs (four females and two males, 221, 244, 270, 298, 312 and 324) were released directly but six (two females and four males, 211, 228, 256, 288, 332 and 353) were held in pre-release cages for five nights before they were released as described by Morris and Warwick (1994). One male (341) died while it was in the pre-release cage. The movements of the hedgehogs were monitored every night after their release for approximately five weeks by radiotracking and less frequently thereafter for approximately four weeks. They were caught for examination and weighed, usually every night for at least the first four weeks, and less often thereafter. Resident wild hedgehogs found during the monitoring procedures were weighed and marked individually by spraying a patch of spines with paint (Morris and Warwick 1994). Behavioural interactions between resident and released hedgehogs were recorded.

Clinical examinations

The 13 hedgehogs were anaesthetised for a clinical examination either on the day of their release or up to 10 days before. Seven of them were re-examined five weeks after the release programme started and four of these were examined again two weeks later.

Anaesthesia was induced in an anaesthetic chamber with 4 per cent halothane (Halothane; RMB Animal Health) and maintained with approximately 2 per cent halothane administered through a malleable face mask. The bodyweight of the hedgehogs was measured and the condition of each animal was scored on the basis of the amount of subcutaneous fat and the muscle thickness of the abdominal wall and limbs as follows: 1 thin, no subcutaneous fat and wasted musculature; 2 moderate, no subcutaneous fat but good muscle thickness; 3 good, some subcutaneous fat; 4 fat, a thick covering of subcutaneous fat.

The oral cavity was examined for signs of disease including dental disease. Scoring systems were used for recording the severity of dental calculus in the oral cavity as follows: 1 mild calculus on some teeth; 2 mild calculus on all teeth; 3 severe calculus on some teeth; 4 severe calculus on all teeth. For gingivitis the scoring was: 1 mild inflammation of some gingival margins; 2 mild inflammation of all gingival margins; 3 severe inflammation of some gingival margins; 4 severe inflammation of all gingival margins.

The chest was auscultated for signs of abnormalities of the cardiovascular and respiratory systems. The abdomen was palpated for abnormalities and to detect pregnancy. The animals were examined for signs of diarrhoea and rectal swabs were taken for bacteriological examination. Faecal samples were also collected when possible for parasitological examination. Before they were released faecal samples were obtained from hedgehogs 256, 288 and 353, and a further 10 samples were taken from cages containing more than one animal. On the second and third examinations, samples of faeces were obtained from two and four animals, respectively. The skin, musculature, eyes and ears were examined and samples for other diagnostic tests were taken as appropriate.

Microbiology and parasitology

Faecal swabs were cultured for bacteria by plating on to 5 per cent horse blood agar (Oxoid CM331; Unipath) and xylose lysine desoxycholate medium (Oxoid CM469). When appropriate, skin samples were plated on to Sabouraud's dextrose agar with chloramphenicol (Oxoid CM41) for fungal isolation. When the faeces were unformed or liquid, blood-free Campylobacter selective agar (Oxoid CM739) was used. The unformed faeces were also examined for the presence of cryptosporidia by the modified Ziehl-Neelsen technique, using a direct smear of a faecal homogenate. Direct smears were prepared from all the faecal samples and examined microscopically for parasite larvae and ova. The McMaster technique was used to estimate the number of nematode eggs per gram of faeces.

Pathology

Detailed systematic post mortem examinations were carried out on four hedgehogs which died during the course of the study. Bacteriological and parasitological examinations were carried out on the rectal contents by the methods described above. Tissues for histological examination were fixed in neutral buffered 10 per cent formalin, embedded in paraffin wax, sectioned and stained with haematoxylin and eosin.

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Results

The results of the clinical and post mortem examinations and the fate of each hedgehog are given in Table 1.

Four of the hedgehogs were obese before their release and had approximately 5 mm of subcutaneous fat (based on the post mortem findings from 341). Their mean (sd) bodyweight was 861.7 (176.4) g whereas, in a previous study, 14 juveniles which had overwintered in the wild weighed 562.0 (114.9) g in April (Morris and Warwick 1994). The released hedgehogs lost up to 38 per cent of their pre-release bodyweight and the heaviest animals lost the most, both in absolute terms and relative to their own initial weight (Morris and Warwick 1994). After approximately 30 days there was little loss in bodyweight and the hedgehogs which were alive at the end of the study had bodyweights substantially greater than the average weight of first year animals that have overwintered in the wild.

