general biologi sambar

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New Zealand Journal of Agricultural Research, 1994, Vol. 37: 79-85002 8-82 33/9 4/37 01-0 079 $2.50/0 The Royal Society of New Zealand 1994 79

General biology of sambar deer {Cervus unicolot) in captivity

G. SEMIADI1-2P. D. MUIR1T. N. BAR RY 2 t

'AgResearchFlock House Agricultural CentreBulls, New Zealand2Department of Animal ScienceMassey UniversityPalmerston North, New Zealand

Abstract Several biological measurements weremade on captive sambar deer over 3 years (198992). Calving occurred from January to November,with a peak in A pril/M ay. M ean calving date was 8May (SD = 71.3 days; n = 31). Calf mortality atbirth was 28 % (12/43), with the major causes be ingadult aggression and inclement weather. Mortalitywas higher in stag calves (41%) than hind calves(6%). Most adult stags were in velvet antler betweenJanuary and April (mean = 125 days; SD = 22.6days) , and in hard antler between May andNovember (mean = 231 days; SD = 40.0 days),during which time rutting behaviour w as observed.Mean dates of velvet stripping and hard antlercasting were 17 April (SD = 14.9 days) and 7December (SD = 35.4 days), respectively. Thesedata are compared with similar data for farmed reddeer. Sambar deer w ere found to be highly nervousand temperamental, but became more settled withregular human contact and feeding with maize andhay.Keywords sambar deer; Cervus unicolor; calvingpattern; antler cycle

"("Corresponding auth orA93047Received 10 July 1993; accepted 15 January 1994

INTRODUCTIONIn spite of a worldwide interest in deer farming,most deer species presently being farmed are oftemperate origin (red deer, Cervus elaphus; fallowdeer, Dama dama; wapiti/elk, Cervus elaphussubsp.). Australia, New Caledonia, and Mauritiusare currently developing systems for the farmingof tropical deer. The most extensive studiesconducted to date with tropical species have beenwith rusa deer (Cervus timorensis) (van Mourik1985; Chardonnet 1988) and axis/chital deer (Axisaxis) (Chapp ie 1989; My lrea 1992). There is a lackof biological data on samb ar deer (Cervus unicolor).The establishmen t of a small semi-domesticatedsambar herd in New Zealand provided theopportunity to observe both biology and behaviourand record aspects of relevance to deer farming.

MATERIALS AND METHODSLocationThe study was carried out at Flock HouseAgricultural Centre, Bulls, Manawatu, NewZealand, latitude 4014 'S and longitude 17516'E.Average annual rainfall is 875 mm with a dryperiod from January to March, and strong westerlywinds during October and November. The meanmonthly temperature ranges from 9 to 20C .AnimalsThe biology and behaviour of two groups of semi-domesticated sambar deer were observed fromNovember 1989 to December 1992. The animalscom prised 10 hinds and five stags transferred fromadjacent Turakina and Rongotea areas, andtransported to Flock House during 1989 and 1990,and 13 hinds and three stags imported from Victoria,Australia, in July 1990.Group A comprised two stags and six hindsfrom the Rongotea area which were set-stocked on0.85 ha of pasture, with access to 0.1 ha of openpampas (Cortaderia sp.). The remaining animals


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80 New Z ealand Journal of Agricultural Research, 1994, Vol. 37(6 stags and 17 hinds) from Turakina and Australia(Group B) were set-stocked on 2.4 ha of pasture,which included 0.1 ha of pine trees (Pinus radiata).The pampas and the pine trees functioned as naturalshelter, particularly against w esterly wind s. Pasturecomposition and availability have been describedby Semiadi e t a l . (1993). Hay and maizesupplements were fed as required during winter.Animals were not handled because of theiraggressive and temperamental nature. All matingoccurred naturally and hard antlers were never cut.In M ay 1992, adult and yearling stags from Grou psA and B were removed because of fightingproblems, leaving one adult stag per group.ObservationsWeekly observations of calving in Group Acommenced in November 1989, increasing to 3times weekly in January 1991. As the number ofcalvings increased, the frequency of observationwas increased to a daily basis. Four adult hinds ineach group could be identified individually.Twenty-four hours after birth, calves were ear-tagged, weighed, removed from their dams, andhand-reared.Fresh faecal samples were obtained from bothgroups on a monthly basis, by collecting 8-12pellets voided per animal from at least 70% of thetotal animals. These were examined for lungwormlarvae (Dictyocaulus viviparus) and gastrointestinalhelminth eggs (Strongylate sp.) by the Departmentof Veterinary Pathology, Massey University. Aveterinarian undertook post-mortem examinationson dead calves.Calculation of dataCalving dates and birth weights were tabulatedmonthly, over 3 years. Calving dates and birthweights for the 1991 season for animals importedfrom Australia were not included, based on theassumption that the period of quarantine/transportmight have altered the normal New Zealandbreeding pattern. However, calf sex and cause ofmortality of these animals were recorded and areincluded in the data presented. Date of antlercasting, period of velvet antler growth, and theperiod of time stags were in hard antler werecalculated from the antler data. Date of velvetstripping was recorded when one of the antlersshowed pronounced velvet stripping. Date of antlercasting was recorded as the average of the twosides. Antler length (length of beam) was measured

