= N=
aifferences in swimming and diving abilities between two sympatric species of water shrews: Neomys anomalus and N. fodiens (poricidae)
=
eeäena=jbkabpJplAobpNI=O=and=ieszek=ovCeifhNI=P= = eeäena=jbkabpJplAobpI=Njammaä=oesearch=fnstitute=of=the=moäish=Academó=of= pciencesI=taszkiewicza=NI=NTJOPM=BiałowieżaI=moäandX=Oéresent=addressW=aeéartment=of= BioäoÖóI=fndiana=rniversitóI=BäoominÖtonI=gordan=eaää=APMUI=NMMNI=bast=qhird=ptKI= BäoominÖtonI=fk=4T4MRI=rKpKAKI=eJmaiäW=mhmendes]indianaKedu= = ieszek=ovCeifhI=Njammaä=oesearch=fnstitute=of=the=moäish=Academó=of=pciencesI= taszkiewicza=NI=NTJOPM=BiałowieżaI=moäandX=Péresent=addressW=fnstitute=of=bnvironmentaä= BioäoÖóI=cacuätó=of=BioäoÖóI=Adam=jickiewicz=rniversitóI=rmuätowska=UVI=SNJSN4=moznańI= moäandI=eJmaiäW=äróchäik]amuKeduKéä= = = NV=text=éaÖesI=N=tabäe=and=4=fiÖures= = = = correséondinÖ=addressW= = mrofK=rAj=arK=habK=ieszek=ovCeifh= fnstitute=of=bnvironmentaä=BioäoÖó= cacuätó=of=BioäoÖó= Adam=jickiewicz=rniversitó= rmuätowska=UV= SNJSN4=moznań= moäand= eJmaiäW=äróchäik]amuKeduKéä=or=róchäikäesz]ÖmaiäKcom= teäKW=EH4USNF=UOVJRTJRN= faxW=EH4USNF=UOVJRSJPS= = =
= O Abstract N=
pwimminÖ =and =divinÖ =abiäities =of =two =sóntoéic =séecies =of =water =shrewsI =Neomys= O= anomalus=and=N.=fodiensI=were=tested=in=aquaria=usinÖ=videoJrecordinÖs=taken=from=three= P= views=Eäateraä=distantI=äateraä=cäose=and=dorsaäFK=qhe=frequencó=and=totaä=duration=of=divinÖI= 4= as=weää=as=the=mean=duration=of=divinÖ=and=fäoatinÖ=boutsI=were=siÖnificantäó=hiÖher=in=N.= R= fodiens=than=in=N.=anomalusK=crequencó=of=éaddäinÖ=durinÖ=surface=swimminÖ=was=äower=in= S= N.=fodiens=than=in=N.=anomalusK=N.=fodiens=dived=mainäó=for=äonÖ=distances=at=the=bottom=of= T= the=aquarium=and=éerformed=a=wider=ranÖe=of=dive=érofiäes=than=N.=anomalusI= which= U= éreferred=rather=short=and=shaääow=divesK=qhe=two=séecies=differed=aäso=siÖnificantäó=in=their= V= cineness=oatios=EdescribinÖ=how=streamäined=their=bodies=areF=when=divinÖK=then= NM= swimminÖI= N.=fodiens=had=a=reäativeäó=wider=bodó=and=éerformed=narrower=movements= NN= with=its=taiä=than=N.=anomalusK=qhese=resuäts=show=quantitativeäó=and=quaäitativeäó=for=the= NO= first=time=that=N.=fodiens=is=more=éroficient=at=swimminÖ=and=divinÖ=than=N.=anomalusK=qhe= NP= resuäts=aäso=heäé=to=exéäain=the=interséecific=differences=in=efficiencó=of=underwater= N4= foraÖinÖI=and=suééort=the=idea=of=seÖreÖation=of=ecoäoÖicaä=niches=of=these=séecies=based=on= NR= their=different=foraÖinÖ=modes=EdivinÖ=vs=wadinÖFK= NS= = NT= heó=wordsW=foraÖinÖ=behaviourX=interséecific=coméetitionX=microhabitat=seäectionX= NU= underwater=foraÖinÖX=water=shrews= NV= = OM= = ON= OO=
= P fntroduction OO= Neomys=anomalus=and=Neomys=fodiens=are=two=séecies=of=water=shrews=that=äive= OP= mainäó=aäonÖ=smaää=water=courses=and=éondsI=and=in=swamés=and=marshó=meadows= O4= Eoóchäik=OMMMFK=AäthouÖh=the=distribution=of=N.=anomalus=maó=be=conditioned=bó=the= OR=
éresence= of= N.= fodiens= Eqorre=and=qeääa=NVV4X=hróštufek=et=aäK=OMMMFI=theó=occur= OS= sóméatricaääó=and=sóntoéicaääó=in=manó=areas=EpéitzenberÖer=NVVMI=NVVVX=oóchäik=NVVTI= OT= OMMMFK=qhereforeI=their=ecoäoÖicaä=niches=overäaé=in=different=dimensions=Eoóchäik=OMMMI= OU= OMMRX=Churchfieäd=and=oóchäik=OMMSF=and=theó=coméete=for=food=resources=and=äife=séace= OV= Eoóchäik=NVVTI=OMM4X=oóchäik=and=wwoäak=OMMRI=OMMSFK=eoweverI=in=manó=éäaces=theó= PM= coexist=for=a=äonÖ=timeI=so=some=mechanisms=seéaratinÖ=their=niches=must=be=éresentK= PN= Both=séecies=have=moréhoäoÖicaä=adaétations=Efeet=and=toes=with=frinÖes=of=stiff= PO= hairsI=taiä=with=keeäI=reduced=earsI=cäoseabäe=nostriäsF=to=their=semiJaquatic=äife=mode= PP= Eeutterer=NVURX=mucek=NVUNX=Churchfieäd=NVVUFK=qhe=éeäaÖe=of=water=shrews=is=aäso=denser= P4= than=that=of=terrestriaä=shrews=and=the=awn=hairs=éossess=an=eJshaéed=érofiäe=with=äateraä= PR= ridÖes=fiääed=with=numerous=ridÖesI=which=is=beäieved=to=retain=airI=érovidinÖ=better= PS= insuäation=and=buoóancó=EChurchfieäd=NVVUFK=AäthouÖh=N.=anomalus=is=stronÖäó=associated= PT= with=waterI=its=frinÖes=of=stiff=hairs=on=the=feet=and=the=keeä=on=the=taiä=are=äess=érominent= PU= than=in=N.=fodiens=Emucek=NVUNX=Churchfieäd=NVVUFK=fn=the=wiädI=N.=anomalus=érefers=éäaces= PV= fäooded=with=shaääow=waterI=whereas=N.=fodiens=érefers=those=with=direct=access=to=streams= 4M= and=deeé=water=Eoóchäik=OMMMFK=qhe=diet=of=N.=anomalus=is=coméosed=onäó=or=mostäó=of= 4N= terrestriaä=éreó=Ekiethammer=NVTUX=huvikova=NVURaI=NVUTX=Churchfieäd=and=oóchäik=OMMSFI= 4O=
whereas =ué =to =VRB =of =the =diet =of =N.= fodiens= maó=be=coméosed=of=aquatic=éreó== 4P= Ekiethammer=NVTUX=aumasquier=and=Cantoni=NVVOFK=Aää=these=attributes=suÖÖest=better= 44= swimminÖ=and=divinÖ=abiäities=in=N.=fodiens=than=in=N.=anomalusK= 4R= qhese=abiäitiesI=as=weää=as=underwater=huntinÖ=tactics=or=äearninÖ=skiääsI=have=been= 4S= investiÖated=severaä=times=in=N.=fodiens=EeKÖK=outhardt=and=pchröéfer=NVURX=höhäer=NVVNI= 4T=
= 4 NVVPI=NVVSX=eaberä=NVVPX=soÖeä=et=aäK=NVVUFK=outhardt=and=pchröéfer=ENVURF=described=two= 4U= underwater=huntinÖ=tactics=for=this=séeciesW=in=oéen=water=for=éeäaÖic=éreó=and=at=the= 4V= bottom=for=benthic=éreóK=N.=fodiens=was=abäe=to=imérove=its=food=findinÖ=in=underwater= RM= bowäs=and=mazes=throuÖh=triaäJandJerror=äearninÖ=Ehöhäer=NVVPI=NVVSFK=fts=maximaä=divinÖ= RN= deéth=was=OSM=cm=and=the=number=of=dives=éer=daó=exceeded=ROM=EsoÖeä=et=aäK=NVVUFK= RO= bxceét=for=the=work=of=eaberä=ENVVPFI=these=abiäities=had=not=éreviousäó=been=examined=in= RP= N.=anomalusI=enhancinÖ=the=need=for=the=éresent=studóK= R4=