Calculus and gingivitis were common findings and were severe in one case in which the gingivae bled on being touched. Eight of the 13 hedgehogs had cuts, abrasions or subcutaneous abscesses or evidence of recently healed injuries before they were released. These minor injuries were treated with antiseptic (povidone iodine) and topical chlortetracycline hydrochloride (Aureomycin topical powder; Cyanamid) when required. Hedgehogs 332 and 353 were treated with ampicillin (Amfipen; Gist-brocades Animal Health) by intramuscular injection before they were released, for an inflamed prepuce and an inflamed foreleg, respectively. All of the injuries were considered to be due to bite wounds caused by other hedgehogs. After they were released, the injuries that were examined during routine nightly monitoring or at subsequent health assessments, were found to have healed without complications.

Nematode ova or larvae were identified in the faeces of two (256 and 353) of the 11 hedgehogs of which the faeces were examined before they were released and in five of the eight animals from which faeces or rectal contents were examined after their release (221, 228, 256, 270 and 332). Escherichia coli, Proteus mirabilis and Plesiomonas shigelloides were commonly isolated from the rectal contents or faeces taken before release but after release the isolates were more diverse and included fungi (Candida species). All the faecal samples were negative for Campylobacter, Salmonella and Cryptosporidia species. Hedgehog 256 was returned to the rehabilitation centre 17 days after its release owing to lethargy and a pronounced loss of bodyweight and it was subsequently euthanased. It was found to have a verminous gastroenteritis, parasitic bronchitis and chronic interstitial nephritis. The gastritis and enteritis were associated with large numbers of nematode ova (Capillaria species) in the small and large intestine and with adults in the small intestine. Klebsiella pneumoniae pneumoniae, E coli and P mirabilis were isolated from the intestine. This animal was found to have a Capillaria species infestation before it was released and was in moderate condition, although comparable with others normally released by the rehabilitation centre.
g
Table 1. Results of clinical and pathological examinations of the hedgehogs on 2/4/93 before their release, and at later dates.
Number of hedgehog and sex Date of examination Bodyweight (g) Condition score Calculus score Gingivitis score Other clinical findings Fate and post mortem findings
211
M
2/4/93 536 3 2 2 - -
6/5/93 505 2 3 1 - -
10/5/95 - - - - - Killed by a badger after five weeks
221
F
2/4/93 1038 4 1 1 Evidence of recent wound on shoulder which was healing -
6/5/93 710 3 1 1 - -
21/5/93 700 2 2 2 100 epg faeces: nematode (strongyle) ova. Positive pregnancy diagnosis  -
12/6/93 - - - - - Survived until the end of the study
228
F
2/4/93 858 3 1 0 Subcutaneous abscess on foreleg  -
6/5/93 685 2 3 1 Recent wound on head, subcutaneous abscess healed; positive pregnancy diagnosis; no nematodes identified in faeces -
21/5/93 695 2 3 1 - -
12/6/93 - - - - Positive pregnancy diagnosis; nematode larvae in faeces Survived until the end of the study
244
F
2/4/93 1036 3 1 1 - -
4/93 - - - - - Radiotransmitter lost four weeks after release
256
F

 

2/4/93

 