6 -c 5 "S 4"oS 33 2 -

1 - Jan FebMar A prMay Jun Jul Aug Sep Oct NovDec

Fig. 1 Monthly calving distribution during 1989-92of captive sambar deer under New Zealand conditions(winter solstice, 20 June; summer solstice, 20 December).

from the base of the coronet to the farthest tip ofthe beam, and beam circumference was measuredfrom c. 5 cm above the brow tine, using flexiblepolypropylene tape.RESULTSCalving pattern and calving intervalSambar births occurred almost all year round,except in July and December, with a peak duringApril and May (Fig. 1). Mean calving date was 8May (SD = 71.3 days; n = 31), with little variationbetween years. Mean calving interval from theidentified hinds was 329 days (SD = 29.7 days; n =6) and in each year all adult hinds gave birth.Birth weight, sex ratio, and mortalityBirth weights for male and female calves weresimilar, 8.1 kg (SD = 1.37; n = 17) and 7.8 kg (SD= 1.72 kg; n = 15), respectively. Pooled data fromlive and dead calves over 3 years showed that thesex ratio of male to female was 1.6:1.0 (n = 43).M ortality of calves within 24 h after b irth was 28%(n = 12/43), with a high proportion of deathsresulting from exposure (33.3%, n = 4) andaggressiveness of adult sambar deer towards thenewborn calves (33.3%; n = 4), but the level ofaccuracy for the causes of mortality may not behigh because of the low num bers of calves involved.Mortality of male and female calves was 41 % (n =11) and 6% (n = 1), respectively.


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Semiadi e t a l .Sambar dee r b io logyAntler dataMean hard ant le r cas t ing da te in adul t s tags was 7December , some 45 days ea r l i e r than young s tags(Table 1) . M ost s tags were in ve lve t ant le r be twe enJanuary and Apri l , ye t young s tags spent varyingpe r iods in ve lve t an t l e r ( r ange 102-155 days ) ,

81depen ding on w hen they were born . The ne t r e su l twas tha t there was a tendency for sambar s tags tobe synchronised in ha rd an t le r be tween May andNovember (Table 2) . For the sambar s tags in thissample , hard ant le r weight of the adul t s tags wastwice tha t of young s tags (Table 1) .

Table 1 Date of hard antler comm encement (mean; S D) in captive adult (> 3 years of age) andyoung (< 3 years of age) sambar deer stags, and length of time (days; SD) in velvet antler and hardantler, during 1990-92, under New Zealand conditions (n = number of observations).

Mean date of velvet stripping'Mean length of time in hard antler (days)Mean date of antler castingHard antler2:Length (cm)Circumference (cm)Weight (g)Mean length of time after antler castingto the first sign of velvet stripping (days) 3

Adult17 Apr (14.9)231 (40.0)7 Dec (35.4)

50.8 (6.92)11.9(1.48)817(297.6)125 (22.6)

n11888888

Young10Jun(84 .6)205(107.8)21 Jan (45.2)

39.5 (4.45)10.1 (1.00)402 (79.7)136(29.8)

n4336663

'Calculated from one sideindividual animals were not identified3Period of velvet antler growth

Table 2 Antler status from identified captive adult (> 3 years of age) and youn g (< 3 years of age) sambar deer inNew Zealand between 1990 and 1992.Stag no.Adult1

