Antipredatoraspectsoffallowdeerbebaviour
duringcalvingseasonatDoñanaNationalPark(Spain)
C. SANJOSÉ and F. BRAZA
Estación Bioldgica deDotiana,CSICApdo1056,41080Sevilla,Spain
Received 26 June 1990,accepted 1 March1991
Thisstudyevaluatessomemorphological,ecologicalandbehavioural aspectsof fallow deer mothers and fawns during the calving season in order to know the antipredatortactics selected in this specieslivinginaparticularly open hahitat at DoñanaNationalPark.Inthisarea(allowdeergivebirthinsynchronyduringthefirst fortnight of June. Atthistimetheymoveawayfrom their matriarchalgroups. A concentration of births in the early afternoon wasdetected,coinciding with the minimalactivityofpredatorsinthearea.Mothersgroomtheiroffspringandingestthe birthremainstopreventtheattractionofpredators. Duringthefirstdaysoflife fawns remain hidden keeping motionless.
KEY WORDS: Damadama, antipredator strategy, carving time, maternal behaviour.
Introduction139
Materials andmethods140
Resulrs143
Discussion145
Acknowledgements146
References147
iNTRODUCTION
The fawns in many species of ungulates are specially liable to suffer from predationduring their first daysoflife. However,only afewworks exist about the mechanisms ofdeveloping anantipredatorstrategy atcalving time, probably due to the difficulty toobserving mothersand their young at birthand during the days immediately after birth.
Mothers and newborn young in those species use a ehiden> strategy (elk,
ALTMANN 1932; fallow deer, GILBERT 1968; Coke’s bartebeest, GosLiNG 1969;
bontebok,DAvID1973, 1975; pronghorn,KITCHEN 1974; reddeer, GIJINNESSet al.
1978; red buck, IRBY 1979). The species of these groups are basically classified as
*cbideroor((followers>dependingonwhetherthe newborn lieconcealed for their first few claysor actively follow their mothers(LENT 1974, RAI.i.s et al. 1986). While
<following> hasbeenviewedasastrategyforavoidingpredatorsinopenhabitats, hidingaisthoughttoreducepredationriskinclosedhabitats(LENT1974,ESTESESTES 1979).
Astothebehaviour offallowdeerduringthecalvingseason,onthebasisofthe
studieswehaveconductedtodate(ALvA.aEzeta!.1975,BitAzAetal. 1988,SANJose
1988,BItizASANJose 1989),wemayclassifyfallow deerinprincipleasehider>’
since the young remain hidden the first days after birth and are only visited
periodicallybytheirmothers,mainlytofeed.Nevertheless,morphological,ecological
andbehavioural aspects,whichdecidethisassignment,havenotbeenevaluated quantitativelyuptonow.Theaimofourstudywastoevaluatesomeoftheseaspects inmothersandfawnsduringthe calvingseasonandtodiscusstheresultsinrelationto theantipredatorstrategiesdevelopedbythisspeciesandthephysicalcharacteristics of thehabitat.
MATERiALS AND METHODS
Observations includedinthispaperwerecarriedoutatDoñana NationalParkfrom1982to
1987.InthisNationalParkof73,000 ha(Fig. I)situated atthemouthoftheriverGuadatquivir,
the wildfallowdeer population occupies anecotone zone whichcorresponds to the transition between thexerophyric schrub and the marshland (BRAZA 1975). In this ecorone wedistinguish between thefollowingbiotopes(according toALLIEReral.1974, andAGuIr.AR AMAT etal.1979)
(Fig.1):
a) marshes:characterized by astrong seasonal dependence and made upof species like
Salicorniaramosissima,Arthrocnemumsp. and Scispusmaritimus;
bI rushareas:formamosaicwits pasturesatthemarshlandborders andpresentadominant vegetation of macus marit&nus;
c) wetmeadows:areasofpastures whichareoccasionallyfloodedandbasicallycharacterized
by the association of Amzena gaditanaand 44sphodelusaestivus;
d) dry meadows:havealowerlevelofthesubterraneanwater and presentacharacteristic association of the speciesUrinea maritima and Anthemis cotula;
e)bracken areas: areas with a dense plant cover where the species Pseridiumaquilinum
dominates;
f)schrubland:ismadeupofvegetationwhich colonizedstabilizedsandsandisbasically characterized byspecieslikeHalirniumhalimifolium,Cistuslibanotis,Ericascoparia,Callunavulgaris andStauracanthusgenistoides,among whichsome isolated cork oaks (Quercussuber)are found.
