Optimal Width of Movement Corridors for Root Voles: Not Too Narrow and Not Too Wide

Author(s): Harry P. Andreassen, Stefan Halle and Rolf Anker Ims

Source: Journal of Applied Ecology, Vol. 33, No. 1 (Feb., 1996), pp. 63-70
Published by: British Ecological Society
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Journalof
AppliedEcology
Optimalwidthofmovement forrootvoles:
corridors
1996,33, nottoo narrowand nottoo wide
63-70
HARRY P. ANDREASSEN, STEFAN HALLE and
ROLF ANKER IMS
DivisionofZoology,Department of Oslo, PO Box 1050 Blindern,
ofBiology,University N-0316Oslo,
Norway

Summary
of male root vole movementsas a functionof corridorwidth
1. The characteristics
weretestedin a 310m longhabitatcorridorconnectingtwohabitatpatches.Detailed
observationsof movementsweremade by meansof radiotelemetry and recordingof
footprints.
in termsoftransference
2. The highestconnectivity, rateofindividualsin thecorridor
system,was observedin theintermediate ofthreecorridorwidthstested(3 m, 1m and
0 4 m).
3. The behaviouralmechanismbehindthe lowerconnectivity of the narrowestcor-
ridorwas a reluctanceofvoles to enterit,whilelinearprogressin thewidestcorridor
was hamperedby a highfrequency of cross-directional
movements.
betweencorridorwidthand movementbehaviourwas unaffected
4. The relationship
by thesimulatedpresenceof competitors and predators.
5. Our resultschallengethe 'the-wider-the-better'principleof movementcorridor
design,and provideelementsforan understanding of the behaviouralmechanisms
underlying themovementecologyof individualsin linearhabitats.
Key-words: corridordesign,Microtus,movementecology.
connectivity,
JournalofAppliedEcology(1996) 33, 63-70

Plummer1993). As veryfewstudieshave been able

Introduction
to obtaindata about animalmovementsin corridors
Habitatfragmentation hasbeenrecognized as a major (Hobbs 1992),theempiricalbasis forcorridordesign
threat
towildlife populations(Diamond1976;Gilpin is poor (Simberloff&Cox 1987;Simberloff etal. 1992;
& Diamond1980;Higgs& Usher1980;Soule 1986; Mann & Plummer1993). For instance,theeffectsof
Lande 1988;Ims & Stenseth1989).As continuous structuralaspects of habitatcorridors,such as cor-
habitatsbecomefragmented, an immediate conse- ridorwidthand continuity,on movementrates are
quenceis thatthemobility of organisms becomes poorly known. Still, specificrecommendationson
restricted(Fahrig& Merriam1985; Stampset al. optimalcorridordesignmaybe foundin theliterature
1987a,b;Fahrig& Paleheimo1988;Burkey1989). (Noss 1987; Harrison 1992; Merriam & Saunders
Thisin turndecreasestheeffective size,and,conse- 1993). The urgentneed forempiricalstudiesaddress-
quently,the viabilityof populations(Soule 1986; ing basic questionsregardingmovementecology of
Boyce1992).It has beenclaimedthatthenegative individuals in linear habitats, has recentlybeen
effectsof habitatfragmentation can be reducedby stressed(see Hobbs 1992; Inglis& Underwood 1992;
connecting isolatedfragments by narrowstripsof Simberloffet al. 1992;Lindenmayer& Nix 1993).
habitat,termed movement corridors(Wilson& Willis We herepresentdata froma studywheremovement
1975;Harris1984;Bennett 1990;Saunders& Hobbs behaviour of the root vole Microtus oeconomus
1991).In thepresent paperwewilladheretothestrict (Pallas) in corridorsofvaryingwidthwas subjectto a
definitionof movement corridorsas linearhabitats detailedanalysis.We used corridorwidthas themain
allowingformovements, butnotforpermanent settle- treatmentvariable,because it is themostbasic struc-
ment. turalcharacteristicoflinearhabitatsuponwhichprac-
The establishment of movement corridors is cur- tical corridordesign inevitablyrequiresa decision.
? 1996British rentlyimplemented as an expensiveconservation Root voles were selectedas the studyorganismsas
Ecological Society strategyworld-wide etal. 1992;Mann&
(Simberloff theyhave proved to be veryusefulmodels forfield