57

4

Young37A

96 5

Year199019911992199019911990199119921990199119921991199219901991199219911992

JVHVVVVVV

VVVVs

FVHVVVVVVVV

VVVVs

MHVVVVVVVVV

VVVVs

AHVHHHHVHVH

HHVVV

MHVHH- 'HHHHH

HVHHVV

JHHHHHH- 1HH

- 'VH- 1S

JHHHHHH

HH

VH

S

AHHHHHH

HH

VH

S

SHHHHHH

HH

SH

S

OHHHHHH

HH

SH

S

NHHHHHH

HVVsHS

DHHVVHVVVVVsHIZl

V = velvet antler, H = hard antler, S = hard spike;- 1 = no recording, stag removed from area.


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82BehaviourRegardless of origin, all sambar deer in this studywere very alert and nervous. Their disquiet wasexhibited by stamping of their front hooves withtails erect. They would walk in a step-by-stepmanner and then run with their heads lowered,la ter forming several small groups . Whenfrightened, the leader would bark, causing allmembers of the group to run spasmodically. In avery stressful situation, they would lie down afterrunning. Because of their alert behaviour, sambardeer would notice any slight change within theirpaddock and they would avoid an area ofdisturbance for some time. Despite their nervoustemperament, it was possible to quieten themthrough regular daily contact and feeding withmaize or hay (0.5 kg/head per day) during winter.Most active rutting behaviour in sambar stagswas observed between la te May and ear lyNovember. Rutting behaviour in sambar stags wasmarked by wallowing, threshing the ground withantlers, head-rubbing on posts and trees, andurinating on their head and antlers. Rutting sambarstags did not appear to produce strong body od ours,as commonly found in red stags, and roaring wasnever heard during daytime. Daytime observations(0900-1000 or 1500-1600 h) indicated thatalthough the dominant rutting sam bar stag collecteda harem, the dom inant stag displayed a high degreeof tolerance toward the presence of other stags inhard antler within the harem. On four occasionsthe dominant stag was seen chasing a rival stagfrom the vicinity of the harem. On four otheroccasions, both dominant and rival stags wereobserved to be mode-fighting (pushing each other),while the hinds were either grazing or lying nearby.Although there was an impression of harmonybetween hard antler stags, two deaths were recordedfrom injuries sustained during fighting. Antlercasting apparently did not change the behaviour ofa dominant stag in keeping the harem, providingthere were still hind(s) in oestrus. A dominant stagwas observed chasing a hind, both before and afterantler casting. However, dominant stags were notaggressive towards stags in velvet antler.

Health statusMonthly checks over 12 months revealed nolungworm larvae or strongylate eggs in faeces ofeither group. On one occasion a young h and-rearedsambar stag (5 months of age), grazing with y ounghand-reared red deer (7 months of age), became

New Zealand Journal of Agricultural Research, 1994, Vol. 37heavily infested with lung worm. The symptomswere heavy coughing, restlessness, refusal to eat,and loss of body weight (-9.4 kg within 27 days).There were 3350 lungworm larvae and 150 strongy-late eggs/g faeces. Treatmen t involved drenchingwith Ivermectin (1VOMEC, Merck, Sharp &Dohne, New Zealand) at 1/3 of the full dosage (5ml; 1000 |j.g) for 3 con secutive d ays, and a boo sterof full dosage (15 ml; 3000 ng) on Day 4. Threedeaths caused by malignant catarrhal fever (MCF)were recorded during the 3 years an ad ult hind inlate winter 1991; a yearling stag in late autumn1992; and a yearling hind in late winter 1992.

