Faecalmarkingbehaviourbyfree-rangingCommongenets Genettagenetta andEgyptianmongooses Herpestesichneutnon insouthwestern Spain
By F.PALOMARES
EstecidnBioldgicaJionana,ConsejoSuperiordeInvestigacionesCsentificas,Seville,Spain
Abstract
.1992
Studied defecation behaviour of the common generand the Egyptian mongoose at the Doñana National Park from November 1985to November 1989.Both speciesfrequentlyusedlatrines and morethanoneindividualcontributedtolatrineformation. Genetsplacedtheirlatrinesintreessituated ontheedgesofthemorefrequentlyusedhabitats,whereasmongoosesdefecatedontheground, and their latrines were found within preferred habitats, near to resting sites. Genets increased their markingbehaviourinFebruary—MarchandNovember—December,andmongoosesinJanuary—Febru ary and September—October.Latrine switching within thesameareawasobserved inmongooses. Datatend toindicatethatlatrinesareusedasacommunicationtoolinbothspecies.
Introduction
Faecesseem tobeusedbymanycarnivores asacommunicationtoolintheirsociallives (MACDONALD1980;GORMANand TROWBRIDGE 1989).Nevertheless, littleisknown about faecalorscentmarkingbehaviourinviverrids(GoR2wIand TROWBRIDGE1989), theonly studiebeingonfree-rangingcivets,Civeuictiscivetta,(BEArERandRANDALL1978)and captive African dwarf mongooses, Helogaleundulate,(RASA 1973),meerkats, Suricata suricata,(Mowc and Soaasrn1986), watermongooses,Atilax paludinosus,(BAKER
1988)andcommongenets,Genettagenetta,(ROEX3ER1980a).Furthermore,faecalmark ingbehaviour isdiscussedonlyoccasionallyinsomeofthesestudies. Here, someaspects offaecalmarking behaviour byfree-ranging common genets and Egyptian mongooses, I-Ierpestes ichneutnon,intheDonana National Park(southwesternSpain)aredescribed.
Studyarea
Thestudywascarriedout intwoareasoftheDonanaNstiooalPark:theBiologicalReserveandthe Coin del Rey. The Park issituated on the western bank oftheGuadalquivirRivtrmouth,in southwesternSpain(approximately37‘N,6°30’W),and includes threemain bioropes: marshesor marismas,scrublandanddunes.Themarshesareusuallyfloodedinwinter andcoveredbyScirpussp. Therearetwomaintypesofscrublaod:adry,xerophyticscrublandvegetatedbyHalimiumsp.and Rosmarinussp.,andadense,mesicscrubland withEricasp.andCallunavulgaris.Therearescattered corkoaks(Quercussuber)alloverthescrubland.Inthedunes,therearemobilebaresandduneswith valleyshetweeo,whichtypicallyarecolonizedby pine(Pinua pinea) forests.Theclimateis Mediterranean subhumid,characterizedbydry,hotsummersandmild,wetwinters.
TheBiologicalReserve(6700ha) issituatedinthemiddleofthePark,andthethreemainbiotopes
describedabovefortheParkcanbefound.TheCotodel Rey,of 3500ha,issituatedinthenorthern partofthePark,andischaracterizedbypinesandEucalyptussp.forestationsonthematorral, and abundantPistacialenriacuspatcheswherethegroundwatertableisnearerthesurface.
226 F Palomayes
Materialand methods
TheBiologicalReservewasexaminedfullyforgenetdefeéationsites,specialattentionbeingpaiedto treesandErieaandCaliunascrubland(themainhabitatofthespeciesintheatudyarea;PALOMARE5 andDEL5BES1988).Bycontrast,threeareasoftheBiologicalReserve(CanodelaRaya=CR,Lueio BoRn=LB,andLagunaSantaOlalla=LO), wheremongooses wereradio-tracked (PALOMARE5and DELIBE51991a),wereexaminedformongnnsefaeces.Greater searchingeffortwasundertakenfrom November 1985 to January 1986. Each defecation site was marked on the terrain. A site was consideredasalatrine iftherewere2ormorefaeces.Latrineswereperiodicallysampled(weeklyfrom November1985toApril 1986, and monthly from then toJanuary 1987).In total, 264and141 samplings were carried out on genet and mongoose latrines, respectively. Radio-trackingat the BiologicalReserveof2genets(anadultdispersingmale,andayoungfemale)and4mongooses(2adult and2young females)from September 1985toApril1986 (PALOMABE5andDELSBE51988,1991a) helpedtointerpret someoftheresultsonfaecalmarkingbehaviour. Habitat typewherelatrineswere found and theirsituation with respect toplacescommonly usedbyradio-trackedindividualswere noted.