63

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All use subject to JSTOR Terms and Conditions
64 experiments involvinghabitatmanipulations(Ims &
Optimalwidthof Stenseth1989; Ims et al. 1993; Wiens et al. 1993).
movement Moreover,althoughwe primarily designedthisstudy
corridors to serveas an empiricalmodel system(sensu Ims &
Stenseth1989) foraddressingbasic biologicalmech-
anisms that impingeon corridordesign(e.g. move-
mentbehaviour),our resultsmay have some direct
implicationsfortheconservationofremnantrootvole
populationsin Europe (van Apeldoornet al. 1992).
We showthatoptimalwidthofmovementcorridors
in termsof habitatconnectivity for root voles is a
compromisebetweennot too narrow and not too
wide. Hence, our resultschallengethe commonpre-
sumptionthat widercorridorsnecessarilyare better
corridors(Noss 1987; Harrison 1992; Merriam &
Saunders 1993).
Methods
STUDY PLOT AND MANIPULATIONS
The studytookplace at EvenstadField Station,south-
eastNorwayduringAugust-October1992.The move-
ment behaviourof individualmale root voles was
studiedin a fencedsystemconsistingof two habitat
patchesconnectedby a 310m long corridor(Fig. 1).
The lengthof the corridorwas about one order of
magnitudelonger than the diameterof an average
male rootvole homerangein non-linearhabitats(Ims
et al. 1993,and unpublished).
The twohabitatpatchesused as releasepoints(size:
5 x 5 m) and thecorridorconsistedof dense,homo-
geneous meadow vegetation,knownto be preferred
habitatby root voles (Ims et al. 1993). An artificial
runwaysystem(Fig. 1) was establishedas 0 1 m wide
vegetation-freepathsalong and acrossthecorridorto
facilitatemovements in theotherwiseverydensegrass
carpet: one 310m long mid-runwayand cross-run-
waysat 10-mintervals.Theserunwaystendedto direct
movementsacross footprintplates (see below). The
area betweenthefencesand thecorridorwas cleared
of vegetativecover by means of herbicides(Fig. 1).
The fenceswere establishedto hinderthe intrusion
of othervoles and mammalianpredatorsduringthe
experiment.We expectedthatthe absence of female
voles should motivateexploratorymovement(Ims
1988) or breedingdispersal (Kawata 1989) in the
experimentalmales. To ensure that the olfactorial
environment in thecorridorat thestartof theexperi-
mentwas not verydifferent to thesituationlateron,
some adult males were temporarilyreleased in the
corridora fewdays beforethefirstexperimental trial
was executed.
We testedthreedifferent corridorwidthsin com-
binationwithabsence/presence ofpredatorsand com-
petitors(Table 1), whichalso may affectmovements
(Saunders & Hobbs 1991). The studystartedwith
trialsin thewidestcorridor(3 m). The two narrower
corridors(I m and 0 4 m) weremade subsequentlyby
mowingand herbicideuse (see timescheduleforthe
varioustreatments in Table 1).
Presenceof competitorsand predatorsweresimu-
lated by six 'barriercages' whichwere permanently
situatedacross thecorridorat fixedintervals(Fig. 1).
The barriercages were made of wire mesh with a
centralpassage tunnelencompassingthemid-runway
of the corridor.The lengthof the barriercages was
adjustedaccordingto thedecreasingwidthofthecor-
ridor.For the treatment'competitor'(Table 1), one
live adult male was keptin each barriercage. Earlier
studiesin which wire mesh cages of a similartype
have been used to simulatepresenceof conspecific
individualsin microtineshave produced significant
responses(Ims 1988, 1990; Nelson 1994). For the
treatment 'predator',the barriercages weresupplied
withfreshfoxscatssealed in small,perforatedplastic
boxes. Simulatedpresenceof predatorsby artificial
additionof predatorscentshave earlierbeen shown

15m 30m 60m lOOm 60m 30m 15m

I i Iiu
Fence 0 Barriercage
- Vegetation - - - Runwaysystem
* Release point A Trackingplate