DISCUSSIONSambar deer demonstrated peak calving in autumn(April-May), with a very large spread (Fig. 1).Peak calving time of wild sambar in Nepal wasreported close to the monsoon season (June-July,local time), also with a very large spread (Mishra1982). This suggests that sambar deer in NewZealand ha ve retained their ancestral calving patternand, unlike temperate d eer, the reproductive patternof tropical deer is not strongly linked to day length.Several other studies support this view. Loudon &Curlewis (198 8) found chital stags in Great Britaindid not respond to melatonin treatment and calvedthroughout the year. Unmated chital hinds in NewSouth Wales, Australia, had continuous oestrouscycles throughout the year (Mylrea 1991). In NewZealand, calves born during late autumn and w intercould face inclement weather conditions and lowavailability of pasture. This is supported by thehigh calf mortality observed in the present study,and similar calf m ortality has been noted with rusaand chital deer in Australia (Mylrea 1991). Thishigh mortality may be responsible for the relativelack of success of sambar in the Manaw atu (Kelton1981). The greater neonatal mortality of sambarstag calves is surprising and it is interesting thatthere is a higher ratio of stag to hind calves born.The p ercentage of calf mo rtality as a result of adultdeer aggressivene ss in the present study was sim ilarto the finding in an earlier study of farmed red deer(Kelly & Drew 1977), 33 versus 27 % , respectively.It was suggested that lack of shelter and highstocking rate were the main cause of the mortalitiesin red deer. The wide spread of calving in sambarcoupled with the likelihood of adult aggressionmeans that single sire mating of sambar deer shouldbe practised and stags removed before calving.


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Semiadi et al.Sambar deer biology 83As the onset of rutting behaviour commencedin May/early June, and assuming the gestationperiod of sambar deer is 8 months (Schaller 1967;Bentley 1978), onset of calving should havecommenced in January. Calving did in factcomm ence at this time, and continued to Novem ber,with a peak in April/May. It would therefore seemthat some sambar stags were capable of mating inany month of the year. A study from Nep al (M ishra1982) indicates that peak mating activity of sam bardeer is between O ctober and Nov ember (local time),when the sambar stag population is in a transitionfrom a high number of stags in velvet antler(October) to a peak number of stags in hard antler(December). This suggests that mating by sambarstags in velvet antler is likely. O ther reports supportthe finding of tropical stags being fertile at any

stage of antler development (Goss 1983; English1992; G. W. Asher pers. comm.). Mylrea (1991)also noted that chital stags in velvet antler couldproduce fertile spermatozoa and m ount an oestroushind. Moreover, sambar stag mating behaviour wasobserved during early velvet antler growth in thepresent study.In contrast, red stags are not fertile while invelvet antler (Wilson 1984), and Clutton-Brock &Albon (1989) showed that although wild red stagsare in hard antler for more than 7 months, the

active breeding season lasted only 1.5 months.This suggests that compared to temperate deer,tropical stags are capable of mating over a much

longer period. Continuous oestrous cycles and theabil i ty of t ropical stags to produce fert i lespermatozoa and to mount hinds at any stage ofantler development would explain such a widespread of calving.Calving and antler data in adult sambar and reddeer are compared in Table 3. Both sam bar and redstags appear to carry their velvet and hard antlerfor similar periods of time, but under New Zealandconditions both antler casting and velvet strippingoccurred 10-12 weeks later in sambar than in redstags. There has been historical debate on the lengthof time that sambar stags in New Zealand carryantlers. However, annual hard antler castingobserved in this study supports the observations ofRudd (1978) for wild New Zealand sambar stags,and the findings of Acharjyo (1983) in India that

sambar stags lost their antlers each year.Sambar deer are known as shy and cunninganimals (Harris 1966). In their natural environment,sambar live in dense habitat, w here natural predatorsare present at all times (K itchener 1961 ;Rice 1986).However, an alert and cautious behaviour has beenretained after 130 years in New Zealan d in spite ofan absence of natural predators, other than human s.Despite their nervous temperamen t, sambar becomequieter under farmed conditions through regularhuman contact and feeding with maize or hay.Malignant catarrhal fever was diagnosed as thecause of death in 3 cases and several other authorshave ind icated that tropical deer may be susceptible

Table 3 Some comparisons of the biology of antler growth and reproduction in captive adult sambar deer and reddeer in New Zealand.