584

2

3

0

200 epg faeces: Capillaria species -
7/5/93 350 - - - - Euthanased on 6/5/93 following return to rehabilitation centre on 25/4/93. Parasitic gastroenteritis; parasitic bronchitis; interstitial nephritis; 10,800 epg rectal contents: Capillaria species; Capillaria species ova and adults in small intestine; Klebsiella pneumoniae pneumoniae, Escherichia coli and Proteus mirabilis isolated from intestine
270
M
2/4/93 1118 4 1 1 Evidence of recent wound on shoulder which was healing; one incisor fractured -
6/5/93 800 2 1 1 - -
21/5/93 760 2 3 2 200 epg faeces: Capillaria species; nematode larvae in faeces -
12/6/93 - - - - - Survived until the end of the study
288
M
2/4/93 908 3 2 0 Subcutaneous abscess on hindleg; other abrasions on legs; tick located on neck -
6/5/93 695 2 1 1 - -
10/5/93 - - - - - Killed by a badger after five weeks
298
F
2/4/93 948 3 4 3 Three teeth with severe wear; two upper molars lost -
6/5/93 785 2 2 1 As above for dentition; no nematodes identified in faeces -
10/5/93 - - - - - Killed by a badger after five weeks
312
F
2/4/93 1014 4 2 0 Ventral abdominal skin abrasions -
6/5/93 795 3 1 1 Positive pregnancy diagnosis -
21/5/93 690 2 1 1 Negative pregnancy diagnosis; no nematodes in faeces -
12/6/93 - - - - - Survived until the end of the study
324
M
2/4/93 816 2 2 0 Subcutaneous abscess in ventral abdominal area -
10/5/93 380 - - - - Died four weeks after release following a road accident. Multiple fractures of the skull, proximal four ribs and left scapula; avulsion of head from cervical spine, associated soft tissue damage and haemorrhaging; thoracic organs lost by post mortem scavenging; stomach and proximal small intestine empty; no abdominal or subcutaneous fat; no nematodes identified from rectal contents
332
M
2/4/93 980 3 1 0 Subcutaneous abscess on foreleg, swelling of prepuce -
28/4/93 - - - - - Dies five days after release following a road accident. 100 epg rectal contents: Capillaria  species; 100 epg rectal contents: nematodes (strongyle); fractures of the maxilla, mandible, cranium, head if the humerus, distal radius, distal ulna and left sixth rib; soft tissue damage and haemorrhage associated; spleen enlarged and congested; semi-digested invertebrates including earthworms in stomach; subcutaneous fat and axillary fat present
341
M
2/4/93 1330 4 2 0 Numerous abrasions of ventral, abdominal and thoracic skin -
13/4/93 1199 - - - - Died under anaesthesia before release. Obese; approximately 5 mm thick subcutaneous fat deposits and a rotund appearance to body due to intra-abdominal fat; intestinal tract empty except for a small amount of hay in the stomach; chronic abcessation of the lung with broncopneumonia; chronic low-grade myocarditis  
353
M
2/4/93 652 3 2 0 Swollen and inflamed left foreleg; subcutaneous abscess of lower jaw; 300 epg faeces: Capillaria species -
4/93 - - - - - Radiotransmitter lost three weeks after release

Hedgehog 324 was killed by a car four weeks after its release. It had already lost 35 per cent of its bodyweight and its condition had been of concern before its death. Its stomach and small intestine were empty, suggesting that it had not fed recently, and there was no subcutaneous or abdominal fat. Hedgehog 341 died while under anaesthesia when its radio-transmitter was being fitted, before it was released. It had chronic abscessation of the lung with bronchopneumonia and a chronic low-grade myocarditis. Its intestinal tract was empty.

No serious aggressive encounters were observed among any of the hedgehogs, resident or released, after the release had occurred (Morris and Warwick 1994). Courtship and mating behaviour between resident and released hedgehogs was observed (Morris and Warwick 1994).

Three animals were found to be pregnant 26, 29 and 49 days after their release, suggesting that they had conceived after they were released, because the gestation period of the hedgehog is 31 to 35 days (Deanesly 1934); furthermore the hedgehogs had been housed as single sex groups while in captivity. In each of the pregnant hedgehogs there were five to 10, 5 to 10 mm diameter, roughly circular, tonic conceptual swellings in the uterus, separated by a similar length of uterine body. One animal (228) was palpated again 15 days later and the swellings had grown to approximately 15 x 40 mm tubular-shaped bodies and showed significantly less uterine tone. Another hedgehog (312) was not pregnant 15 days after a positive diagnosis but it is not known whether it had given birth or had aborted.

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Discussion

The study identified two substantial health problems: periodontal disease and injuries. These problems indicate that the animals' management while in captivity could be improved. All the 13 hedgehogs had peridontal disease when they were first examined, and hedgehog 298 had severe calculus and gingivitis and had lost several teeth. This was a surprising finding considering their young age. The incidence and prevalence of periodontal disease in wild hedgehogs has not been investigated but it would seem unlikely that it would be high in young animals. It is possible that the soft diet offered at the rehabilitation centre led to a build up of plaque on the teeth, and subsequently to the formation of calculus and gingivitis (Kertesz 1993). The diet included puppy dog food (Puppy Chum; Pedigree Pet Foods), day-old chicks and a mix designed for insectivorous birds (Prosecto; J. Haiths). A more natural diet of beetles, caterpillars, earthworms, flies, spiders, centipedes and slugs (Yalden 1976) might be better, but the production and supply of this diet would be impractical. At the time of the first health assessment it was considered that the preferable course of action for the hedgehogs would be to return them to the wild where they would receive a more natural diet which would improve their oral health. There is a clear need to investigate and provide a more suitable diet for captive hedgehogs. Dental disease is painful and in the long term could affect the survival of hedgehogs by reducing their feeding efficiency. In addition, many people place food in their gardens for wild hedgehogs (Morris 1983) and it is important to ensure that such supplementary diets do not cause dental disease.