Fig.1. Designofthestudy plot.Thetwohabitat andthecorridor

patches consistedofdense,homogeneous meadow vegetation
(shadedarea).Theareaoutsidethecorridor (whiteareainsidethefence)consisted ofbareground.Theenlarged sectionof
? 1996 British thecorridor (inset)showsthedesignoftheartificial
runway releasepointandthepositioning
system, platesinthe
oftracking
Ecological Society, 3 mwidecorridor. Runways as 0 1mvegetation-free
wereestablished pathsalong(mid-runway) andacrossthecorridor
(cross-
JournalofApplied runways at 10-mintervals).Trackingplateswereplacedin bothtypesofrunways forthe0-4m
(exceptin thecross-runways
Ecology,33, 63-70 corridor)as wellas alongtheedgesofthecorridor.

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All use subject to JSTOR Terms and Conditions
65 Table 1. Experimentalprotocol:Competitor,otheradultmalesin thebarriercages (Fig. 1); Predator,foxscatsin thebarrier
H. P. Andreassen, cages; Control,emptybarriercages. Ten different
male rootvoles wereused in each of the9 treatmenttypes
S. Halle & Numberof samplerecords
R. A. Ims
Treatment Period* Radiotrackingt Footprintst

Width,3 m
Control 1-9 Aug 10 5
Predator 10-16 Aug 10 2
Competitor 18-26 Aug 9 5
Width,1m
Competitor 26 Aug-i Sep 10 5
Control 2-7 Sep 10 5
Predator 8-15 Sep 10 4
Width,0-4m
Predator 19-23 Sep 10 5
Competitor 24-8 Sep 9 4
Control 29 Sept-3 Oct 10 4

* The variablelengthsof the experimentalperiods(e.g. periods > 5 days) werecaused by instancesin whichindividual
malesin a trialwerenotcaughtbefore08.00h thenextday.
t Reductionin thesamplesize (from10 to 9) of radiotracking
recordsfortwoof thetreatments werecaused bydysfunction
of radiotransmitters.
t The reductionsin thenumberoffootprint recordsfroma maximumoffive(= numberofexperimental dayspertreatment)
forfourtreatment typeswerecaused by heavyrainfallsmakingfootprint platesunreadable.

to affectspace use in microtines(Jedrzejewskiet al. (1977). Devices for recordingfootprints(tracking

1993). For the 'control'treatment,the barriercages
were leftempty.Beforethe onset of each treatment,
the barriercages were carefullywashed to remove
remainingscent fromthe previoustreatment.Fur-
thermore,the cages were always placed on plastic
sheetsto preventthesoil beneaththecages becoming
contaminatedby scent.
TREATMENT AND MONITORING OF THE
STUDY ANIMALS
On each day,tworadiocollared(Biotrack,SS-2 trans-
mitters),adult male voles werereleasedat 08.00h in
the two habitatpatchesin the opposite ends of the
corridor(one male per patch, cf. Fig 1). Each trial
was terminated at 18.00h. Thereafter,
themaleswere
caughtbymeansoflivetrapsactivatedinthecorridor.
Trials weregenerallyconductedon consecutivedays,
provided that males from the previous trial were
caughtbefore08.00h thenextmorning.
Longitudinalmovements inthecorridorweremoni-
toredby locatingthe males (to the nearestmetrein
the longitudinaldirectionof the corridor)by means
of a probe antenna(Andreassenet al. 1993) at every
fullhour from09.00h to 17.00h. The resultingnine
radio-trackingpositionsobtainedpermale wereused
to estimatethe distancemoved,the speed of move-
mentand thetimespentin thecorridorforeach male
(forfurtherdescriptionoflongitudinalmovement par-
? 1996British
Ecological Society, ameters,see footnotesto Table 2).
JournalofApplied Cross-directionaluse of thecorridorwas registered
Ecology,33, 63-70 bythefootprint recordingtechniqueofKing & Edgar
plates) were situatedalong the corridoredges and
in the mid-runway and the cross-runways insidethe
corridor(Fig. 1). The use of corridoredges and the
two runwaytypesby voles werescoredaccordingto
thenumberofplateswithfootprints at theend ofeach
trial.Footprintcountswererecordedperexperimental
day ratherthan per individualsince footprintsof a
pair of individualsreleasedsimultaneously could not
be distinguished whenevertheyhad passed each other
inthecorridor.Radiotrackingshowedthatsuchcross-
ing occurredin at least 29% of the trials.Due to
the generallow sample size of footprintrecordsand
missingdata, leadingto unbalanceddesigns(Table 1),
statisticaltestsofcompetitor/predator treatmentwere
not appliedto thefootprint data.
The males used in thisstudywerelaboratoryborn
F2-F4 generationdescendantsof animalscaughtin
Pasvik, north-eastNorway. We chose to use lab-
oratory-raised animalsto standardizepriorexposure
to factorswhich mighthave influencedmovement
behaviour.Males werethechosensexas theyaremore
suitable'models' thanfemalesforthistypeof study;
theyappear to be more motivatedto disperse(Ims
1989;Kawata 1989),and theyare lesssubjectto short-
termbehaviouralchanges due to reproductivehor-
mone cyclesthatwould have introducedextraneous
variancein thedata.
Beforethetrialsin thecorridor,maleswerehoused
separatelyin laboratorycages exposed to constant
light,to mimicthe natural lightregimeduringthe
summerin north-east Norway,as wellas to avoid any
effectsof changingphotoperiodin the course of the