Antler growthMean time of hard antlercommencement (SD)Mean days in hard antler (SD)

Mean hard antler casting date (SD)Mean days in velvet antler (SD)ReproductionMean calving date (SD)Gestation period (days)Birth weight (kg)Oestrous cycle length (days)

Sambar deer17 A pr(14.9)23 1(40.0)7 Dec(35.4)125(22.6)8 May(71.3)24 07.8-8.117

AuthorPresent studyPresent studyPresent studyPresent study

Present studyBentley (1978)Present studyEnglish (1988)1

Red deer9 F e b(6.1)24 92 5 S e p120

8 Dec(12)23 36.2-9.618.2

AuthorFennessy & Mackintosh (1992)Fennessy & Mackintosh (1992)M u i r & S y k e s ( 1 9 8 8 )Fennessy & Suttie (1985)

Moore etal. (1988)Kelly & Moore (1977)Moore etal. (1988)Kelly & Moore (1977)

Australia


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84 New Zealand Journal of Agricultural Research, 1994, Vol. 37to MCF (Soraja et al 1987; G. W. Asher pers.comm.). Although high resistance to internalparasites has been reported in rasa deer (Chardonnet1988; Woodford 1991), the heavy lungworm burdenin a young stag suggests that young sambar deermay be susceptible when exposed to a source ofinfection. Normal worm burden levels in NewZealand deer farming are 0-72 lungworm larvae/gfaeces (Wilson 1985), compared to 3350 lungwormlarvae/g faeces in the infected sambar grazing incontact with red deer. Otherwise, no significanthealth problems were evident with set-stockedsambar deer during the 3 years of the present studywhere they were not in contact with other deerspecies.

The pattern of red deer feed requirements(calving in late spring/early summer) generallydoes not fit with the spring peak of pastureproduction in New Zealand. Advancing the calvingseason of red deer through hybridisation with otherdeer species has been suggested as a means ofbetter matching feed supply with feed demand.Hybridising red deer with Pere David's deer(Elaphurus davidianus) or rasa deer has beensuggested as a means of advancing the calvingseason (Tate et al. 1992). Present observationsindicate that sambar deer, on average, calve 7months earlier than red deer in New Zealand andthe gestation period is similar for sambar and reddeer (Table 3). The production of a fertile hybridsambar x red deer could thus be valuable to theNew Zealand deer industry.

ACKNOWLEDGMENTSLawrence and Pat Rowe and Clive Jermy are thankedfor providing sambar deer for research. We thank PeterBurke, Manager, Flock House Deer Unit, for his helpwith these studies. Massey University thanks AgR esearch(formerly MAF Tech) for the provision of financialassistance. G. Semiadi received sch olarship support fromthe Government of Indonesia and from the New ZealandMinistry ofForeign Affairs and Trade.

REFERENCESAcharjyo, L. N. 1983: Observations onaspects of antlercasting in captive sambar deer. Pp. 23-28 in:Antler development in Cervidae, Brown, R. D.ed . Caesar Kleberg Wildlife Research Institute.Kingsville, Te xas.Bentley, A. 1978: An introduction to the deer of

Australiawith special reference to Victoria.The Koetoeng Trust, Melbourne.

Chardonnet, P. 1988: Etude de factibilite technique eteconomique De L'elevage D e'Cerfs en Nouvelle-Caledonie. IEMVT, Cedex. France.Chappie, R. S. 1989: The biology and behaviour ofchital, Axis axis deer in captivity. Unpubl. PhD

thesis, University of Sydney, Sydney, Australia.Clutton-Brock, T. H.; Albon, S. D. 1989: Red deer inthe high lands. London, BSP Professional Books.English, A. W. 1988: Diseases of deer. Vade Mecumseries A no. 11. Post Graduate Foundation inVeterinary Science. University of Sydney,Sydney, A ustralia.English, A. W. 1992: Managem ent strategies for farmedchital deer. Pp. 189 -196 in: The biology of deer,Brown, R. D. ed. New York, Springer-VerlagPublications.Fennessy, P. F,; Suttie J.M. 1985: Antler growth:nutrition and endocrine factors. Pp. 239 -25 0 in :Biology of deer production, Fennessy, P. F.;Drew, K. R. ed. The Royal Society of NewZealand bulletin 22.Fennessy, P . F.; Mack intosh, C. G. 1992: Hybridisationof red deer and Pere David's deer. Pp. 181-186in: Proceedings of a Deer Course forVeterinarians no. 9, Wilson, P. R., ed.NewZealand Veterinary Association Deer Branch.Goss, R. J. 1983: Deer antlers. Regeneration, functionand evolution. London, Academic Press.Harris, L. H. 1966: Hunting sam bar deer. New ZealandForest Service, Wellington.Kelton, S. D. 1981: Biology of sambar deer in NewZealand, with particular reference to diet in aManawatu flax swamp. Unpubl. MSc thesis,Massey University, Palmerston North, NewZealand.Kelly, R. W.; Drew, K. R. 1977. The behaviour andgrowth of deer on improved pastures. Pp. 20-25