Allfaeceswerecollectedfordietanalysis(PALOMARE5andDEL5EE51991b),buttheirnumberand positionwere noted before collecting.TotestpossibleinfluencesoffaecalremovaLongenetmarking behaviour,somefaeceswere left atthelatrineon64samplingoccasions,whereasinanother200 occasionsallofthefaeceswerecollected.Removaldidnotaffectpresenceorabsenceoffaecesonthe followingsamplingoccasion(G-testofindependence;G=0.002,p<0.001).
GenetlatrineswerealsointensivelysearchedforduringJune1987intheCotodelKey.Sporadic informationonmarkingbehaviourofbothspecieswasalsoobtainedfromMarch1987toNovember
1989whileradio-tracking24mongoosesand10genetsandcollectingfaecesforscatanalysisinthe
samearea(PALOMARZ5 andDamESl991a).
Results
Bothgenetsandmongoosesmadelatrines.Twenty-seven genetdefecationsites werefound ontheBiologicalReserve,with anumberoffaecesrangingfrom Ito27.Sevenofthese sitescontainedonly1scat;fifteenwereperiodicallysampled.Morethanonegenetseemed to use the latrines. The radio-tracked male was trapped at latrine 320, which was continuouslyused(Tab.1) despitethefactthatthisindividualneverreturned tothearea during5months ofradio-tracking(Tab.1).
Forty-onemongoose defecation siteswere found on the BiologicalReserve, with a
numberoffaecesrangingfrom1to40.Twenty-oneofthesesiteshadonly1scat;fifteen wereperiodically sampled. Different mongooses alsodeposited theirfaecesonthesame latrine. On 6January 1985,2mongooses(anadultradio-trackedfemaleandanuntagged animal)left3faecesafewcentimetersfromeachother,closetoarestingsite.AttheCoto delReyIobservednumerouslatrines usedbydifferentmongooses belongingtothesame familygroup.
Defecationsiteground
Genetfaeceswerefound intrees(44.4%on maintrunks,22.2%on secondary trunks,
14.8 %onraptor nestsand7.4% ontreebranches), onthickets orhedges(7.4%)andon theground (3.7%;n=27sites).Faecesintreesweresituatedonaverage4.2mhigh(SD=
2.8, range=2.5—14.0,n=22).Treeswerealwayscorkoaksexceptforonepine.
Whenusinglatrines, genetstended todeposit faecesoverthewhole availablesurface and,asarule,freshfaeceswereneverdeposited overother freshones,although overold ones.The onlylatrine found ontheground had9faecesdeposited oversemifallenrush busheswith thefurthestseparated by25m.
Atleast100cork oaksweresearchedfor genetlatrinesintheCoto delKey,butnone were found.Ontheotherhand,only3latrines(with3,6and21faeces)usedforafewdays werefound inthisareaduring radio-tracking activities.Scatteredfaecesweresometimes found,all ofthemsituatedontheground.