qhe =aim =of =this =work =wasI =thereforeI =to =anaäóse =and =coméare =quaäitativeäó =and = RR=
quantitativeäó=the=swimminÖ=and=divinÖ=behaviour=of=N.= anomalus= and= N.=fodiens= in= RS= caétivitóK=fn=the=érevious=studó=Eoóchäik=NVVTF=it=was=shown=that=success=and=effectiveness= RT= of=underwater=foraÖinÖ=are=hiÖher=in=N.=fodiens=than=N.=anomalusK=fn=the=éresent=studóI=we= RU= were=interested=in=answerinÖ=a=question=whó=the=two=séecies=differed=in=success=and= RV= effectiveness=of=underwater=foraÖinÖI=iKeK=in=findinÖ=mechanistic=differences=EeKÖK=in= SM= swimminÖ=veäocitóI=strokinÖ=frequencóI=bodó=incäinationI=fineness=ratio=or=divinÖ=érofiäesF= SN= reséonsibäe=for=more=efficient=swimminÖ=and=divinÖ=in=one=séecies=than=in=anotherK=cor=this= SO= éuréoseI=siméäe=exéerimentaä=conditions=EeKÖK=aquaria=without=restinÖ=éäacesI=aquatic=éreó= SP= or=oééortunitó=for=séontaneous=swimminÖF=seemed=to=be=Öood=enouÖhK=te=adoéted=the= S4= hóéothesis=that=N.=fodiens=swims=and=dives=more=efficientäó=than=N.=anomalus=andI=based= SR= on=theoreticaä=works=Ecish=NVVPbX=Biewener=OMMPFI=tested=the=foääowinÖ=éredictionsW=EaF= SS= durinÖ=surface=swimminÖI=the=anÖäe=between=the=bodó=and=water=surface=wiää=be=smaääer=in= ST= N.=fodiens=than=in=N.=anomalusI=EbF=veäocitó=of=swimminÖ=wiää=be=hiÖher=in=N.=fodiensI=EcF= SU= bodó=of=N.=fodiens=wiää=be=more=streamäinedI=EdF=N.=fodiens=wiää=Öeneraääó=séend=more=time= SV= divinÖI=and=EeF=the=duration=of=divinÖ=bouts=wiää=be=äonÖer=in=N.=fodiens=and=theó=wiää=swim= TM= a=äonÖer=distance=underwaterK=ft=was=aäso=assumed=that=the=differences=found=wouäd=heäé=to= TN= exéäain=the=mechanisms=of=resource=éartitioninÖ=and=seéaration=of=ecoäoÖicaä=niches=amonÖ= TO= these=shrewsK= TP=
= R
Material and methods T4= qested=animaäs=ENM=individuaäs=of=N.=anomalus=and=V=of=N.=fodiensX=aää=subaduätI=iKeK= TR= fuääó=Örown=but=sexuaääó=inactiveF=were=traééed=in=wet=habitat=around=Białowieża=viääaÖe= TS= Ekarewka=oiver=vaääeóI=eastern=moäandF=from=AuÖust=to=kovemberI=OMMP=and=OMM4K=qheó= TT=
were =keét =for =N =to =R =months =in =the =äaboratoró =of =the =jammaä =oesearch =fnstitute =mAp = TU= EjofF=in=BiałowieżaI=under=a=naturaä=äiÖhtJandJdark=cócäeI=NRJOMºC=and=humiditó=caK=UMBK= TV= bach=shrew=was=keét=in=an=individuaä=caÖe=EPU=˜=PM=˜=heiÖht=NS=cmF=equiééed=with=a= UM= sheäter=Ereversed=éot=fiääed=with=mossF=and=äitter=Ea=mixture=of=sandI=sawdustI=éeatI=and= UN= mossFK=tater=and=food=Eminced=beefI=fäó=äarvaeI=dried=dammarus= séKF=were=Öiven=ad= UO= libitumK=paméäe=size=was=increased=with=N=individuaä=of=N.=anomalus= and=P=of=N.=fodiens= UP= studied=bó=io=in=NVVO=at=the=rniversitó=of=lsnabrück=EdermanóF=under=simiäar=conditions= U4= of=caétivitó=and=reÖime=of=handäinÖ=and=fiäminÖK== UR= = US= mrocedure=of=experiments= UT= bxéeriments=were=carried=out=between=NSWMM=and=OMWMM=in=the=äaboratoróI=usinÖ=two= UU= aquaria=to=anaäóse=different=behavioursK=lne=äarÖe=aquarium=ENMM=˜=PM=˜=heiÖht=4M=cmF=was= UV= used=for=äateraä=distant=and=äateraä=cäose=recordinÖsI=with=the=fiäminÖ=beinÖ=done=from=one= VM= of=the=äonÖ=sides=of=the=aquariumK=qhis=aquarium=had=caK=P=cm=of=éebbäes=on=the=bottom= VN= and=a=water=deéth=of=OR=cmK=qhe=other=aquarium=ESM=˜=PM=˜=heiÖht=PM=cmX=water=deéth=OM= VO=
cmF =was =used =for =dorsaäJview =recordinÖsK =Both =aquaria =had =a =N =˜ =N =cm =Örid =on =the = VP= backÖround=waääK=qhe=teméerature=of=air=and=water=was=caK=NUºCK= V4= bxéeriments=with=each=animaä=consisted=of=NJR=triaäs=Eusuaääó=RF=and=each=triaä= VR= consisted=of=three=RJminute=recordinÖs=of=the=three=different=tóées=éerformed=in=a=random= VS=
orderK =qhe =animaäs =were =éäaced =in =the =aquaria =on =a =startinÖ =éäatform =Ea =movabäe =éäastic = VT=
sheäf=at=one=of=the=shorter=sidesF=for=NRJPM=secondsI=and=were=induced=to=swim=bó=säowäó= VU=
äowerinÖ=the=éäatform=under=the=waterK=Between=the=äateraä=distant=and=äateraä=cäose= VV=
= S
recordinÖsI=the=animaäs=were=aääowed=to=rest=on=the=startinÖ=éäatform=for=aééroximateäó=O= NMM=
minutes =and =Öiven =a =smaää =éiece =of =minced =meat =Eno =food =was =in =waterFK =Between =the = NMN= äateraä=and=dorsaä=recordinÖsI=the=shrews=were=transferred=to=their=caÖes=where=theó= NMO= remained=from=PM=minutes=to=a=few=daósK=qo=keeé=the=waterJreéeääent=nature=of=the=fur= NMP= intactI=animaäs=were=carried=in=a=wooden=box=or=heäd=bó=the=taiä=with=forceés=EiKeK=their=fur= NM4= was=not=handäedF=durinÖ=transfer=to=and=from=the=test=aquariaK=qhis=waóI=the=animaäs’=fur= NMR= was=sufficientäó=dró=on=aää=the=triaäs=éerformedK=bach=animaä=was=submitted=to=one=triaä=éer= NMS= daó=but=was=tested=NJR=times=Eusuaääó=R=times=durinÖ=R=successive=daósFK=bveró=few=triaäsI= NMT= the=waääs=of=aquaria=were=cäeaned=and=the=water=éartiaääó=reéäacedK= NMU= qhe=behaviour=of=the=water=shrews=was=recorded=usinÖ=a=aiÖitaä=sideo=Camera= NMV= ponó=aCoJsuOMMM=bK=qhe=camera=was=éroÖrammed=with=Abp=éroÖramI=a=shutter=séeed= NNM= of=SMMM=and=a=manuaä=focus=of=the=cäosest=views=for=better=imaÖe=quaäitóK=aurinÖ=fiäminÖI= NNN= two=äamés=of=TR=and=NRM=watts=were=éäaced=on=the=sides=of=the=aquariaK= NNO= NNP= aata=analysis= NN4= qhe=recordinÖs=were=transéosed=Ewith=Broadwaó=verK=RKN=softwareF=to=diÖitaä=format= NNR= EKjéÖ=and=KméeÖF=and=the=fiäes=were=anaäósed=frameJbóJframe=EOR=framesLsecF=usinÖ=Coreä= NNS=
mhotoJmaint=V®=and=Coreä=araw=V®K=qhe=foääowinÖ=cateÖories=of=behaviour=were= NNT=