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66 Table 2. The effectof corridorwidthon movementand corridoruse parameters.Data are lumpedforthe treatment
factor
Optimalwidthof absence/presenceofcompetitor/predator.Values are presentedas mean +-standard errorof themean
movement Corridorwidth
corridors Parameters 3m Im 0 4m

Longitudinalmovements*
Maximumdistancereached(m) 75 + 16 205 + 22 35 + 15
Cumulativedistancetraversed(m) 163 + 33 440 + 57 51 + 21
Movementspeed/efficiencyt
Time spentin corridor(h) 79+ 0't 59 + 04 3-9 + 11
Maximumspeed (m/h) 61 + 11 160 + 19 93 + 24
Averagespeed (m/h) 21 + 4 76 + 11 51 + 13
Progressrate(m/h) 10 + 2 40 + 10 35 + 10
Cross-directional
movementst(footprint
counts)
Edge counts 1 42 + 0 51 0 50 + 0 20 0 21 + 0 11
Edge/mid-runway ratio 0 23 + 0-09 0 03 + 0-01 0 09 + 0-07
ratio
Cross-runway/mid-runway 1 56 + 0 31 0-36+ 0 05

* Longitudinalmovementparametersare based on the completedata set of radiotrackingrecords(Table 1). Maximum

distancereached= maximumdistancereachedfromreleasepatch.Cumulativedistancetraversed= cumulativemovedduring
thetrialas revealedby hourlyradiotracking.
t Movementspeed/efficiency parametersincludeonlytrialsin whichmales enteredthecorridor(N3.. = 26, NJ...= 30 and
No 4, = 8). Time spentin corridor= totaltimeperioda male was foundin thecorridor.Maximumspeed = longestdistance
movedduringone hour.Averagespeed = totaldistance/time spentin corridor.Progressrate = maximumdistance/time spent
in corridor.
t Cross-directional
use parametersare based on footprint countrecords(Table 1) fromthefootprint platessituatedalong
thecorridoredges (edge counts),in themid-runway and thecross-runways (Fig. 1). No estimateof theratiovariable(cross-
runway/mid-runway), is givenforthe0 4 m corridor,sincefootprints
werenot recordedin thecross-runways forthiscorridor
width.