Table1.Numberof faecesby monthand latrinein commongenets from November1985to
January1987
Latrinenumber / NO / DE / JA / FE / MA / AP / MA / Months
313 / JU / AG / SE / OC / NO / DR / JA
320 / 202 / 72 / 4 / 5 / 4 / 3 / 2 / 2 / 1 / 3 / 23 / 18 / 8 / 2 / 2
215 / 32 / 02 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 1 / 0 / 0
204 / 02 / 22 / 1 / 0 / 1 / 0 / 0 / 0 / 0 / 0 / 1 / 0 / 1 / 2 / 3
308 / 192 / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 4 / 7 / 1
306 / 82 / 2 / 3 / 2 / I / 1 / 4 / 0 / 0 / 0 / 0 / 0 / 0 / 0
314 / 52 / 0 / 2 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 1 / 0 / 0
210 / Y / 1 / 0 / 1 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 1 / 0 / 1
415 / 32 / 0 / 2 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / —
281 / 92 / 1 / 8 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0
200 / 62 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0
300 / 272 / 4 / 12 / 4 / 3 / 1 / 6 / 0 / 0 / 1 / 3 / 7 / 4
261 / 2 / 3 / i / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0 / 0
418 / 142 / 8 / 4 / 0 / 0 / 3 / 13 / 1 / 0 / 1 / 2 / 3 / 2
388 / 32 / 0 / 0 / 0 / 2 / 0 / 2 / 8 / 7 / 4
282 / 42 / 1 / 0 / 0 / 0 / 0 / 0
1.5 / 2.2 / 2.5 / 0.6 / 0.4 / 0.7 / 1.4 / 0.5 / 1.6 / 1.5 / 1.9 / 2.0 / 1.2
1.7 / 2.4 / 3.7 / 1.3 / 0.9 / 1.3 / 3.7 / 0.9 / 5.9 / 4.6 / 2.7 / 2.9 / 1.5
thesedatawerenotusedtoobtain themonthly mean.
Allmongoosefaeceswerefoundontheground.Of17latrines,35.3%werebybramble edgesorother thick vegetation; 23.5% underthissamethick vegetation; 17.6% under pineshrubs,but clearlyvisible;11.8%overflattened grassandsurroundedby rushes, Juncussp.;5.9%onpathsamongvegetation;and5,9%atanopengroundelevationinthe middle of apool. Most of the detected latrines (76.5%) werein clearlyvisiblesites, whereas23.5%werehidden.However, densevegetationprobablymadeitdifficultforthe observer tofind thelatter. Scattered faeceswere mainly found on themiddle ofpaths within themostusedhabitats andonthoseleadingtorestingsites.
inmongooses,latrinesurfacesizesvariedgreatly,fromthosewithfaecesseparatedbya few centimeters, to others where there wasa continuumof faeces with the furthest separated by45m.
Placeoflatrines
Genetsmainlyinhabit mesicscrubland patchesintheBiologicalReserve(PALOMARE5and Delibes 1988) and preferentially deposited their faeces on the edges of these patches (69.2%;n=26;X2=3.85,p=0.0498).Insidethe1.4km2ofthefemalehomerange(the only individual that hadadcfined range; PALOMARJ5andDELIBE51988),three latrines werefound,situated bytheedgesofmesicscrubland.
Mongooses usemesicscrubland and brambles (Ri-sbussp.)torest(PALOMARE5and
Damns1990)andmostfaeceswerefoundverynearorinsidethishabitat.Thirteen of17 latrines (thosewith higher number offaecesandusedlonger)werelessthan50mfrom known nocturnal restingsitesandinsidehomerangecoreareasofradio-trackedindividu als.Duringintensive24-hour tracking periods mongooses wereobserved on5occasions defecatingclosetorestingsites.Thenumber oflatrinesinthehome rangesofthethree mostradio-trackedmongooseswere6(homerangeof1.7ken2),7(homerangeof4.8km2) and2(homerangeof5.7km2).
228 F.Palomares
Temporalpatternoflatrineuse
On average,the number offaecesatgenetlatriner was higher inFebruary—Marchand November—December,and lower from April to August, with aslight increaseinJuly (Tab.1).Of15latrines periodicallysampled,only5werecontinuouslyusedforatleast4 months, and only 1wasalways used (Tab.1).Two latrines (onewith 9faeceson the ground andanother with15faecesinaraptor nest)madebytheradio-tagged malewhile roaming bythe duneswerelocated. They werecloseto restingsites.Whenrestingsites weredeserted,thelatrines werenotusedanylonger.