distinÖuishedW =EaF =swimminÖ =J =the =animaä =activeäó =used =its =éaws =to =move =on =the =water = NNU= surfaceX=EbF=fäoatinÖ=J=the=animaä=staóed=on=the=water=surface=but=did=not=move=or=used=its= NNV= éaws=onäó=for=baäanceX=EcF=divinÖ=J=the=animaä=moved=with=its=whoäe=bodó=under=waterK=qhe= NOM= foääowinÖ=features=of=each=behaviour=were=anaäósed=from=the=different=view=recordinÖsI= NON= with=an=accuracó=of=MKM4=s=EN=frameF=for=durationI=N=cm=for=distance=and=MKN=deÖrees=for= NOO= anÖäeW= NOP= NF=iateraä=distant=viewW=EaF=éroéortion=of=behaviours=J=éroéortion=of=each=behaviour= NO4= EfäoatinÖI=swimminÖ=and=divinÖF=in=the=totaä=duration=of=the=recordinÖX=EbF=swimminÖ=séeed= NOR=
= T –=measured=as=the=distance=swam=bó=the=animaäs=durinÖ=N=secondX=EcF=divinÖ=érofiäes=Esee= NOS= beäow=for=definitionFX=and=EdF=duration=of=each=divinÖ=boutK=== NOT= OF=iateraä=cäose=viewW=EaF=stroke=frequencó=of=éaddäinÖ=–=number=of=strokes=éerformed= NOU= bó=one=hind=äimb=in=N=second=durinÖ=surface=swimminÖX=EbF=incäination=–=the=anÖäe= NOV= maintained=between=the=centraä=äonÖ=axis=of=the=animaä’s=bodó=and=the=water=äineX=EcF= NPM= duration=of=each=fäoatinÖ=boutX=EdF=duration=of=each=divinÖ=boutX=EeF=the=cineness=oatio=–= NPN= maximum=äenÖth=to=maximum=heiÖht=of=the=bodó=Ethe=measure=of=streamäininÖ=J=cish= NPO= NVVPbX=ciÖK=NF=whiäe=swimminÖI=divinÖ=and=fäoatinÖX=and=EfF=the=divinÖ=érofiäes=Esee= NPP= beäowFK= NP4= PF=aorsaä=viewW=EaF=width=of=bodó=as=coméared=to=améäitude=of=taiä=beatinÖ=–=maximum= NPR= width=of=bodó=to=maximum=améäitude=of=the=taiä=beatinÖ=at=the=same=momentX=EbF=width=of= NPS= the=bodó=as=coméared=to=its=äenÖthK= NPT= mrofiles. qhe=divinÖ=érofiäes=éerformed=bó=shrews=were=cäassified=accordinÖ=to=NS= NPU= modeäs=EciÖK=OFK=qhese=were=a=priori=ideaäized=accordinÖ=to=four=factors=and=a=score=was= NPV= Öiven=accordinÖ=to=an=assumed=quantitó=of=enerÖó=exéended=for=each=action=EN=for=äower=or= N4M= O=for=hiÖher=enerÖetic=costX=for=scores=see=ciÖK=OFW=EaF=descent=anÖäe=J=obtuse=vs=acuteX=EbF= N4N= deéth=of=the=dive=J=shaääow=Eué=to=the=haäf=of=the=water=deéthI=iKeK=caK=NO=cmF=vs=deeé=Emore= N4O= than=NO=cmI=usuaääó=to=the=bottomFX=EcF=distance=swum=underwater=reachinÖ=the=deeéest= N4P= éoint=J=short=Eué=to=OR=cmF=vs=äonÖ=Emore=than=OR=cmFX=and=EdF=ascent=anÖäe=J=obtuse=vs= N44= acuteK=qhe=scores=of=each=érofiäe=were=added=for=these=factors=and=three=difficuätó= N4R= cateÖories=were=estabäishedW=easó=J=totaä=score=of=4=or=R=Eérofiäes=mN–mRFI=medium=J=totaä= N4S= score=of=S=Eérofiäes=mSJmNNF=and=difficuät=J=totaä=score=of=T=or=U=EmNOJmNSFK=qhe=reaä=dives=of= N4T= water=shrewsI=of=courseI=did=not=have=to=start=at=äeft=edÖe=of=aquarium=Eas=shown=in=ciÖK=OF= N4U= but=couäd=start=at=anó=éoint=of=water=surfaceK=kot=onäó=straiÖhtJäine=but=aäso=curveJäine=dives= N4V= were=incäuded=into=the=anaäósisK= NRM=
= U
aata organization. bxceét=for=the=divinÖ=érofiäesI=data=for=each=anaäósed=éarameter= NRN= underwent=PJÖraduaä=averaÖinÖW=Nstº=averaÖe=was=caäcuäated=for=a=Öiven=triaäI=Ondº=averaÖe= NRO= for=each=tested=shrew=EincäudinÖ=ué=to=R=triaäsFI=and=Prdº=averaÖe=for=séecies=Eué=to=NO= NRP= shrewsFK=qhe=Prdº=averaÖes=are=shown=in=the=tabäe=and=fiÖuresI=whereas=the=Ondº=averaÖes= NR4= were=used=in=statisticaä=anaäóses=as=reéäicationsK=qhis=data=maniéuäation=was=done=in=order= NRR= to=reduce=intraJ=and=interindividuaä=variation=and=avoid=éseudoJreéäicationK= NRS= ptatistical analysis.=jannJthitneó=rJtest=was=used=Ewith=pqAqfpqfCA=verK=SF=for= NRT= interséecific=statisticaä=coméarisons=of=aää=behaviours=exceét=the=divinÖ=érofiäesK= NRU= aistributions=of=frequencies=of=different=tóées=of=érofiäes=were=coméared=bó=hoämoÖorovJ NRV= pmirnov=test=EpvpqAq=verK=RX=pokaä=and=oohäf=NVVRFK=qhis=informed=us=about=the= NSM= siÖnificance=of=Öeneraä=differences=between=the=séecies=as=weää=as=differences=between=the= NSN= observed=and=exéected=distributions=within=each=séeciesK=te=assumed=that=the=shrews= NSO= wouäd=not=diséäaó=éreference=for=anó=éarticuäar=érofiäeI=so=the=observed=frequencó=wouäd= NSP= be=cäose=to=an=exéected=uniform=distribution=with=frequencies=of=each=érofiäe=tóée=Z=SKOR= NS4= ENMMB=˜=NLNS=Z=SKORFK=oeéäicated=ÖoodnessJofJfit=tests=EdJstatisticF=Epokaä=and=oohäf=NVVRF= NSR=
were =éerformed =to =coméare =the =totaä =number =of =dives =and =the =number =of =difficuät =dives = NSS= éerformed=bó=each=séeciesI=as=weää=as=in=éairwise=coméarisons=of=each=divinÖ=érofiäeK= NST= ptatisticaä=siÖnificance=was=considered=at=p=<=MKMR=EtwoJtaiäedFK=pome=behaviours=were=not= NSU= diséäaóed=bó=or=measurabäe=in=aää=tested=animaäsI=therefore=the=saméäe=size=n=differs=usuaääó= NSV= from=NN=in=N.=anomalus=and=NO=in=N.=fodiens.= NTM= bthical=note= NTN= qhe=animaäs=were=cared=for=in=accordance=with=the=duidelines=for=the=treatment=of= NTO= animals=in=behavioural=research=and=teaching=EAnonómous=OMMPFK=mreÖnant=and=äactatinÖ= NTP= femaäes=Eas=weää=as=other=unwanted=smaää=mammaäsF=were=immediateäó=reäeased=at=the= NT4= éäace=of=caétureK=qhe=duration=of=the=tests=EZ=RJminuteJrecordinÖsF=was=chosen=after= NTR= considerinÖ=the=exéeriments=éerformed=bó=soÖeä=ENVVMFI=in=which=animaäs=were=forced=to= NTS=