Atpresent, theIberian lynx(Lynxpardina)istheonlypredator abletokillhalf-growndeerin Doñanaand, asweknow, thelynxhasvery littleinfluenceon theearlymortaliry offallowdeer fawns(BELTRAN et al. 1985). Of other potential predators of fallowdeer calves (fox, genet, ichneumon,wildboar, Imperialeagle)rheanalysesdonotrevealsignificant deer presenceintheir diet (VENERO 1982, PAL0MARE5 1986, RAu 1988, FERRER 1989).
During4years(1983-1986), fromthe25thofMaytothe20thofJune weconductedacensus bycaratleastonceaweekafractionoftheecotoneareawhichmeasured5.5kmlongand1.5km wide(Fig.1).
Generallytwoobserversrecordthenumberofsolitaryfemales(animalscharwere arleast50 awayfromanyotherfallowdeer).Theproportion ofsolitaryfemaleswascalculatedwithrespectro the rotal number of females present in the area the same day.
Themeanpopulationsizeduring theperiod1983-1986was264.44(*7.89)deerwirhamean totalnumberoffemalesof123.92(±6.45).Themeancalf-ratiowas60.37(±2.05)(from BRAZAet al. 1990).
The rutting season takes placeduring the first fortnight of October (BRAZA er al.1986).
intipredator behaviourin fallow deer141
riArea ofCensus
Fig.1.
DoñanaNationalParkandstudyarea(above).Themainbiotopesconsideredinthe studyarc
roportionol theecoronezone(below).
Ct
0
U)
0
30-
20
10
L
I PI
25515255
MayJuneJuly
198a
1984
1985
1986
Fig.2
Variationsin the percentageof solitary females
Femalesgivebirth toasinglefawnattheendofspring(BRAZA eral.1988),afteragestation period o236days (SAN Jos1988),
Duringthecalvingseason,intheareaofcensuswesearcheddailytolocalizenewbornfawns.
This wasdone on foot and sometimes counting on the assistance of a vehicle or a guard on
Antipredator behaviourin fallowdeer 143
horseback. When wefoundafawnwerecorded alltheenvironmental characteristics oftheplace whereitwashiddenaswellasthebehaviourshownbythemother(if nearby)andthe fawnatour approach.Oncecaptured thefawnwasmarkedandmeasured (BRAZA etaI.1988, SAN josI etal.
1989).
When itwasnotpossibletodetect themomentofbirth weusedvariouscriteria toestimate thecondition vnewbornn,e.g. the presenceofremainsofthe placenta, thestillmoistfur ofthe fawn, the umbilicalcord’s degree of healing, the hardening of thehoovesand the presence of cartilageontheirpoints.Thesecriteriahavefrequently beenusedtodetermine theageofnewborn fawnsof other deer species (i.e. HAUGEN SnAKE 1958).
Finally,inordertomakemoredetailedobservations,wefollowedina‘mainobservationarea’ (aprox.150ha)themovementsofthepregnant females.Thiscontrol wascarriedoutbymeansofa binoculartelescope(20(40)x(80(500)froma30mhighobservation towercontinuallyfromdawnto nightfall.
Weusednon parametric statistics (SIEGEL 1956)toevaluate thelevelofsignificanceofthe
results.
RESULTS
Asudden increase intheproportionofsolitaryfemaleswas observedeach year at thebeginning ofJune (Fig.2).Thedistributions ofsolitary femalesindifferentyears arecoincident(83X84,84X85, 85X86, t=1; P<0.01; KendalLCoefficienceof Concordance).Only108 birth dates ofthe190fawnscapturedcould beknown with precision. The peak of solitary females coincides with the average date of birth estimated.Withtheexceptionof1982,withameandateofbirththe5thofJune,for the rest ofthe years (1983-1987)most of birthsoccurredwithin the3 first days of June: 1=1(±3.7),isiS(±3.8),*=3(±2.2),Yc=2 (±3.8)and i=2.5(±4.3)
respectively.