Numbers of faeces on mongoose latrines wcre higher in September—Octoberand
February,with lower but similar numbers for the rest of the year (Tab.2). Usually mongoosesdidnotkeeplatrinesforlong.Latrineswerecommonly usedduringanisolated month, oratmost,for2or3consecutivemonths, although afteratimetheycould bere used.Twolatrineswereusedduring7 months (Tab.2).
Table2.Number offaecesbymonthandlatrineinEgyptainmongoosesfromNovember1985to
January1987
LatrineMonths
NPlaceOC NO DE JAFE MA AP MA JU JU AG SE OC NO DR
1B
2B
30
4B
50
60
70
8CR
9CR
100 ii0
12B
130
14B ISB
23221110
8003000
3001111
2063000
420
310
30———
20000
26139
$
00010000
000012000
00000000
00001000
000
000
—000———
00000000
00000100
00000000
113041000
963217 23650
2201————
9213000
403216a
9.0 1.5 2.0 3.54.7 2.5 2.1 2,2 2,4 2.9 1.8 8.2 3.5 1.9 0.0
9.7 1.8 2.1 6.8 10.5 5.2 3.5 4.4 6.2 8.7 4.5 13.5 9.6 4.9 0.0
B=LucioBolin,0= LagunaSantaOlalla,CR=CaflodeIaRaya.
aLatrine wasflooded.
Latrineswitchingbymongooses
Mongooses seemed to uselatrines alternatively on asame area.For instance, atBolin, latrine1wasthemostusedfrom October 1985toMarch1986,together latrines2and4; latrine12wasthemost usedfrom Mayto August 1986,andalthough mongooses never stopped defecating on latrine 12, latrine 15 was the most used from September to November 1986(seeFig.).AtOlalla,latrines3,5,6and7 were themost usedfrom the beginningofthestudytoApril1986,butfromMayon,onlylatrines11and13wereused (Fig.).
LUCIOBOLINLAGUNASANTA OLALLA
MONTHSMONTHS
L1 / 123L2 / LLJL4 / L3 / IZJL5 / LIJL.6 / LSJLTISDL12 / [XJL14 / us / LXILiD / Lii / CLu
Monthly contributionofeachlatrine(I.)tototalnumberoffaecescollectedofEgyptian mongoosesin
LucioBolinandLagunaSantaOlalla, bothintheDoñanaBiologicalReserve
Discussion
Theuoffaecesforcommunicationhasbeenwidelysuggestedinseveralcarnivorespecies
(MACDONALD1980;Gosu%t±u’tandTROWBRIDGE 1989),andthismightbealsothecasefor
genetsandmongooses sincetheyusedlatrinessituatedatspecificplaces.ROEDER (1980a)
concluded thatcaptivegenetsusetheirfaecesforexchanginginformation,andHEEETZ et al.(1984)found aspecificcomponentinthesubstancesecreted bytheperianalglandof
maleEgyptianmongooses.Theposition ofthisglandaround theanus,suggeststhatfaeces maybeimpregnated with differentodoursatleastbetweenmalesandfemales.Mongoose latrinesituation,nearrestingsitesandinsidecoreareas,suggeststhatfaecesmaybeusedas aself—recognitionoftheterrain,asterritorialmarkingofcoreareas,which areexclusive amongadultindWi&ntsofthesamesex(PALOMAKESandDEUWES1993a),andtofaciiite communicationamongfamilygroup members(EWER 1973;GOSLING andMcKAY1990).
DÜCXER (1965) remarkedthatViverrinaeusuallyusepermanentlatrines,whereas Herpestinaetendtoswitchtheirlatrinesoften.R0EDER(1980a)pointedoutthatgenetsuse permanent latrines, aspontaneouslatrine switch being very rare. In this study these tendencies were observed in genets and mongooses, although for the former itwas observed moresporadically than permanent latrines. Theformation oftemporal latrines seemsdifferentineachspecies.Roaminggenetssuchastheradio-trackedmale,couldbe thecauseinthisspecies.With mongooses, itcould bemainlyproduced byacontagious defecation by some mongooses travelling together, a frequent event in Doflana (PALOMARE5 andDELIBES 1993a).This could beascertained onseveraloccasionsinthe Coto dclRey.