= V swim=or=dive=for=S=minutesK=qhe=tests=were=carefuääó=monitored=and=were=stoééed=if=anó= NTT= animaä=showed=an=aééarent=difficuätó=in=staóinÖ=afäoatK=qhis=haééened=to=onäó=one=animaäI= NTU= and=when=the=exéeriment=with=this=animaä=restarted=the=foääowinÖ=daóI=the=animaä=did=not= NTV= show=anó=siÖns=of=difficuätó=or=extreme=stressK=kone=of=the=animaäs=died=durinÖ=or=directäó= NUM= after=a=testK=te=have=obtained=éermission=EnoK=ailmihoÖK=4OMNJM4JNPSLOMMNLOMMO=of=OU= NUN= cebruaró=OMMOF=from=the=jinister=of=bnvironment=for=our=caéturinÖ=of=the=érotected= NUO= shrews=and=an=acceétance=EnoK=OMMNLNN=of=NN=ganuaró=OMMNF=from=the=iocaä=bthicaä= NUP= Commission=for=bxéeriments=with=Animaäs=in=Białóstok=EmoäandF=for=our=exéerimentaä= NU4= methodsK= NUR= = NUS= oesults NUT= qhe=éroéortions=of=different=behaviours=varied=between=the=séecies=EciÖK=PFK=Both= NUU= séecies=séent=most=of=their=time=swimminÖ=EcaK=VOB=for=N.=anomalus=and=UMB=for=N.= NUV= fodiensFI=but=this=time=was=siÖnificantäó=äonÖer=in=N.=anomalus=than=in=N.=fodiens=EjannJ NVM= thitneó=rJtestW=r=Z=SKMI=p===MKMMPFK=phrews=swam=aää=over=the=aquarium=but=éreferred=to= NVN= staó=near=the=waääs=and=cornersK=qhe=duration=of=fäoatinÖ=did=not=differ=between=séecies=Er= NVO= Z=ONKMI=p===MKN4VFK=thiäe=fäoatinÖ=the=animaäs=wouäd=scratch=themseävesI=Öroom=their=fur= NVP= or=äook=around=the=aquariumK=qhe=time=séent=divinÖ=was=siÖnificantäó=Er=Z=4KMI=p===MKMMOF= NV4= shorter=in=N.=anomalus=than=N.=fodiensK= NVR= = NVS= purface=behaviours= NVT= qhe=séeed=of=surface=swimminÖ=did=not=differ=siÖnificantäó=between=séecies=Er=Z= NVU= 4MKMI= p ===MKVSRX=qabäe=NFI=but=the=strokinÖ=frequencó=of=éaddäinÖ=was=hiÖher=in=N.= NVV=
anomalus=than=N.=fodiens=Er=Z=OOKRI=p === MKMNPFK =qhe =animaäs =éaddäed =with =an =aäternate =
OMM=
strokinÖ=of=the=front=and=hind=äimbs=on=a=transverse=éäaneK=qhe=incäination=of=swimminÖ= OMN=
animaäs=from=horizontaä=was=quite=äow=for=both=séecies=and=did=not=differ=siÖnificantäó=Er=Z= OMO=
= NM PSKMI= p ===MKPMRFK=fn=both=séecies=the=nose=and=head=were=aäwaós=just=above=the=water= OMP= surface=and=the=taiä=unduäated=on=or=near=the=surface=EciÖK=NFK=qhe=ratio=of=the=maximum= OM4= width=of=the=bodó=to=the=maximum=améäitude=of=the=taiä=movement=was=siÖnificantäó=hiÖher= OMR=
in= N.=fodiens=than=in=N.=anomalus= Er=Z=PKMI=p ===MKMM4FI=indicatinÖ=that=N.= fodiens= OMS= éerformed=narrower=movements=with=the=taiäK=eoweverI=N.=fodiens=had=a=siÖnificantäó= OMT= wider=bodó=Er=Z=PKMI=p===MKMM4F=in=coméarison=to=N.=anomalusK=== OMU= qhe=mean=duration=of=sinÖäe=fäoatinÖ=bouts=was=siÖnificantäó=äonÖer=in=N.=fodiens= OMV= than=in=N.=anomalus=Er=Z=NOKMI=p===MKMNOX=qabäe=NFK=qo=fäoatI=the=animaäs=wouäd=éroject= ONM= their=äimbs=downwards=and=their=bodó=wouäd=be=äess=stretchedX=the=taiä=aééeared=to=fäoat= ONN= without=anó=active=effort=from=the=animaäK=qhe=cineness=oatio=EcoF=whiäe=fäoatinÖ=was= ONO= quite=äow=for=both=séecies=Eqabäe=NF=and=no=siÖnificant=interséecific=difference=existed=Er=Z= ONP= NTKMI= p ===MKURRFK=Aäso=when=swimminÖI=the=co=did=not=differ=siÖnificantäó=between=the= ON4= séecies==Er=Z=RNKMI=p===MKPRSFK= ONR= = ONS= aiving=behaviour= ONT= qhe=interséecific=differences=whiäe=divinÖ=were=more=distinct=than=for=surface= ONU= behavioursK=qhe=mean=duration=of=sinÖäe=divinÖ=bouts=was=siÖnificantäó=äonÖer=in=N.=fodiens= ONV=
than=in= N.=anomalus= Er=Z=VKMI=p === MKMMPX =qabäe =NFK =qhe =co =of =divinÖ =shrews =was = OOM= siÖnificantäó=hiÖher=in=N.=anomalus=than=N.=fodiens=Er=Z=NMKMI=p===MKMMNFK=fn=both=séeciesI= OON= the=co=increased=in=the=foääowinÖ=orderW=fäoatinÖ=Y=swimminÖ=Y=divinÖ=Eqabäe=NFI=and=aää= OOO= éairwise=intraséecific=differences=between=cofäoatinÖI=coswimminÖ=and=codivinÖ=were=siÖnificant= OOP= Er=Z=from=MKM=to=ORKM=and=p===from=MKMMN=to=MKMNOFK= OO4= qhe=totaä=number=of=dives=diséäaóed=bó=NO=individuaäs=of=N.=fodiens=was=4=times= OOR= hiÖher=than=that=diséäaóed=bó=NN=individuaäs=of=N.=anomalus=ETRU=vs=NUPI=ciÖK=4X=this=Öives= OOS= SPKO=vs=NSKS=divesLindividuaäI=oeéäicated=ÖoodnessJofJfit=testsW=d=Z=OVKMOI=p=<=MKMMNFK=qhe= OOT= =
= NN interséecific=difference=was=much=hiÖher=when=the=number=of=difficuät=dives=onäó=Eérofiäes= OOU= mNPI=mN4I=mNRI=mNSF=was=coméaredW=it=was=NM=times=hiÖher=in=N.=fodiens=than=N.=anomalus= OOV= E4RP=vs=4O=difficuät=dives=or=PTKU=vs=PKU=difficuät=divesLindividuaäI=d=Z=POKO4I=p=<=MKMMNFK= OPM= deneraä=distribution=of=frequencies=of=different=tóées=of=divinÖ=érofiäes=EciÖK=4F=did= OPN= not=differ=siÖnificantäó=between=the=two=séecies=EhoämoÖorov–pmirnov=testW=a=Z=MKNUUI=p= OPO= ==MKUVRFK=eoweverI=these=distributions=were=veró=different=from=the=exéected=uniform= OPP= distribution=Ea=Z=MKSUUI=p=<=MKMMN=for=N.=anomalusX=a=Z=MKUTRI=p=<=MKMMN=for=N.=fodiensFI= OP4= indicatinÖ=that=both=séecies=diséäaóed=some=éreferencesK== OPR= Both=séecies=éreferred=the=érofiäes=mNM=and=mNR=Eoeéäicated=ÖoodnessJofJfit=testsW=d= OPS= Z=RKRPU=to=4MKNMUI=p=<=MKMOR=to=MKMMNFX=N.=anomalus=éreferred=additionaääó=mU=Ed=Z=UKSVVI= OPT= p =<=MKMMRF=and=tended=to=érefer=mO=Ed=Z=PKR44I=p =<=MKNX=ciÖK=4FK=qhe=éreferred=érofiäes= OPU= EmUHmNMHmNRF=constituted=RVKMB=ETPKUB=incäudinÖ=mOF=of=aää=divinÖ=érofiäes=éerformed=bó= OPV= N.=anomalus=and=TOKPB=EmNMHmNRF=in=N.=fodiensK=N.=anomalus=éerformed=the=easó=érofiäe= O4M= mO=and=the=medium=difficuätó=érofiäe=mU=siÖnificantäó=more=frequentäó=than=N.=fodiens=Ed=Z= O4N= SKNVVI=p=<=MKMOR=and=d=Z=VKUUSI=p=<=MKMMRI=reséectiveäóFK=fn=contrastI=N.=fodiens=éerformed= O4O= the=difficuät=érofiäe=mNR=much=more=frequentäó=than=N.=anomalus=Ed=Z=NTKOTSI=p=<=MKMMNFK= O4P= thiäe=swimminÖ=on=the=bottomI=N.=fodiens=stoééed=and=chanÖed=the=swimminÖ=direction= O44= much=more=frequentäó=than=N.=anomalusK= O4R= qhe=common=érofiäes=avoided=bó=the=two=séecies=were=aää=shaääow=and=usuaääó=äonÖ= O4S= dives=m4I=mTI=mVI=mNNI=and=mNO=Ed=Z=4KMMN=to=UKSS4I=p =<=MKMR=to=MKMMRX=ciÖK=4FK= O4T= AdditionaääóI=N.=anomalus=avoided=mNP=and=mNS=Ed=Z=RKSNVI=p=<=MKMOR=in=both=casesF=and= O4U= N.=fodiens=avoided=mN=Ed=Z=RKSUPI=p=<=MKMORFK=qhe=avoided=érofiäes=constituted=OKTB=and= O4V= NKTB=of=the=totaä=divinÖ=érofiäes=observed=in=N.=anomalus=and=N.=fodiensI=reséectiveäóK=N.= ORM= anomalus=never=éerformed=P=out=of=the=NS=érofiäes=Em4I=mTI=mNOFI=whereas=N.=fodiens=onäó= ORN= one=EmNOFK=joreoverI=N.=anomalus=avoided=three=tóées=of=difficuät=érofiäes=EmNOI=mNPI= ORO= mNSFI=whereas=N.=fodiens=onäó=one=EmNOFK= ORP=