In41casesitwaspossible torecord exactly the timeofbirth.From thesecases wecouldcalculatetheaveragehourwhich is14:16(±3.16). Mostbirths(x =25.54; P’cO.OOi) were detectedbetween 15:00 hr and 17:00 hr (Universal time) (Fig.3).
Inallthebirthsobserved intheir entirity(n=20),themother turns towardsthe
fawn, whichTieson thegroundimmediatelyafter birth,andstartstolickitactively,
removing andingesting theamnioticmembraneswhich stillcover the young. Finally,
D
0
C?
cy
CC
C C)
C.)
a)
3—Distributionofbirthsovertheday’ight hourt(Universal time)
rush marsh
dry meadow
‘wetmeadow
Eischrub bracken
Fig. 4.Habitat preferenceshown by femaLesfor hiding their fawn (n=190).
thefemale ingests the newborn’s faeces. The newborn fawn hasadark brown coat with white speckles on its back, the belly being entirely white. Its colour gets progressively lighter in the course of the 1st week of life.
During the first days oflife the fawn lies hidden.Its motherlies down some metresofforgoesawayto joinagroup ortograzealone. The biotope ofthehidding
place was not selected randomly (xi =511.45;P<0.001). Most of the fawns
captured were found in a rush area Cx? =467.46; P<O.0O1)(Fig.4). The fawn
remains motionless and adopts the typical flat-on-the-groundposture,frequently
curled up totally concealingits extremitiesand head, althoughthe tendencyto run awaywhen aperson isapproachingrapidly iscorrelatedwith the fawn’s age(r I; P<0.001)(Fig. 5),increasing and surpassingaprobabilityof 50%from the 3rdday of life.
01190fawnscaptured,for89itwaspossibletoobserve themother’sbehaviour when weapproachedthe fawn. In 76% ofallcasesthe female went awayfrom the placewheretheyoungwashidden.In28casesthefemalestayed neartheplacewhere the young was being marked at an average distanceof 72.5 m (±45.2).
/no flight
Itighi
lawn age (days)
Eig. 5.TemporaL variation in the fawn’s reaction at the approach of a person on foot
Antipredator behaviourin fallow deer 145
When analysing the fawns’ and mothers’ behaviour in the main biotopes (marshes, rushareasandmeadows) thetendencywassimilar; so,theflight probability byfawns younger than4dayswassignificantlylow(*=0.12;SD=0.12;x 5.44;
P<0.05,yJ=14.74;P<O.O1and Ic =12;Pc0.01 repectively);themother’s flight
tendency washigh in the three biotopes(*=0.74;SD=0.36) but due to the small sample itwas significant only for rush areas (x? 11;P<o.01).
Finally, countingon 13 cases of fawns ohserveddaily, the average date of integrationinto thegroup wascalculated.The criterionbeing the ageof the young when seen in agroup for the second time. This value wasof 13.61days (±4.31).
DISCUSSION
Most offallow deer birthstake place in Doñanaduringthe first fortnightof June. Inthischaracteristicthe population coincideswith mostdeer populationsin temperateareas presentinga reproductiveperiod which is markedly seasonal and
extraordinarily synchronized (Vos 1960, MITCHELL LINCOLN 1973, BERGERUD
1975, CLUTTON-BROCK GUINNESS 1975).
Two general hypothesis are invoked to explain birth synchrony: (i) optimal
timing with respect toseason mayenhancethe survival and growth ofoffspring as wellas the survival and future reproductivesuccess of the mother (DAUPHINE MCCLURE 1974, BUNNELL1980,DUNBAR 1980, CLUTTON-BROCICet al. 1982, RUT- BERG 1987) and(ii) females maysynchronizebirthsinorder toreduce predationon newborns either bysatiatingor confusing predators(DAUPHINEMCCLURE 1974, EsTEs ESTES 1979, LOTT1981, BERGERUD et a!. 1984).