experiment.All males used were sexually mature,
adult animals(> 50 g and scrotaltestes),and thusof
adequate size forthe2 5 g radiocollars.
Twenty-six differentindividualswereavailable for
thisstudy.Most ofthesehad thusto be usedin several
trials(X = 2 6 + 0 1 [SE] trialsper male), but never
more than once per treatment combination.Gener-
ally,10 differentmalesweretestedpertreatment com-
bination(Table 1). Males used in morethanone treat-
ment were released in alternatingpatches with the
longestpossible inter-trial intervalto avoid habitu-
ation to thestudyregime.Numberof trialsper male,
when included as a covariate in ANCOVA-models,
neverexplainedmorethan4% ofvariance(P > 0 50)
of any of themovementindicesestimated(Table 2).
Thisresultindicatesthatneitherhabituationnorother
possibleconfoundingtimeeffects, e.g. scentaccumu-
lation (see also Discussion) were importantin this
study.Males partneredin trialsinfrequently crossed
each otherin the corridor(29% of all trials) so it
is unlikelythatdirectinterference betweenthe free-
rangingmalesinfluenced theresults.Hence,we findit
justifiableto treateach trialpermale as thestatistical
unit(cf.Table 1) in theanalyses.
Results
Corridorwidthhad a substantialeffecton all move-
? 1996British
EcologicalSociety, mentparameters(Table 2) (two-wayANOVAS, all par-
JournalofApplied tialr2> 0 25, P < 0 001). Presence/absence
of pred-
Ecology,33, 63-70 atorsand competitors wererelatively
unimportant for
all parameterstestedwithrespectto thistreatment (all
partialr2< 0 04, P > 0 05).
Voles generallymoved furthest and fastestin the
intermediate width (1 m) corridor (Table 2).
Maximum distancesreached by voles in the inter-
mediatewidthcorridorwere on average threeand
six timeslongerthan in the widerand the narrower
corridors,respectively. In the intermediatecorridor
morethanone-thirdofthevolesreachedtheopposite
end, while this rarelyhappened in the wideror the
narrowercorridor(Fig. 2).
The low successof the narrowestcorridorin con-
nectingthe habitatpatcheswas caused by reluctance
of voles to enterit (Fig. 2); the narrowestcorridor
(04 m) was enteredonly in 27% of the trials,while
the two widercorridorswereenteredin themajority
of thetrials(100% and 83% entranceratein the 1m
and the 3m corridor, respectively)(X2= 40.3,
d.f. = 2, P < 0 001). This apparentaversionby voles
to thenarrowestcorridoris corroboratedbytheshort
time spent in this corridorby voles that actually
enteredit; on average,males enteringthewidestcor-
ridorspenttwiceas muchtimeinthecorridoras males
enteringthenarrowestcorridor(Table 2).
The speed of linear movementwas considerably
lowerformales in the 3 m wide corridorsystemthan
in the two narrowersystems.For instance,the pro-
gressrate(expressedas maximumdistancereachedin
thecorridordividedby timespentin corridor)in the
1m wide corridorwas on averagefourtimeshigher
thanin the3 m widecorridor,whereasthisparameter