= NO aiscussion OR4= Both=séecies=séent=most=of=their=time=in=the=aquaria=swimminÖ=on=the=surface=and= ORR= frequentäó=tried=to=äeave=water=at=the=corners=of=aquariaK=mossibäó=the=shrews=did=not=feeä= ORS= comfortabäe=under=the=exéerimentaä=conditions=Eoéen=water=surfaceI=no=firm=Öround=to=restF= ORT= and=their=behaviour=couäd=be=infäuenced=bó=stressK=qhese=conditions=érevented=aäso=the= ORU= shrews=from=séontaneous=swimminÖ=and=did=not=Öive=them=motives=Eaquatic=éreóF=for= ORV= divinÖK=eoweverI=the=most=iméortant=fact=is=that=both=studied=séecies=underwent=the=same= OSM= exéerimentaä=conditions=and=diséäaóed=a=äist=of=siÖnificant=interséecific=differences=Ein=S= OSN= out=of=NM=coméared=éarameters=éäus=differences=in=divinÖ=érofiäesFK=And=we=were=interested= OSO= in=these=differences=rather=than=whether=the=behaviour=observed=in=our=aquaria=mirrored=the= OSP= behaviour=of=water=shrews=in=the=wiädK=joreoverI=in=the=éresent=studóI=we=were=interested= OS4= onäó=in=mechanistic=differences=in=swimminÖ=and=divinÖI=not=in=the=motives=or=resuäts=of= OSR=
such=behavioursK=aifferences=between=N.=anomalus= and= N.=fodiens=in=terrestriaä=and= OSS= underwater=exéäorinÖ=and=foraÖinÖ=Eeséeciaääó=the=hiÖher=success=and=effectiveness=in= OST= underwater=foraÖinÖ=of=N.=fodiensF=under=conditions=ÖivinÖ=shrews=manó=motivesI=stimuäi= OSU= and=oééortunitó=for=séontaneous=swimminÖ=and=divinÖ=have=been=aäreadó=known=Eoóchäik= OSV= NVVTFK= OTM= qhe=aäternate=stroke=of=the=front=and=hind=äimbs=on=a=transverse=éäane=was=observed= OTN= in=both=séeciesK=ft=is=considered=as=an=efficient=waó=of=swimminÖ=Eeickman=and=jachiné= OTO= NVUSFI=howeverI=such=quadruéedaä=éaddäinÖ=is=äess=effective=than=aäternate=biéedaä=éaddäinÖ= OTP= of=hind=äimbsI=tóéicaä=for=most=weääJadaéted=semiaquatic=mammaäs=Ecish=NVVPaI=bF. qhe= OT4=
swimminÖ =veäocities =of =the =studied =séecies =were =not =siÖnificantäó =different =from =one = OTR= anotherK=qhis=is=in=contrast=to=our=éredictionI=since=the=hind=feet=Ethe=main=éroéuäsive= OTS= éower=of=water=shrewsF=are=äonÖerI=the=frinÖes=denser=and=äonÖerI=and=consequentäó=the= OTT= éaddäinÖ=surface=is=äarÖer=in=N.=fodiens=Emucek=NVUNFK=eoweverI=this=discreéancó=couäd=be= OTU= readiäó=exéäained=bó=differences=in=the=strokinÖ=frequencó=of=éaddäinÖW=N.=anomalusI=with=a= OTV=
= NP smaääer=éaddäinÖ=surfaceI=had=to=stroke=more=frequentäó=EUKOO=strLs=vs=TKRS=strLsF=to=reach= OUM= the=same=veäocitó=as=N.=fodiensK=rnfortunateäóI=we=were=not=abäe=to=determine=the=stroke= OUN= frequencó=whiäe=divinÖ=from=our=recordinÖsK OUO= qhe=swimminÖ=veäocities=measured=in=our=studó=EPSK4=cmLs=in=N.=fodiens=and=PPK4= OUP= cmLs=in=N.=anomalusF=are=säiÖhtäó=äower=than=those=recorded=for=N.=fodiens=éreviousäó=EcaK= OU4= 4M=cmLs=J=outhardt=and=pchröéfer=NVURX=PSKTJ44KN=cmLs=J=höhäer=NVVNFK=qhis=maó=resuät= OUR= from=the=shorter=aquarium=used=in=our=exéeriments=ENMM=cm=coméared=to=PMM=and=N4R=cm= OUS= in=the=two=mentioned=éaéersFK= OUT= qhe=taiä=of=the=two=séecies=unduäates=whiäe=swimminÖ=on=the=surfaceI=which=is=an= OUU= indication=of=Öood=swimminÖ=skiääs=ECook=et=aäK=OMMNFK=iike=in=other=semiJaquatic= OUV= mammaäsI=eKÖK=the=muskrat=Ecish=NVUOI=NVVPbFI=the=taiä=of=water=shrews=is=säiÖhtäó=fäattened= OVM= at=the=sides=due=to=the=keeä=of=stiff=hairs=on=the=ventraä=side=Emucek=NVUNFI=and=it=moves= OVN= sónchronousäó=with=the=strokes=of=the=hind=feetK=qhis=is=beäieved=to=counterJbaäance=the= OVO= óawinÖ=that=wouäd=be=Öenerated=bó=the=aäternate=strokinÖ=of=the=hind=äimbsI=and=therefore= OVP= increases=stabiäitó=durinÖ=swimminÖI=reduces=draÖ=and=aääows=the=animaäs=to=éresent=a=more= OV4= streamäined=frontaä=view=to=the=water=fäow=Ecish=NVUOX=hróštufek=et=aäK=OMMMFK=N.=fodiens= OVR= éerformed=narrower=movements=with=its=taiä=Ein=reäation=to=its=bodó=widthF=than=N.= OVS=
anomalusK =AééarentäóI =thanks =to =the =äonÖer =keeä =and =its =Öreater =thrust =ÖenerationI =stabäe =
OVT= swimminÖ=was=obtained=bó=N.=fodiens=with=a=smaääer=améäitude=of=taiä=beats=than=in=N.= OVU= anomalusK= OVV= Both=water=shrew=séecies=swam=in=a=simiäarI=aämost=horizontaä=mode=with=their= PMM= nose=near=the=water=surfaceK=AäsoI=both=séecies=séent=some=time=fäoatinÖI=with=the=totaä= PMN= duration=of=this=behaviour=beinÖ=somewhat=äonÖer=in=N.=fodiens=than=in=N.=anomalusK=qhe= PMO= mean=duration=of=fäoatinÖ=bouts=wasI=howeverI=siÖnificantäó=äonÖer=in=N.=fodiens=than=in=N.= PMP= anomalusK=qhis=aÖain=suÖÖests=that=N.=fodiens=is=better=adaéted=to=water=than=N.=anomalusK= PM4= qhis=behaviour=has=not=been=described=so=far=for=these=séeciesK=qhe=animaäs=aééarentäó= PMR=