Withrespect topredators,asalready mentionedintheIntroduction,at present therearenolargepredatorsintheareaand there isonlythelynx(Lynx pardina)tobe mentioned,whichattacks youngindividualsinautumnandwinter,affectingmortality rates toaminimum degree (DELIBES1980, BELTRANet al.1985). However,until the forties there were wolves (Canis lupus) living in Doñana; so, predation was an important force selecting for breedingsynchrony in fallow deer.
Furthermore,thecycleofforageavailability,inanarealikeDonanawithavery short spring andaverydrysummer, isalsoprobably animportantfactor selectingthe time of the calving season. The fact that none of the offspringof late breeding survived the summer seems toconfirmthis hypothesis.Asinother ungulatespecies (i.e.FESTA-BIANCHET1988)inadequatenutritionis suggested asthecauseofmortality of late newborn calves.
At Doñana a peak of births has been detected coincidingwith the period of minimaldiurnal activity ofpredatorsinthearea(DELIBES1980, ALVAREZetal.1983, PALOMARES 1986,BELTRAN 1988,RAU 1988). Asimilardiurnal birth distributionhas been described for other fallow deer populations (CHAPMAN CHAPMAN 1975, STERBAKLOSAK 1984)andforotherungulatespecies(E5TEs1966,GOSLING 1969). ESTES(1966)suggests that the morning peak allowsthe calves togaincoordination before night fall, when predatorsare more active, but, since wehave nodata from nocturnalobservations, in our case it is not possible hypothesize.
The rapid removal ofthe placenta byplacentophagiaalsolessens the possibility that predatorswill be attracted to the slowly-developing calf. In some species a diurnal distributionofbreedinghasbeen interpretedasapossible thermalregulation
(Len1981) but we think that this could be less adaptativein Doñanawhere the mean minimal temperature itt May-June never falls bellow 10 OC in June (i=z12.51*1.72°C during the period 1983-1986).
The efficiency of *clying-outnasan antipredator behaviourisincreased by the initial cleaning of thefawn and by the motherconsuminghis faeces. These factors reduce thechance of predatordetectingacalfbyitssmell. This behaviourhasbeen reportedfor many ungulatespecies (GOSLING 1969, ESPMARK 1971,JACKSONet al.
1972, KITCHEN 1974, AUTENRIETH FICHTER 1975, GUBERNIK 1980, TRILLMICH
1981, SADLEIR 1984).
On thebasisofbehaviouralaspectsofantipredatortactics offallowdeer during thecalvingseason, thesespeciescould beclassified as<hidersince the newborn lie concealed for their first fewdaysoflife. Accompaniedby theconcealmentofyoung calves, the isolation of the female reduces the chance of predation by making detectionbypredatorsmoredifficult.Thisbehaviourisusuallyshown byspecies that calveonshrublandswhichvegetationprovidesagood concealment(ALTMANN 1963, HAWKINSKLIMSTRA1970,JACKSoNet al. 1972,Wmmet al.1972,HEIDEMANN
1973, MEIER1973,CHAPMANCHAPMAN1975,CLUTTON-BROCKGUINNESS1975, NELSONMEal1981,OZOGAVERME1986). Femalesofspeciessuchaswildebeest that calve on short-grassareas which offer less conceal, do not seek isolation for parturitionand thereisnolying-out behaviour(ESTES1966). InDoñana,fallowdeer have adapteda<hiden> strategy toanopen habitat,selecting the rush areas asthe biotope that offers thehighest cover, and then displaying atypical thiders.behaviour asshown byother fallowdeer livinginforests(GanERT 1968,CHAPMAN CHAPMAN
1973).
On theotherhand, thecryptic colour ofthefallowdeer newborn calvesand the particular position that they adopt lying-out could be interpreted as a complete strategy ofcamouflage (TINBERGENet al.1967) and permits an understandingofthe motionless behaviour displayed in the open biotopes(marshes and meadows).
Furthermore,thesuccessofthe<hider,strategy inungulatesdepends inpart on the mother’s habitity to minimize the informationshe transmitsabout her young’s hiding place while remainingclose enough to distract or drive away a predator (ALTMANN 1963, LANGMAN 1977,MACCONNELL-YOUNT SMITH1978,PRATT ANDERSON1979,TRUE’rr 1979, STEIGERSFLINDERS 1981, BYERSBYERS 1983). At theapproach ofapotentialpredator(aperson on foot) most fallowdeer mothers run awayfrom the place where the young ishidden,attractingthe attentionofthe predator by their flight.