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 25 Corridorwidth:3m
67 questions about corridordesign,for which all the
20
H. P. Andreassen, requiredelementsof properexperimental designs(i.e.
15
S. Halle &
R.A.Ims
10 + randomization
independent
oftreatment
measurements)
combinationsas to ensure
are satisfied,
has recently
been discussedby Inglis& Underwood(1992).
Our studylacks some statisticalrigourof a proper
25 Corridorwidth:1m experiment in thesensethatwe werenot able to have
201. more than one corridorsystem.To obtain proper
15.,
randomizationof treatmentcombinationswith the
10 + same sample size would have required45 corridor
5 F 171 systems.The present study should thereforebe
regardedas a thoroughlycontrolled,manipulative,
251+ observationalstudyratherthana properexperiment.
Corridorwidth:0-4m
20O. However,we are not able to imagineconfounding
15fl variables that could have produced the very pro-
nounced patternof our results.With respectto the
iof
101 most obvious confoundingfactorof the study,i.e.
time,our resultsare not compatiblewitha consistent
timebias, e.g. broughtabout by an accumulationof
0 60 120 180 240 300
scentthroughtime.Such a bias shouldhavegenerated
Fig.2. Frequencydistribution of maximumdistances
reached(20-mintervals)fromthereleasepoint(see Fig.1) a generaldecreaseor increasein movementestimates
forthethreetested widths.
corridor Blackbarsattheendof bothwithinand betweenthetreatments.
thecorridor givethenumber oftimesmalesdid notleave Our resultscorroboratethreeearlierstudiesshow-
theirreleasepatchor reachedtheoppositepatch,respec- ing that structuralaspects of corridorsaffectmove-
Arrows
tively. denotethemediansofthethreedistributions. mentratesin smallmammals(Lorenz& Barrett1990;
Merriam& Lanoue 1990; La Polla & Barrett1993).
However,our data do not lend supportto thenotion
did not differ betweenthe 1m and 0 4 m
significantly thatwidercorridorsnecessarilyconnecthabitatpat-
corridor(Table 2). chesmoreefficiently thannarrowcorridors(e.g. Har-
The footprint recordsindicatedan increaseduse of rison1992).Of threecorridorwidthstested,theinter-
corridoredges(edge counts;Table 2) withincreasing mediate width stimulatedthe highestfrequencyof
corridorwidth.The two ratios'edge counts/mid-run- long-distance movements. Similar results have
way counts' and 'cross-runwaycounts/mid-runway recentlybeen obtained for meadow voles Microtus
counts', whichboth may serveas indicesfor cross- pennsylvanicus (La Polla & Barrett1993)undermore
directionaluse of the corridor,had highestmean natural,butlesscontrolledconditions.In themeadow
values in the 3 m wide corridor(Table 2). This indi- vole study,therewereless movementsin 5 m thanin
cates that therewere more cross-directional move- 1m wide corridorsconnectinghabitatpatches with
mentsin thewidestcorridorthanin thetwonarrower free-ranging subpopulationsof voles. Whereas the
corridors. meadowvole corridorswereonly10m long(La Polla
& Barrett 1993) and probably supported mainly
within-homerange movements,the long-distance
Discussion
movementsin our 310m long corridorare probably
Many observationalstudieshave indicatedthatlinear morerepresentative forexploratorymovementsdur-
structuresin fragmentedlandscapes may play an ing dispersal(sensuStenseth& Lidicker1992). Typi-
importantrole as movementcorridorsand hence, cally exploratorymovements,e.g. duringdispersal
influencethe abundance and spatial structuring of events,take place on groundwhichis novel to the
fragmentedpopulations (e.g. Wegner & Merriam movinganimal(thisstudy),whilewithin-home-range
1979; Middleton & Merriam 1983; Hansson 1987; movements(the studyof La Polla & Barrett1993)
Szacki 1987;Verboom& van Apeldoorn1990;Zhang are normallybased on spatial memory(Ims 1994).
& Usher 1991; Fitzgibbon1993). However,veryfew However,therelationship betweencorridorwidthand
studieshave assessed movementratesof individuals movementrate was the same for the two Microtus
as functionsofqualitativeand quantitativeaspectsof species, despite the fact that theywere studied in
such corridors,whichin turncould serveas empiri- different settings,presumablyperformingdifferent
cally based guidelinesfor design of movementcor- movementtypes at different spatial scales (Wiens
ridorsforconservationpurposes.Preferentially, such 1989; Johnsonet al. 1992). This bears promisefor
guidelinesshould stemfromexperimental studiesin generalizations.The potential robustness of our
? 1996British
Ecological Society, which the causal relationship between corridor resultsis further emphasizedby thefactthatthegen-
JournalofApplied characteristics
and connectivitycould be established. eral relationshipbetweencorridorwidthand move-
Ecology,33, 63-70 The greatdifficulties
of conductingstudiesaddressing mentbehaviourin M. ceconomuswas notchangedby