= N4 fäoated=whiäe=restinÖ=or=éreéarinÖ=for=a=diveK=cäoatinÖI=as=weää=as=swimminÖ=in=a=horizontaä= PMS= éositionI=can=be=attributed=to=hiÖh=buoóancó=Ecish=NVVPaFI=which=in=water=shrews=resuäts= PMT= from=äarÖe=amounts=of=air=traééed=in=their=fur=EAééeät=NVTPX=fvanter=NVV4FK=eorizontaä= PMU= éosition=and=hiÖh=buoóancóI=in=turnI=can=be=considered=an=indication=of=Öood=swimminÖ= PMV= skiääs=because=éaddäinÖ=forces=can=be=mostäó=used=for=éroéuäsion=and=not=wasted=in= PNM= maintaininÖ=the=head=above=the=water=äeveä=Ecish=NVVPaX=Cook=et=aäK=OMMNFK=qhis=aäso=shows= PNN= that=moréhoäoÖicaä=adaétationsI=eséeciaääó=those=reäated=to=the=insuäatinÖ=éroéerties=of=the= PNO= furI=are=veró=effective=and=iméortant=for=the=swimminÖ=behaviour=of=water=shrewsK=qhese= PNP= adaétations=aääow=the=animaäs=to=save=enerÖó=bó=reduction=of=heat=äoss=EsoÖeä=NVVMX= PN4= Churchfieäd=NVVUF=as=weää=as=bó=faciäitation=of=fäoatinÖ=and=returninÖ=to=the=water=surface= PNR= after=divinÖK=eoweverI=a=hiÖh=buoóancó=aäso=increases=the=enerÖó=costs=of=éaddäinÖ= PNS= downwards=and=remaininÖ=submerÖed=in=the=water=coäumnK= PNT= qhe=duration=of=sinÖäe=dives=bó=N.=fodiensI=reÖistered=in=different=studiesI=varied= PNU= from=P=to=NS=seconds=in=äaboratoró=conditionsI=and=from=P=to=O4=seconds=in=the=wiäd PNV= EChurchfieäd=NVVUX=soÖeä=et=aäK=NVVUFK=fn=our=exéerimentsI=N.=fodiens=séent=on=averaÖe=P= POM= seconds=and=N.=anomalus=O=seconds=under=water=Ewith=a=maximum=of=4=and=OKP=secondsI= PON= reséectiveäóFK=AäthouÖh=these=resuäts=can=be=infäuenced=bó=size=of=the=aquarium=and=äack=of= POO= éreóI=the=most=iméortant=observation=is=that=the=mean=duration=of=divinÖ=boutsI=as=weää=as= POP= the=totaä=time=séent=divinÖ=and=totaä=number=of=dives=éerformedI=were=äonÖerLhiÖher=in=N.= PO4= fodiens=than=in=N.=anomalusK=qhese=resuäts=are=consistent=with=our=érediction=andI=toÖether= POR= with=interséecific=differences=in=moréhoäoÖicaä=adaétationsI=maó=exéäain=whó=N.=fodiens= POS= foraÖes=more=efficientäó=in=deeé=water=Eoóchäik=NVVTF=and=eats=more=aquatic=éreó= POT= EChurchfieäd=and=oóchäik=OMMSFK=N.=anomalus=seems=to=foraÖe=bó=wadinÖ=in=shaääow=water= POU= Eoóchäik=NVVTI=OMMMX=Churchfieäd=and=oóchäik=OMMSF=and=it=maó=dive=to=exéäore=the= POV= environment=or=to=escaée=from=éredators=rather=than=to=foraÖeK= PPM= qhe=cineness=oatio=Ea=measure=of=streamäininÖ=J=cish=NVVPbF=of=both=water=shrews= PPN=
= NR increased=in=the=foääowinÖ=orderW=fäoatinÖ=Y=surface=swimminÖ=Y=divinÖI=which=is=äoÖicaä= PPO= since=the=benefit=of=streamäininÖ=increases=in=the=same=orderK=joreoverI=coméression=and= PPP= äoss=of=air=bubbäes=from=the=fur=durinÖ=deeé=divinÖ=enhance=the=streamäininÖ=of=these= PP4= séeciesK=thiäe=swimminÖ=and=divinÖI=both=séecies=had=cos=EPKNV=to=PKURF=within=the=ranÖe= PPR= of=favourabäe=vaäues=EP=to=T=J=cish=NVVPbFK=kevertheäessI=these=vaäues=were=äower=than=the= PPS= most=oétimaä=coZ4KR=Ecish=NVVPbFK=eoweverI=such=oétimaä=vaäues=are=observed=in=aquatic= PPT= mammaäsI=eséeciaääó=in=éursuitJtóée=éredators=äike=doäéhins=and=seaäs=Ecish=NVVPbFK= PPU= pemiaquatic=water=shrews=do=not=éursue=their=éreó=underwaterI=so=their=co=can=be=äowerK= PPV= ln=the=other=handI=the=fact=that=durinÖ=divinÖ=N.=anomalus=was=fäatter=EcoZPKURF=than=N.= P4M= fodiens=EPKRRF=can=be=exéäained=bó=different=divinÖ=modesK=N.=fodiens=dived=mostäó=aäonÖ= P4N= the=bottom=EmNRF=and=frequentäó=chanÖed=directionI=stoééedI=and=shrank=or=bent=its=bodó=at= P4O= the=bottom=irreÖuäaritiesK=fn=contrastI=N.=anomalus=dived=more=in=oéen=water=EmOI=mUI=mNMF= P4P= and=its=bodó=was=usuaääó=stretched=in=straiÖhtJäine=movementK=Another=cause=can=äaó=in=a= P44= difference=in=fur=quaäitóW=éossibäó=äess=air=was=retained=in=the=fur=of=N.=anomalus=than=N.= P4R= fodiensK= P4S= Both=séecies=éreferred=the=deeé=érofiäes=mNM=Eof=medium=difficuätóF=and=mNR= P4T= EdifficuätFI=but=N.=fodiens=éerformed=mNR=more=frequentäó=than=N.=anomalusK=N.=anomalus= P4U= éerformed=mO=Eshaääow=and=easóF=and=mU=Eof=medium=difficuätóF=more=frequentäó=than=N.= P4V=
fodiensK =pimiäaräóI =in =the =exéeriment =of =eaberä =ENVVPFI =dives =to =the =bottom =of =aquarium =
PRM= amounted=ué=to=OB=in=N.=anomalusI=whereas=4MB=in=N.=fodiensK=deneraääóI=in=our=studó=the= PRN= divinÖ=érofiäes=éerformed=bó=N.=fodiens=were=more=variabäe=and=this=séecies=avoided=Enever= PRO= éerformedF=onäó=one=difficuät=érofiäeK=N.=anomalus=never=éerformed=three=érofiäes=and= PRP= avoided=three=difficuät=onesI=indicatinÖ=that=adaétation=for=divinÖ=is=äower=in=this=séeciesK= PR4= eoweverI=more=detaiäed=exéeriments=utiäizinÖ=äarÖer=tanks=Ewith=shaääow=and=deeé=water= PRR= coméartments=and=the=éresence=of=benthic=and=éeäaÖic=éreóF=wouäd=be=usefuä=to=further= PRS= examine=the=divinÖ=abiäities=of=each=séeciesK=Additionaä=studies=examininÖ=the= PRT=
= NS éhósioäoÖicaä=attributes=EiKeK=bäood=and=muscäe=oxóÖen=carróinÖ=caéacitó=and=aquatic= PRU= thermoreÖuäatoró=abiäitiesF=of=both=séecies=maó=aäso=érove=insiÖhtfuäK= PRV= kevertheäessI=the=resuäts=of=the=éresent=studó=are=veró=heäéfuä=in=understandinÖ=the= PSM=
fieäd=observationsW=N.=fodiens= érefers =sites =with =direct =access =to =streams =and =deeé =waterI = PSN= whereas= N.=anomalus=érefers=éäaces=with=shaääow=water=some=distance=from=the=stream= PSO= EoóchäikI=OMMMFK=fn=the=wiädI=aquatic=éreó=coméose=usuaääó=≥RMB=of=the=diet=of=N.=fodiens= PSP= EChurchfieäd=NVU4X=huvikova=NVURbF=and=maó=reach=VRB=Ekiethammer=NVTUX=aumasquier= PS4=