Finally, the effectivenessof lying-out in avery youngcalf dependson aflight distance ofalmostzero(TINBERGENet al,1967,GOSLKNG1969). Apredatormaythen passclosebywithoutthecalfrevealing itspresence byflight. Incontrast,afewdays later, fallowdeercalvesjumpand runawaywhen approachedtowithin afewmetres. This isprobably correlatedwithimprovedchance ofescapefrom predatorsby flight.
ACKNOWLEDGEMENTS
ThisstudywassupportedbytheComislén IoterininisterialdeCienciasy TecnologIa(C1CYT)
and theJunta de Andalucia.
REFERENCES
AGIJILARAMATJ.,MONTESC.,RAMIREZ L.TORRESA.1979.Pargue NacionatdeDoñana. Mapa
ecoidgico.Madrid:Instituto para Ia Conservacio’n deLa Naturaleza.
ALLIERC.,GONZALEZBERNALDEZF.RAMIREZL.1974.Donana. Mapaecologico.Sevill.a:Estacidn
Biolc5gica de Doiiana(ConsejoSuperiordeInvestigaciones CientIficas).
ALTMANNM.1952. Socialbehaviorofelk,Cervuscanadensisnelsoni,intheJacksonHole areaof
Wyoming. Behaviour4: 111-143.
ALTMANNM. 1963. Naturalistic studies of maternal care in moose and elk, pp. 233-253. In: Rheinfold ML.,Edit. Maternal behavior in mammals. London: Rheinfold H.L.
ALVAREZF.)BEAm F., AZCARATET., AGUILERA E. MAaTINR.1983. Circadian rhythms ina
vertebrate community of Doñana National Park. Antis XV CongresoInternacionalFauna
Cinegdtica Silvestre15: 379-387.
ALVAREZF.,BRA7.A F.NORZAGARAY A.1975.Etogramacuantificadodclgame(Damadama)en
libertad. DoilanaAcres Vertebrata2: 93-142.
AUTENRIETSI R.E. &.FJCHTERE. 1975. On the behavior and socialization of pronghorn fawns.
Wildlife Monographs42: 1-11.
BELTRAN j.F.1988. Ecologia yconducta espacio-temporaldcl lince ibérico (Lynx pardina)enci
Parque Nacionalde Doñana. Tesis Doctora4tlnivenidad deSevilla.
BELTRANJ.F., SANJosiiC., DELIBESM.BRAZA F.1985. AnanalysisofIberian lynxpredation
upon fallowdeerin the CotoDoñana,SWSpain.Proceedings oftheXVIIInternationalCongress of Game Biologists17: 173-181.
BERGERUnA.T. 1975. The reproductive season of Newfoundland caribou. CanadianJournalof
Zoology53: 1213-1221.
BERGERUn A.T.,BUTLER 11.E. MILLERDR. 1984.Antipredator tacticsof calvingcaribou:
dispersion in mountains. CanadianJournalof Zoology62: 1566-1575.
BRAZA F. 1975. Censodel gamo Dama dama)en DoEana. NaturaliaHispdnica3:1-27.
BRAZAF.,GARCIAJ.E.ALVAREZF.1986.Rutting behaviouroffallowdeer.ActaTheriologica31:
467-478.
BRAnF.SANJosC. 1989. An analysisof mother-young behaviour of fallowdeer during lactation period. BehaviouralProcesses 17: 93-106.
BRAZAF.,SANJostC.BLosaA.1988.Birth measurements, parturirion datesand progenysex
ratio of Damadamain Doñana. Journalof Mammalogy69: 607-610.
BRAZA F.,SANJoshC.,BLOM A.,CASESV.GARCIAj.E.1990.Populationparametersof fallow
deer at DoEana National Park (SW Spain). Acta Theriologica35:279-290.
BUNNELLT.L.1980.FactorscontrollinglambingperiodofDali’ssheep.Canadian JournalofZoology
58: 1027-1031.
BIERS J.A.BIERS K.Z. 1983. Do pronghorn morhers reveal the locations of their fawns?