Evenlackingdetailedinformation,themongoose behaviour observed inDoñana does notseemtocoincidewith thatreported byBEN-YsAcovandYou-Toy(1983)inIsrael,
230 F.Patomares
where mongooses appear to usepermanent latrines. In Israel, mongooses seemto use permanent resting sites (BEN-YAACov and YOM-T0V 1983), but in Doflana they often change reiting sites (PALOMAItES and DELIBE51993b). Sincemongooses tend to make latrines near restingsites, thisbehaviour might explainsuch differences.AlsoinIsrael, BEN-Ycovand Y0M-Tov(1983)foundscatteredfaeces,whichwereascribedtoroaming individuals.
Defecation sitesusedbygenetsinDoflana(mainly trees)havebeenlittlereported in
otherareas,whererocksarethecommonest(e.g.LIVErandReaDER1987).However,this differenceisprobably becauserocksarenot availableinDofiana.On theother hand,an interestingfindingwasthedifferentfaecalmarkingbehaviourfound betweengenetsliving intheBiologicalReserveandCorndelKey.Habitat compositionineachareaisdifferent, andthereforethismightbethereasonforthedifferences.Variationsinmarkingbehaviour attributed tohabitatchangesare frequentinothercarnivores(MACDONALD1980;SMITHet al.1989;DELIBESetal.1991).
Two annualpeaksinthenumber offaecesatlatrines wereobservedinbothspecies. ThoseofFebruary—MarchforgenetsandJanuary—Februaryformongoosescouldbedue to mating, which takes placeatthesetimes(AYMERTCH1982; PALOMARE5and DELtBES
1992).Moreover, anincreaseinmarkingbehaviourandinspectionofscentmarkingduring thematingperiodhas beenobservedincaptivegenets(ReaDER1980a,I980b),and tigers (SMITHetal.1989).ThepeaksdetectedinNovember—DecemberforgenetsandSeptem ber—Octoberfor mongooses could becausedby the dispersal ofyoung animalsin the former (see MACDONALD and MASON1987, for the otter, Lutralutra),and because mongooses travel more often infamily groups at that time (PALOMARESand DELIBES
1993a),there beingmoreindividualsabletodeposit theirfaecesatthesamesites.
Acknowledgements
TheresearchwassupportedbyDGICYT(projectPB87-0405).TheauthorhadapostdoctoraLgrantof ConsejoSuperiordeInvestigacionesCientIficas.C.DCLACouxvhelpedinthefieldworkwhile searching forgenetlatrinesintheCotodeL Rey area. M. Daisnas,S.M. MACDONALDand an anonymousrefereeprovidedusefulcomments onanearlydraftofthismanuscript.N.BU5TAMANTE
‘reviewedtheEnglishversionandC.KELLERtranslatedtheGermansummary
Zusamtnenfassung
MarkierungdurchDeflikarionbeifreilebendenGinsterkatzen,Genertagenetta,unitMangusten,
1-lerpestesid.sneurnon,inSlidwest-Spanien
DieDefãkationsgewohnheitenvonGinsterkatzenundMangustenwurden imNationalparkDonana vonNovember1985bisNovember 1989studiert. GinsterkatzenundMangustenlagernihrenKotoft inLatrinen ab.AlssolchebenutzenGinsterkatzenHohlstelleninBlumenimGrenzbereichihrer Habitate, wahrend Mangusteo ihrenKotamBodenablagern undihreLatrinen sichimBereichdes bevorzugten HabitatsundinderNIhevonSchlafpLatxenbefinden.Ginsterkatzenmarkierenintensi
vervonFebruar bisMarxundvonNovember bis Dezember,undMangusten vonJanuar biaFebruar undSeptember bisOktober. Beibeiden Artc-nwird dieselbeLatrine vonverachiedenenIndividuen benutzt. Esscheint, daLatrinen zumfnformationsaustauschzwischenIndividuen&enen.
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Author’saddress:Dr. FitANcssco PALOMARES,Eseaeión Bioltigica Doflana, Consejo Superior de
Investigaciones Cientfficas,Avda.MariaLuiaaa/n,E-41013Sevilla,Spain