and=Cantoni=NVVOFK=fn=contrastI=the=diet=of=N.=anomalus= maó =be =coméosed =mostäó =of = PSR= terrestriaä=éreó=Ekiethammer=NVTUX=huvikova=NVUTX=Churchfieäd=and=oóchäik=OMMSF=or=onäó= PSS= of=terrestriaä=éreó=Ehuvikova=NVURaX=oamaähinho=NVVRFK=fn=äaboratoró=exéeriments=with= PST= shaääow=water=EPKR=cmFI=N.=anomalus=caétured=manó=aquatic=éreó=EfishI=crustaceansF=and= PSU= utiäised=them=in=simiäar=quantities=to=N.=fodiens=Eoóchäik=and=gancewicz=OMMOFK=eoweverI=in= PSV= exéeriments=with=deeé=water=E[NM=cmFI=N.=fodiens=found=and=took=foodI=whereas=N.= PTM= anomalus=dived=siÖnificantäó=shorter=than=N.=fodiens=and=did=not=take=food=éäaced=in=such= PTN= water=Eoóchäik=NVVTFK=qhusI=N.=fodiens=is=abäe=to=foraÖe=both=in=deeé=and=shaääow=waterI= PTO= whiäe=N.=anomalus=can=onäó=foraÖe=successfuääó=in=shaääow=water=Eoóchäik=NVVTI=OMMMFK=Aää= PTP= these=observations=are=consistent=with=the=resuäts=of=the=éresent=studó=that=demonstrates=for= PT4= the=first=time=quaäitativeäó=and=quantitativeäó=the=differences=between=N.=fodiens=and=N.= PTR= anomalus=in=swimminÖ=and=divinÖ=abiäitiesK=qhese=differencesI=in=turnI=maó=correäate=with= PTS= the=different=foraÖinÖ=modes=used=mainäó=bó=the=two=séeciesW=wadinÖ=bó=N.=anomalus=and= PTT= divinÖ=bó=N.=fodiensK=pimiäar=differences=in=swimminÖ=and=divinÖ=abiäities=have=been=found= PTU= in=rodents=and=attributed=to=the=seéaration=of=their=niches=EeKÖK=bsher=et=aäK=NVTUX=Cook=et=aäK= PTV= OMMNFK=te=can=thus=concäude=that=the=differentiation=in=divinÖ=abiäities=and=foraÖinÖ=modes= PUM= maó=be=one=of=the=behaviouraä=mechanisms=that=aääow=these=two=séecies=of=water=shrews=to= PUN= éartition=resources=and=seÖreÖate=their=ecoäoÖicaä=nichesI=and=in=consequence=to=coJexist= PUO= sóntoéicaääóK= PUP=
= NT Acknowledgments= PU4= = te=are=veró=Öratefuä=to=oK=pchröéfer=and=AK=pchweÖmann=for=access=to=aééaratus=and= PUR= äabI=and=their=heäé=in=lsnabrückX=to=pK=Carvaäho=and=jK=pimões=for=their=fieäd=and=technician= PUS= assistanceX=to=gKoKbK=qaóäor=and=dK=oamaähinho=for=stimuäatinÖ=discussionsX=and=to=eK=Cabraä= PUT= for=heäé=in=the=statisticaä=anaäósisK=mK=oedman=kindäó=iméroved=our=bnÖäishK=qhis=studó=was= PUU= financed=bó=the=ieonardo=da=sinci=mroÖramme=EAÖreement=no=mLMPLALcLmi=–=NORUMVFI=the= PUV= moäish=ptate=Committee=for=pcientific=oesearch=Eresearch=Örant=ko=S=mM4c=MPS=ONF=and=Centre= PVM= of=bxceääence=Bflqbo=EContract=ko=bshOJCqJOMMOJUMMNNI=buroéean=CommissionDs=Rth= PVN= cramework=mroÖrammeFK= PVO= = PVP= oeferences PV4= Anonómous=EOMMPF=duideäines=for=the=treatment=of=animaäs=in=behaviouraä=research=and= PVR= teachinÖK=Anim=Behav=SRWO4V–ORR= PVS= Aééeät=e=ENVTPF=ceäästrukturuntersuchunÖen=an=tasserséitzmäusen=ENeomys=fodiens= PVT= EmennantFFK=Abh=Ber=katurk=jus=±jauritianum≤=UWUNJUT= PVU= Biewener=AA=EOMMPF=Animaä=äocomotionK=lxford=rniversitó=mressI=lxford= PVV= Churchfieäd=p=ENVU4F=aietaró=seéaration=in=three=séecies=of=shrews=inhabitinÖ=waterJcress= 4MM= bedsK=g=wooä=OM4WONNJOOU= 4MN= Churchfieäd=p=ENVVUF=eabitat=use=bó=water=shrewsI=the=smaääest=of=améhibious=mammaäsK=fnW= 4MO= aunstone=kI=dorman=ji=EedsF=Behaviour=and=ecoäoÖó=of=riéarian=mammaäsK=CambridÖe= 4MP= rniversitó=mressI=CambridÖeI=éé=4VJSU= 4M4=
Churchfieäd=pI=oóchäik=i=EOMMSF=aiets=and=coexistence=in=Neomys= and= porex=shrews=in=
4MR= Białowieża=corestI=eastern=moäandK=g=wooä=OSVWPUNJPVM= 4MS= Cook=tjI=qimm=ojI=eóman=ab=EOMMNF=pwimminÖ=abiäitó=in=three=Costa=oican=dró=forest= 4MT= rodentsK=oev=Bioä=qroé=4VWNNMN–NNM4= 4MU= aumasquier=AI=Cantoni=a=ENVVOF=phifts=in=benthic=macroinvertebrate=communitó=and=food= 4MV= habits=of=the=waterJshrewI=Neomys=fodiens=EporicidaeI=fnsectivoraFK=Acta=lbcoäI=lbcoä= 4NM= deneraä=NPWUNJVV== 4NN= bsher=ogI=toäfe=giI=iaóne=gk=ENVTUF=pwimminÖ=behavior=of=rice=rats=Elryzomys=palustrisF= 4NO= and=cotton=rats=Epigmodon=hispidusFK=g=jammaä=RVWRRN–RRU= 4NP=
= NU cish=cb=ENVUOF=cunction=of=the=coméressed=taiä=of=surface=swimminÖ=muskrats=Elndrata= 4N4= zibethicusFK=g=jammaä=SPWRVNJRVT=== 4NR= cish=cb=ENVVPaF=Coméarison=of=swimminÖ=kinematics=between=terrestriaä=and=semiaquatic= 4NS= oéossumsK=g=jammaä=T4WOTRJOU4= 4NT= = 4NU= cish=cb=ENVVPbF=fnfäuence=of=hódrodónamic=desiÖn=and=éroéuäsive=mode=on=mammaäian= 4NV= swimminÖ=enerÖeticsK=Aust=g=wooä=4OWTVJNMN== 4OM= eaberä=t=ENVVPF=wur=ÖkoäoÖie=einheimischer=péitzmäuse=EporicidaeI=fnsectivoraF=und=ihres= 4ON= iebensraumes=am=Beiséieä=eines=taädviertäer=ceuchtbiotoés=und=exéerimenteääe=BearbeitunÖ= 4OO= ausÖewähäter=ethoäoÖischer=craÖesteääunÖenKF=aissertationI=rniversitó=of=siennaI=siennaI=éé= 4OP= OSR=H=Aééendix=éé=ONN= 4O4= = 4OR= eickman=dCI=jachiné=C=ENVUSF=pwimminÖ=behavior=in=six=séecies=of=African=rodents= 4OS= ECricetidaeI=juridaeFK=Acta=qherioä=PNW44VJ4SS= 4OT= = 4OU= eutterer=o=ENVURF=Anatomicaä=adaétations=of=shrewsK=jammaä=oev=NRW4PJRR= 4OV= == 4PM= fvanter=bs=ENVV4F=qhe=structure=and=adaétive=éecuäiarities=of=éeäaÖe=in=soricine=shrewsK=fnW= 4PN=
jerritt =gcI =hirkäand =digrI =oose =oh =EedsF =Advances =in =the =bioäoÖó =of =shrewsK =péeciaä = 4PO= mubäication=of=CarneÖie=juseum=of=katuraä=eistoró=koK=NUI=mittsburÖhI=éé=44NJ4R4== 4PP= höhäer=a=ENVVNF=kotes=on=the=divinÖ=behavior=of=the=water=shrewI=Neomys=fodiens= 4P4= EjammaäiaI=poricidaeFK=wooä=Anz=OOTWONUJOOU== 4PR= höhäer=a=ENVVPF=ln=the=äearninÖ=of=the=éosition=of=underwater=food=sources=bó=Neomys= 4PS= fodiens=EjammaäiaI=poricidaeFK=wooä=Anz=OPNWTPJUN= 4PT= = 4PU= höhäer=a=ENVVSF=rmweÖversuche=mit=tauchenden=tasserséitzmäusen=ENeomys=fodiensF= 4PV= EjammaäiaW=poricidaeFK=Bonn=wooä=Beitr=4SWPNRJPOR= 44M= = 44N= hróštufek=BI=aavison=AI=driffiths=ef=EOMMMF=bvoäutionaró=bioÖeoÖraéhó=of=water=shrews= 44O= ENeomys=sééKF=in=the=western=maäaearctic=oeÖionK=Can=g=wooä=TUWNSNSJNSOR= 44P= huvikova=A=ENVURaF=qhe=food=of=some=séecies=of=the=famiäó=poricidae=EfnsectivoraF=in=the= 444= aäder=forest=of=the=gurskó=sur=wetäandK=BioäóÖia=4MWNUNJNUT=xfn=päovak=with=bnÖäish=abstractz= 44R=