BehavioralEcologyand Sociobiology13:147-156.
CHAPMAN D.CHAPMANN N. 1975.Fallow deer (Damadama). Lavenham,Suffolk; DaltonT.
CLUnON-BROCK T.H. GUINNESS F,E. 1975.Behaviourof red deer(Cervuselaphus)atcalving
Lime. Behaviour55:286-300.
CLunoN-BR0CKT.H.,GUINNESSF.E.ALBONS.D.1982.Reddeer:behaviourandecologyoftwo
sexes. Edinbuh: EdinburghVnivenity Press.
DAUPHINET.C.MCCLURE R.L.1974. Synchronous matinginCanadian barren-ground caribou.
Journalof Wildlife Management38: 54-66.
DAwnJ.H.M.1973.Thebehaviourofbontebok,Damaliscusdorcasdorcas(Pallas1766),withspecial
reference to territorial behaviour. Zeitschrift fürTierpsychologie33: 38-107.
DAvmJ.H.M.1975.Observations on matingbehaviour,patturition,sucklingand themother-young bondin thebontebok (Damaliscusdorcasdorcas).JournalofZoology,London177:203-223. DELIBESM.1980. Feedingecology ofrheSpanishlynxintheCotoDoftana.ActaTheriologica25:
309-324.
DUNBAR R.J.M. 1980. Demographicsandlife-history variablesofapopulation ofgeladababoons
Theropithecus gelada.Journalof Animal Ecology49: 485-506.
148C. SanJoséand F. Braza
ESI’MkRKY. 1971. Mother-young relationship and ontogeny of behaviour in reindeer (Rangifer tarandusL.).ZeitschrifI fürTierpsychologie29:42-81.
ESTESRD.1966. Behaviour andlifehistory of the wildebeesr(ConnochaetestaurinusBurchell).
Nature212:999-1000.
ESTESRD.& E5TF,sR.K.1979.Thebirthandsurvivalofwildebeestcalves.Zeitschrift für
Tierpsychologie50: 45-95.
FERRER M. 1989. Eláguilaimperial. Basebibliograficadeespecicsamenazadas.Sevilla:Juntade
1ndalucia, AgenciadelMedio Ambiente.
FEST&-BI&NCHET M.1988. Birthdate and survival in bighorn lambs (Oviscanadensis).Journalof
Zoology,London214:653-661.
GILBERTB.K.1968.Development ofsocialbehaviorinthe fallowdeer(Damadama).Zeitschriftfür
Tierpsychologie 25:867-876.
GOSLING L.M.1969.Parturitionandrelated behaviour incoke’sharrebeest (Alcelaphusbuselaphus
co/eelGunther).JournalofReproductionandFertility6:265-286.
GUBERNIKD.J. 1980. Maternal imprinting or maternal dabellingmingoats?AnimalBehaviour28:
124-129.
GuINNESSFE., GIBSONR.M. CLUTTON-BROCK Ti-I.1978. Calving time of reddeer(Cerr’us
elaphus)on Rhum. Journalof Zoology, London 185: 105-114.
FIAUGEN AD.SPEATCEW.D. 1958. Determining ageofyoungfawnwhite-railerdeer.Journalof
Wildlife Management22: 319-321.
HAwKINSRE.KLIMSTRA W.D.1970.Apreliminarystudyofthesocialorganizationofwhite-
tailed deer. Journalof Wildlife Management34:407-419.
I-IEI0EMANNG. 1973.Zur Biologiedes Damwildes(CervusdamaL1758). Hambmg &Berlin:P.
Parey.
itaY L.R,1979.Reproductioninmountainred-buckReduncafulvorufala.Mammalia43:191-213.
ACKSONR.M., WHITEM.KNOWLTON F-F. 1972. Activiry patterns ofyoungwhite-tailed deer
lawnsin SouthTexas. Ecology53:262-270.
(ITCHEND.W. 1974.Socialbehavior andecologyofthepronghorn.WildlifeMonographs38: 1-96.
ANGMANV.A.1977.Cow-callrelationships ingiraffe(Giraffacamelopardalisgiraf-fa).Zeitschriftfür
Tie,psychologie43:264-286.