Cuts, abrasions and subcutaneous abscesses, probably caused by bites from other hedgehogs, were common before the hedgehogs were released, and although these minor injuries responded well to treatment, methods of husbandry that reduce the chances of aggressive behaviour should be used. The requirements for the housing of hedgehogs in captivity have not been well documented although there is some information (Morris 1967).

The health of the hedgehogs after their release was generally good but several findings indicated suboptimal welfare.  Both Capillaria species and strongyle type (Crenosoma species) nematodes have been previously recorded from hedgehogs in the UK (Boag and Fowler 1988, Majeed and others 1989, Keymer and others 1991). In the present study ova of Capillaria species nematodes were more commonly found in the faeces than the ova and larvae of strongyle nematodes. Although the species were not identified and the life cycles are not known for certain, it is likely that most of the hedgehogs had an infestation of egg-producing adults before their release because there was only a short interval between the animals' release and the detection of nematodes in the faeces. The verminous gastroenteritis and parasitic bronchitis found post mortem in hedgehog 256 suggests that further monitoring of parasitic infestations during the over-wintering period in the rehabilitation centre would be useful and possibly that those found to be infested should be treated before release to increase their chances of survival in the wild. In general, the bacteriological and mycological examinations yielded no organisms considered to be of clinical significance but the intestinal flora of hedgehog 256 may have played a secondary role in the pathogenesis of the gastroenteritis observed post mortem.

The poor condition of hedgehog 324 may have had a bearing on its inability to avoid a collision. Unfortunately, the carcase had been partially eaten by scavengers and it was not possible to establish the cause of the loss of bodyweight. Despite the severe head and other injuries inflicted on hedgehog 332 by a vehicle they were not evident externally in the field at night; the animal was found alive and only noted as dead two hours later. It is known that large numbers of hedgehogs die in road collisions and, in a review, Morris (1993) quoted estimates for the UK of between 100,000 and 1.3 million such casualties annually. The lesions of bronchopneumonia and myocarditis found post mortem in hedgehog 341 were thought to have been severe enough to have caused its death under anaesthesia; it had been particularly active in the pre-release cage in an attempt to escape, which may explain the absence of intestinal contents, and the resultant stress may have affected its survival under anaesthetic. The benefits or otherwise of the use of pre-release cages in this study were discussed by Morris and Warwick (1994).

The ability to reproduce is considered to be a sign of good welfare (Moberg 1985). Although three hedgehogs were found to be pregnant, neither their litter sizes nor the survival of the young were measured.

If possible, obesity is avoided in captive animals. However, obesity may have improved the survival chances of the hedgehogs released in this study. Assuming a daily maintenance requirement of 145 kcal/day, then, for example, 300 g of fat deposits would provide for approximately 19 days without food (Kirkwood 1991). The three hedgehogs which weighed more than 1000 g before being released survived to the end of the study, but an analysis of weight in comparison with survival is complicated by the causes of death; the risk of predation and road accidents may not be affected by weight.

Three hedgehogs were killed by badgers. In two cases the predation was established from an examination of the carcase and in the third case the badger was encountered in the act (Morris and Warwick 1994). The relative susceptibility of the hedgehogs to this cause of death was discussed by Morris and Warwick (1994).

There was no evidence that the release of the hedgehogs adversely affected the welfare of the resident wild hedgehogs. Despite the release of 12 additional animals at a site where hedgehogs were present, no aggressive encounters were observed between the released and wild animals (Morris and Warwick 1994). A single wound, thought to be the result of an aggressive interaction between hedgehogs, was found on one female (228). It is likely that the resident animals were already exposed to the infectious agents identified in the hedgehogs before their release; nematodes have been described in wild hedgehogs (Boag and Fowler 1988). Studies in which both the released and the resident animals are monitored in detail are required to clarify this issue.