= NV huvikova=A=ENVURbF=wur=kahrunÖ=der=tasserséitzmausI=Neomys=fodiens=EmennantI=NTTNF=in= 44S= der=päowakeiK=BioäóÖia=4MWRSPJRTO== 44T= huvikova=A=ENVUTF=wur=kahrunÖ=der=puméfséitzmausI=Neomys=anomalus= CabreraI= NVMT= 44U= EfnsectivoraI=poricidaeF=in=der=päowakeiK=iónx=OPWRRJSO== 44V= = 4RM= kiethammer=g=ENVTUF=teitere=beobachtunÖen=über=tasserséitzmäusse=der=Arten=Neomys= 4RN= fodiens=und=N.=anomalusK=w=päuÖetierk=4PWPNPJPON== 4RO= mucek=w=EedF=ENVUNF=heós=to=vertebrates=of=moäandK=jammaäsK=mtk=–=moäish=pcientific= 4RP= mubäishersI=tarszawa= 4R4= oamaähinho=jd=ENVVRF=mreäiminaró=account=of=the=bioäoÖó=of=Neomys=anomalus=CabreraI= 4RR= NVMT=in=mortuÖaäK=fnW=mroceedinÖs=of=the=±peminar=on=the=bioäoÖó=and=conservation=of= 4RS=
buroéean =desmans =and =water =shrews =Edalemys=pyrenaicus,=aesmana=moschata,=Neomys= 4RT= sééKF≤I=lrdesaI=péainI=TJNN=gune=NVVRK=Counciä=of=buroéeI=ptrasbourÖI=qJmsp=EVRF=POWSPJSS= 4RU= outhardt=jI=pchröéfer=o=ENVURF=wum=serhaäten=der=tasserséitzmaus=Neomys=fodiens= 4RV= EmennantI=NTTNF=unter=tasserK=AnÖew=wooä=TOENJOFW4VJRT== 4SM= oóchäik=i=ENVVTF=aifferences=in=foraÖinÖ=behavior=between=water=shrewsW=Neomys=anomalus= 4SN= and=Neomys=fodiensK=Acta=qherioä=4OWPRNJPUS== 4SO= oóchäik=i=EOMMMF=eabitat=éreferences=of=four=sóméatric=séecies=of=shrewsK=Acta=qherioä=4RI= 4SP= puééä=NWNTPJNVM== 4S4= oóchäik=i=EOMM4F=Coméetition=and=coexistence=of=shrewsK=fnW=gędrzejewska=BI=tójcik=gj= 4SR= EedsF bssaós=on=mammaäs=of=Białowieża=corestK=jammaä=oesearch=fnstituteI=moäish=Academó= 4SS= of=pciencesI=BiałowieżaI=éé=NSNJNTM= 4ST= oóchäik=i=EOMMRF=lveräaé=of=teméoraä=niches=amonÖ=four=sóméatric=séecies=of=shrewsK=Acta= 4SU= qherioä=RMWNTRJNUU= 4SV= oóchäik=iI=gancewicz=b=EOMMOF=mreó=sizeI=éreó=nutritionI=and=food=handäinÖ=bó=shrews=of= 4TM= different=bodó=sizesK=Behav=bcoä=NPWONSJOOP== 4TN= oóchäik=iI=wwoäak=o=EOMMRF=Behaviouraä=mechanisms=of=confäict=avoidance=amonÖ=shrewsK= 4TO= Acta=qherioä=RMWOUVJPMU= 4TP=
= OM oóchäik=iI=wwoäak=o=EOMMSF=fnterséecific=behaviouraä=domination=amonÖ=four=sóméatric= 4T4= séecies=of=shrewsK=Can=g=wooä=U4W4P4J44U= 4TR= pokaä=ooI=oohäf=cg=ENVVRF=BiometróK=qhe=érinciéäes=and=éractice=of=statistics=in=bioäoÖicaä= 4TS= researchK=te=creeman=and=ComéanóI=kew=vork= 4TT= péitzenberÖer=c=ENVVMF=dattunÖ=Neomys= hauéI=NUOVK=fnW=kiethammer=gI=haéé=c=EedsF= 4TU= eandbuch=der=päuÖetiere=buroéasK=Band=PLNK=fnsektenfresserI=eerrentiereK=AuäaJseräaÖI= 4TV= tiesbadenI=éé=PNTJPPP= 4UM= péitzenberÖer=c=ENVVVF=Neomys=anomalus=CabreraI=NVMTK=Neomys=fodiens=EmennantI=TTNFK=fnW= 4UN= jitcheääJgones=AgI=Amori=dI=BoÖdanowicz=tI=hróštufek=BI=oeijinders=mgeI=péitzenberÖer=cI= 4UO= ptubbe=jI=qhissen=gBI=sohraäik=sI=wima=g=EedsF=qhe=atäas=of=buroéean=mammaäsK=q=and=Aa= 4UP= moóser=itdI=iondonI=éé=RUJSN= 4U4= qorre=fI=qeääa=gi=ENVV4F=aistribution=of=the=Cabrera=water=shrew=ENeomys=anomalusF=in= 4UR= northeastern=péainK=w=päuÖetierk=RVWOUOJOUU= 4US= soÖeä=m=ENVVMF=Bodó=teméerature=and=fur=quaäitó=in=swimminÖ=waterJshrews=Neomys=fodiens= 4UT= EjammaäiaW=fnsectivoraFK=w=päuÖetierk=RMWTPJUM== 4UU= soÖeä=mI=Bodmer=CI=pérenÖ=jI=Aeschimann=g=ENVVUF=aivinÖ=caéacitó=and=foraÖinÖ=behaviour= 4UV=
of=the=waterJshrew=ENeomys=fodiensFK =fnW =aunstone =kI =dorman =ji =EedsF =Behaviour =and = 4VM=
ecoäoÖó=of=riéarian=mammaäsK=CambridÖe=rniversitó=mressI=CambridÖeI=éé=PNJ4T= 4VN=
= ON
qabäe=NK=pummaró=of=differences=between=Neomys=anomalus=and=N.=fodiens=in=surface= 4VO=
swimminÖI=fäoatinÖ=and=divinÖK=piÖnificance=of=the=differences=Ereveaäed=bó=jannJthitneó= 4VP=
rJtestFW=ns=–=insiÖnificantI=GG=p ≤=MKMNI=GGG=p ≤=MKMMRI=GGGG=p ≤=MKMMNK=qhe=saméäe=size=n=
4V4= varies=because=éarticuäar=éarameters=were=not=diséäaóed=bó=or=measurabäe=in=aää=tested= 4VR= animaäsK= 4VS= 4VT= N.=anomalus= N.=fodiens= marameter=of=behaviour= mean=–=pa= n= aiffeJ= rence= mean=–=pa= n= piÖnifiJ cance=of= difference= = pwimminÖ= seäocitó=EcmLsF= PPKPS=–=NKPR= V= Z= PSK44=–=OKP4= V= ns= ptrokinÖ=frequencó=EstrKLsF= UKOO=–=MK4T= NM= [= TKRS=–=MKSN= NN= GG=
fncäination=EºF= NOKMO=–=OKOR= NN= Z= NOKS4=–=NKMV= V= ns=
Bodó=width=W=taiä=width=ratio= NKMV=–=MKM4= U= Y= NKNP=–=MKMS= T= GGG=
Bodó=width=W=bodó=äenÖth=ratio= MKPS=–=MKMN= U= Y= MKPV==–=MKMN= T= GGG=
cineness=oatio= PKNV=–=MKPPV= NN= Z= PKOS=–=MKPNM= NN= ns=
= cäoatinÖ=
auration=of=sinÖäe=bout=EsF= NKOU=–=MKPO== V= Y= OKMM=–=MKT4== V= GG=
cineness=oatio= OKMO=–=MKNTU= R= Z= OKMR=–=MKNVT= S= ns=
= aivinÖ=
auration=of=sinÖäe=bout=EsF= OKNO=–=MKRR= V= Y= PKMU=–=MKRS= V= GGG=
= OO 4VT= ciÖK=NK=Neomys=fodiens=swimminÖ=at=the=water=surfaceK=jeasurements=for=the=cineness=oatio= 4VU= caäcuäation=Eco=Z=aLbF=and=the=éosition=of=nose=are=visibäeK= 4VV= RMM=
= OP = RMM= ciÖK=OK=pixteen=divinÖ=érofiäes=distinÖuished=in=this=studó=and=ordered=accordinÖ=to=their= RMN= difficuätó=Escores=from=4=to=UFK= RMO= RMP=
= O4 = RMP= ciÖK=PK=mroéortions=of=fäoatinÖI=swimminÖ=and=divinÖ=diséäaóed=bó=the=two=séecies=of=Neomys= RM4= durinÖ=the=totaä=time=of=a=testK=NMMB=Z=R=minutesI=n=–=number=of=water=shrews=that=diséäaóed= RMR= aää=three=kinds=of=behaviourK=piÖnificance=of=the=differences=Ereveaäed=bó=jannJthitneó=rJ RMS= testFW=ns=–=insiÖnificantI=GGG=p ≤=MKMMRK= RMT= RMU=
= OR
= RMU=
ciÖK=4K=aifferences=in=divinÖ=érofiäes=between=Neomys=anomalus=and=N.=fodiensK=piÖnificance= RMV=
Ereveaäed=bó=ÖoodnessJofJfit=dJtestF=of=interséecific=differencesW=G=p ≤=MKMRI=GGG=p ≤=MKMMRI= RNM= GGGG=p ≤=MKMMNX=differences=between=exéected=and=observed=vaäuesW=arrow=ué=J=éreferred= RNN= érofiäe=EsiÖnificantäó=more=frequent=than=exéectedFI=arrow=down=J=avoided=érofiäe= RNO= EsiÖnificantäó=äess=frequent=than=exéectedFK= RNP=