£NTP.C.1974.Mother-infantrelationshipsinungulates,pp. 14-55.In:GeistV.Walther P.R.,
Edits.Thebehaviourofungulatesanditsrelation ro management. Moges, Switzeland: InternationalUnion forConservationof Nature.
OTTD.F. 1981.Sexual behavior and intersexualstrategies in Americanbison. Zeitschrift für
Tieqsychologie56:97-114.
IACCONNELI.-VOUNTE.SMITHC.1978. Muledeer-coyote interactionsin northcentral Color
ado. Journalof Mammalogy59: 422-423.
IEIF.R E.VON 1973. BeitragezurgeburtdesDamwildes(CervusdamaL.).ZeitschriftfürSaugetier
kunde 38: 348-373.
IITCHELB.LINCOLNG.A.1973.Conceptiondatesinrelation toagecondition intwopopulations
o1red deer in Scotland. Journalof Zoology, London 171: 141-152.
IELSON M.E.ME.CII L.D. 1981.Deersocialization andwolfpredation inNorrheasternMinne
sota. Wildlife Monographs77:1-47.
‘ZOGAJJ.VERMEL.J.1986. Relationofmaternal agetofawn-rearing successin white-tailed
deer. JournalofWildlife Management50:480-486.
SLOMARESF.1986.EcologiadeIaginetaydelmeloncilloenelParque NacionaldeDoñana.Tesis de Licenciatura,UniversidaddeGranada.
{ATTD.M.ANDERSON Vii.1979.Giraffecow-calfrelationshipsandsocialdevelopmentofthe
calf in theSerengeti.Zeitschrift fürTierpsychologie51:233-251.
&LLS K.,KRANZ K. LUNDRIGAN B.1986. Mother-young relationships in captive ungulates:
variability and clustering. AnimalBehaviour34: 134-145.
tuJ.1988. Ecologiadelzorro, Vulpesvulpes,en IaReserva Biolégicade Donana, Iluelva,SO
España. TesisDoctoral,UniversidaddeSevilla.
.JTBERG A.T. 1987.Adaptive hypotheses ofbirth synchrony inruminants: as interspecific test.
The AmericanNaturalist5: 693-710.
SADLF.IRR.M.F.S. 1984.Ecologicalconsequenceoflactation. ActaZoologicaPennica171:179-182. SANJosé C.1988. Estrategia reproductiva delashembrasdegamo(Damadama).TesisDoctoral,
Unive&dad ComplutensedeMadrid.
SANJOSéC.,BRAZAF.VARELAI.1989.Captura ymarcajedecrIasdegamo eneiParqueNacional
de Donana. ActasIX BienalRealSociedadEspañola HistoriaNatural9 (1):289-301.
SIEGELS.1956.Nonparametric statisticsforthebehavioralsciences.NewYork:McGraw-HillBook
Co., 312 pp.
STEIGERS W.D. JR & FuNonisJ.T. 1981. Mortality and movements of mute deer fawns in
Washington. Journalof Wildlife Management44:381-388.
STERBA 0.& KLOSAK IC. 1984. Reproductive biologyof fallowdeer, Damadama.B. Female reproduction. Acta ScientiarumAcademiaeScientiarum Bohemoslovacae,Brno18: 1-46.
TINBERGEN N., IMPEKOvEN M. FRANCE D. 1967. Anexperimeot onspacing-out asadefence against predation. Behaviour28: 307-321.
TRIu.MIcHF.1981.Mutualmother-puprecognition inGalapagosfursealsandsealions:cuesused and functional significance. Behaviour78: 21-42.
TRIJETTJ.C. 1979. Observations ofcoyote predation on muledeer fawnsinArizona. Journalof
Wildlife Management43: 956-958.
VENEROJ.L. 1982.DictadelosgrandesfitófagossilvestresdclParque Nacionalde Doflana.Tesis
Doctora4tiniversidaddeSevilla.
VosA.nt 1960. Behaviorofbarren ground caribou on theircalvinggrounds. JournalofWildlife
Management24: 250-258.
WIuTEM.,KNOWLTON F.F. BLAZF.NERW.C. 1972. Effectsofdam-newbornfawnbehavioron
captureand mortality. Journalof Wildlife Management36: 897-906.