The release process in this study was unusual in several ways; one animal (256) found to be ill after its release was returned to the rehabilitation centre; another (298) remained within a farm building for three days and had to be 'rescued' while a third was retrieved after having moved out of the monitored area of farmland (Morris and Warwick 1994). These interventions were necessary for welfare or practical reasons, but in similar non-monitored releases these animals might have suffered compromised welfare or died. Similarly, the hedgehog which died while under anaesthesia, with cardiovascular and respiratory lesions, would have been released without knowledge of its poor health.

Interventions for the welfare of wild animals are becoming more common as popular concern for wildlife grows. The potential for disease to affect both the rehabilitated animals and their wild counterparts should not be underestimated, and the husbandry and veterinary care in rehabilitation centres should be of a high standard to reduce this possibility. The released animals may affect the welfare of the wild population by, for example, acting as a source of infectious or parasitic disease, or by competitive or aggressive interactions. In release programmes for hedgehogs there is a need to identify and monitor parasitic infestations before the hedgehogs are released and treatment should be considered in cases where heavy infestations may be compromising health. The development of a diet suitable for maintaining hedgehogs in captivity, which does not cause periodontal disease would be of value. Investigations into the housing of hedgehogs in captivity should pay particular attention to group size and social structure to prevent or reduce aggressive interactions and fight wounds. Mortality due to road accidents and predation reduced the numbers of juvenile hedgehogs released in this study. Further studies are required to determine the fate of juvenile hedgehogs after their release and to investigate the effect that the release of hedgehogs has on the welfare of the wild population.

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Acknowledgements

The authors would like to thank Ms Louisa Gammidge, Miss Tracy Howard, Miss Judith Howlett and Mr Hugh Warwick for their technical support and Mr Wayne Smith-Gillard for his help with this study. They are also grateful for the assistance given by Mr Colin Seddon, the staff of the RSPCA Wildlife Hospital, West Hatch, Somerset and Mr Paul Yates in providing facilities for, and help during, the assessment and recording of the health of the animals. Mr and Mrs Swanston generously allowed access to their farm for the study.

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References

Boag B & Fowler P A (1988) Journal of zoology, London 215, 379

British Wildlife Rehabilitation Council (1993) Wildlife Casualty Recording Scheme Quarterly Return, January-March 1993

Deansley R (1934) Philosophical Transactions of the Royal Society B 223, 239

Harris S (1989) Proceedings of the Inaugural Symposium of the British Wildlife Rehabilitation Council. Eds S. Harris, T. Thomas. London, British Wildlife Rehabilitation Council. p 94

Kertesz P (1993) Veterinary Dentistry and Oral Surgery. London, Wolfe Publishing. p 73

Keymer I F, Gibson E A & Reynolds D J (1991) Veterinary Record 128, 245

Kirkwood J K (1991) Journal of Nutrition 121 (IIS), S29

Kirkwood J K (1993) Veterinary Record 132, 235

Lewis J C M & Stocker L (1993) Veterinary Record 133, 23

Majeed S K, Morris P A & Cooper J E (1989) Journal of Comparative Pathology 100, 27

Moberg G P (1985) Animal Stress. Ed G. P. Moberg. Bethesda, American Physiological Society. p 245

Morris B (1967) The UFAW Handbook: the Care and Management of Laboratory Animals. 3rd edn. Edinburgh, E & S Livingstone. p 478

Morris P A (1983) Hedgehogs. London, Whittet Books. p 70

Morris P A (1984) Journal of zoology, London 203, 291

Morris P A (1993) A Red Data Book for British Mammals. London, Mammal Society. p 10

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

Morris P A & Warwick H (1994) Animal Welfare 3,163

Yalden D W (1 976) Acta Theriologica 21, 401


A. W Sainsbury, BVetMed, CertLAS, MRCVS,  A. A. Cunningham, BVMS, MRCVS, J. K. Kirkwood, BVSc, PhD, MRCVS, S. K. Macgregor, HNC, MISCT, Veterinary Science Group, Institute of Zoology, Zoological Society of London, Regent's Park, London, NW1 4RY, P. A. Morris, PhD, Department of Biology, Royal Holloway and Bedford New College, University of London, Egham, Surrey TW20 0EX

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