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68 circumstances suchas simulatedpresenceofpredators
Optimalwidthof or competitorsin the corridor.The apparentunim-
movement portanceof thesetwo potentiallymodifyingfactors
corridors mighthave been due to a generalfailureto simulate
predators and competitorsadequately. However,
similarmethodsused in earlierstudies(althoughnot
conductedincorridors)producedclearresponses(Ims
1988,1990;Jedrzejewski et al. 1993;Nelson 1994).
The detailedobservationsmade duringthepresent
studysuggestseveral behaviouralmechanismsthat
may account for the variationin connectivity with
corridorwidth.The low efficiency of the narrowest
corridorin transferringvolesbetweenhabitatpatches
seemedto be caused by a behaviouralaversionbythe
voles to enterthecorridor(cf. the low entrancerate
in the 04 m corridor;Fig. 2). Moreover,voles that
enteredthe corridorspent less time in the narrow
and theintermediate corridorcomparedto thewidest
corridor(Table 2). For the intermediate widthcorri-
dor, however,the short residencytime probably
reflectsthehighlinearmovementrateratherthanany
aversionbehaviour(cf. 100% entranceratein the 1m
wide corridor;Fig. 2). The voles frequently reached
theoppositeend quicklywhentestedin the 1m wide
corridor.The low linearmovementspeedin thewidest
corridor,despitea relativelyhighentrancerate(83%),
may have been caused by a highfrequencyof cross-
directional(zig-zag) movements.This was indicated
by the highproportionateuse of cross-runways and
edgesin the3 m widecorridor(cf. theratio-variables
based on footprint counts;Table 2). Zig-zaggingmay
be a movementmode which is commonlyused by
animals in the absence of physicalorientationcues
(Bell 1991; Ims 1994). In movementcorridors,the
edgesare likelyto functionas themaincue facilitating
directionalmovements. Whileanimalsmovingin nar-
rowcorridorsmay be able to perceivetheedgeseven
fromtheinteriorof thecorridor,zig-zagmovements
may be necessaryin widerlinearstripsof habitatto
obtainfrequent visualcontactswiththeedges.In fact,
animalswillprobablynotperceivea linearpatchas a
movementcorridorabove a givenwidth,but as an
elongatedhabitat(see also La Polla & Barrett1993).
It has recently
beenemphasizedthatgeneralprinciples
in landscape ecology will most likelyemergefrom
detailed studies unravellingthe behaviouralmech-
anismsinvolvedin the movementecologyof species
(Johnsonet al. 1992;Wienset al. 1993;Ims 1994). In
particular,we believe that this argumentapplies to
the generationof principlesof movementcorridor
functionand design.
Mortalityratesof animalsin movementcorridors
have neverbeenestimated,althoughitis an extremely
importantfactordetermining whethercorridorsfunc-
tion as links or sinks (sensu Pulliam 1988) in the
dynamicsof fragmented populations(Fahrig& Mer-
? 1996British
EcologicalSociety, riam1985;Henein& Merriam1990).Still,onepremise
JournalofApplied underlying the presumptionthatwidercorridorsare
Ecology,33, 63-70 bettercorridorsis an inverserelationshipbetween
This content downloaded from 139.133.69.143 on Wed, 9 Jul 2014 08:43:02 AM
All use subject to JSTOR Terms and Conditions
mortality rateand corridorwidth.Movementsin nar-
rowcorridorsarebelievedto be moreriskythanmove-
mentsin wide corridorsdue to a higherinfluenceof
variousedge effects,such as predation(Simberloff&
Cox 1987; Noss 1987; Andren & Angelstam1988;
Henein & Merriam 1990; Saunders & Hobbs 1991;
Harrison 1992; Merriam & Saunders 1993; Paton
1994). The voles' reluctanceto enterthe narrowest
bya low entrancerate;
corridorin our study(reflected
27%), may reflectrisk-aversionbehaviour (McNa-
mara & Houston 1987). However, our resultsalso
pointto thepossibilityof a higheraccumulatedmor-
talityrisk in wide corridorssince voles spentmore
timein the widestcorridor,and since theyused the
corridoredges more intensively (cf. numberof edge
designedto mea-
counts;Table 2). Studiesspecifically
surehow mortality rateschangewithcorridorwidth,
will be an importantnextstep towarda knowledge
about whichcharacteristics may determinewhether
corridorsfunctionas links or sinks in a landscape
context.
Conclusion
We have shownthatthe efficiency of linearhabitats
as corridorsconnectingisolated habitat patches is
dependenton theirwidth,but in a differentmanner
thanoftensupposed.Furthermore, we have identified
what may be importantbehavioural mechanisms
underlyingmovementpatternsin linearhabitats.
Acknowledgments
OttarBj0rnstad,JorunFauske, Barbara Halle, Edda
Johannesen,Piotr Pawlak, Gytis Racius and Anna-
BarbaraUtelliparticipated duringthefieldwork.Bar-
bara Halle computerizedthe data. Ottar Bj0rnstad,
Gary Fry,LennartHansson,Karine Hertzberg,Har-
ald Steen and Nils Chr. Stensethprovidedvaluable
commentsto draftsof the manuscript.This studyis
part of the project 'The effectof habitatcorridors
on biodiversityin agriculturallandscapes' which is
supportedfinancially bytheResearchCouncilofNor-
way (NFR).
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