in: Deer farming in New Zealand: progress andprospects, Drew, K. R.; McDonald, M. F. ed.Wellington, Editorial Services Limited.Kelly, R. W.; Moore , G. H. 1977: Reproduct iveperformance in farmed red deer. New Zealandagricultural science 11: 179-181.Kitchener, H. J. 1961: The sambar deer Cervusunicolor equinus. Malaya n n ature journal 15:52-61 .Loudon, A. S. I.; Curlewis, J. D. 1988: Cycles of antlerand testicular growth in a seasonal tropical deer

(Axis axis). Journal of reproduction and fertility83 : 729-738.


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Semiadi et al.Sambar deer biology 85Mishra, H. R. 1982: The ecology and behaviour ofChital (Axis axis) in the Royal Chitawan N ationalPark, Nepal, with comparative studies of hogdeer {Axis porcinus), sambar (Cervus unicolor)and barking deer {Muntiacus mu ntjak). Unpubl.PhD thesis, University of Edinburgh, Edinburgh,

United Kingdom.Moore, G. H.; Littlejohn, R. P.; Cowie, G. M. 1988:Liveweights, growth rates and mortality offarmed red deer at Invermay. New Zealandjournal of agricultural research 31: 293-300 .Muir, P. D.; Sykes, A. R. 1988: Effect of winter nutritionon antler development in red deer (Cervuselaphus):a field study. New Zealand journal ofagricultural research 31: 145-150.Mylrea, G. E. 1991 : Reproduction in tropical species.Pp. 249-261 in: Proceedings of a Deer Course

for Veterinarians no. 8. Wilson, P. R. ed . NewZealand Veterinary Association Deer Branch.Mylrea, G. E. 1992: Natural and artificial breeding offarmed chital deer (Axis axis) in Australia.Unpubl. PhD thesis , University of Sydney,Sydney, Australia.Rice, C. G 1986: Observations on predators and preys atEravikulam National Park, India. Journal o f theBombay National History Society 83: 283-305 .In : Biological abstracts 1987 84: A B - 2 4 5 .Rudd, J. 1978: The antler growth pattern in the Bay of

Plenty Sambar herd. New Zealand wildlife 7:33-35 .Schaller, G. B. 1967: The deer and the tiger. Chicago,The University of Chicago Press.

Semiadi, G.; Muir, P. D.; Barry, T. N.; Veltman, C. J.;Hodgson, J. 1993: Grazing patterns of sambardeer (Cervus unicolor) and red deer (Cervuselaphus) in captivity. New Zealand journal ofagricultural research 36: 253-260 .Soraja, S.; Logathan, P.; Indrasegaran, S.; Gon, Ch.;Sheik-Omar, A. R. 1987: An outbreak ofmalignant catarrhal fever in Javan deer (Cervustimorensis). Kajian veterinar 19: 179-183.Tate, M. L.; Manley, H. C ; E merson, B. C ; Fennessy,P. F. 1992: Interspecies hybrids of deera ru m i-nant resource for gene mapping and quantitativetrait studies. Proceedings of the New ZealandSociety of Animal Production 52: 137-142.van Mourik, S. C. 1985: Behaviour and physiology offarmed rusa deer (Cervus rusa timorensis).Unpubl. PhD thesis, University of Melbourne,Victoria, Australia.Wilson, P. R. 1984: Antler growth and control. Pp. 3 1 -35 in: Deer Refresher Course. Proceedings no.72 . Post Graduate Foundation in VeterinarySc ience , Unive r s i ty of Sydney , Sydney ,Australia.Wilson P. R. 1985: Disease prevention strategies:Experience on a small deer unit. Pp. 83-86 in :Biology of deer production, Fennessy, P. F.;Drew, K. R. ed. The Royal Society of NewZealand bulletin 22.Wo odford K. B. 199 1: Repro duct ive cyc les andperformance of rusa deer in the tropics andsubtropics. Pp. 262-268 in: Proceedings of aDeer C ourse for Veterinarians no. 8, Wilson,P. R. ed . New Zealand Veterinary AssociationDeer Branch.

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