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
1996,33,
63-70

forrootvoles:
corridors
Optimalwidthofmovement
nottoo narrowand nottoo wide
HARRY
ROLF

P. ANDREASSEN,
ANKER
IMS

STEFAN

HALLE

and

N-0316Oslo,
DivisionofZoology,Department
of Oslo, PO Box 1050 Blindern,
ofBiology,University
Norway

Summary
1. The characteristics
of male root vole movementsas a functionof corridorwidth
weretestedin a 310m longhabitatcorridorconnectingtwohabitatpatches.Detailed
and recordingof
observationsof movementsweremade by meansof radiotelemetry
footprints.
in termsoftransference
2. The highestconnectivity,
rateofindividualsin thecorridor
ofthreecorridorwidthstested(3 m, 1m and
system,was observedin theintermediate
0 4 m).
of the narrowestcor3. The behaviouralmechanismbehindthe lowerconnectivity
ridorwas a reluctanceofvoles to enterit,whilelinearprogressin thewidestcorridor
of cross-directional
movements.
was hamperedby a highfrequency
4. The relationship
betweencorridorwidthand movementbehaviourwas unaffected
and predators.
by thesimulatedpresenceof competitors
5. Our resultschallengethe 'the-wider-the-better'
principleof movementcorridor
of the behaviouralmechanisms
design,and provideelementsforan understanding
themovementecologyof individualsin linearhabitats.
underlying
corridordesign,Microtus,movementecology.
Key-words:
connectivity,
JournalofAppliedEcology(1996) 33, 63-70

Plummer1993). As veryfewstudieshave been able

to obtaindata about animalmovementsin corridors
Habitatfragmentation
hasbeenrecognized
as a major (Hobbs 1992),theempiricalbasis forcorridordesign
etal. 1992;
threat
towildlife
populations
(Diamond1976;Gilpin is poor (Simberloff&Cox 1987;Simberloff
& Diamond1980;Higgs& Usher1980;Soule 1986; Mann & Plummer1993). For instance,theeffectsof
Lande 1988;Ims & Stenseth1989).As continuous structuralaspects of habitatcorridors,such as corhabitatsbecomefragmented,
an immediate
conse- ridorwidthand continuity,on movementrates are
becomes poorly known. Still, specificrecommendationson
of organisms
quenceis thatthemobility
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 forempiricalstudiesaddressquently,the viabilityof populations(Soule 1986; ing basic questionsregardingmovementecology of
Boyce1992).It has beenclaimedthatthenegative individuals in linear habitats, has recentlybeen
effects
of habitatfragmentation
can be reducedby stressed(see Hobbs 1992; Inglis& Underwood 1992;
Simberloff
et al. 1992;Lindenmayer& Nix 1993).
isolatedfragments
connecting
by narrowstripsof
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
definition
of movement
corridors
as linearhabitats detailedanalysis.We used corridorwidthas themain
formovements,
butnotforpermanent
settle- treatment
allowing
variable,because it is themostbasic strucoflinearhabitatsuponwhichpracment.
turalcharacteristic
The establishment
of movement
corridors
is cur- tical corridordesign inevitablyrequiresa decision.
rentlyimplemented
as an expensiveconservation Root voles were selectedas the studyorganismsas
world-wide
etal. 1992;Mann&
strategy
(Simberloff
theyhave proved to be veryusefulmodels forfield

Introduction

? 1996British
Ecological Society

63

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64
Optimalwidthof
movement
corridors

experiments
involvinghabitatmanipulations(Ims &
Stenseth1989; Ims et al. 1993; Wiens et al. 1993).
Moreover,althoughwe primarily
designedthisstudy
to serveas an empiricalmodel system(sensu Ims &
Stenseth1989) foraddressingbasic biologicalmechanisms that impingeon corridordesign(e.g. movementbehaviour),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 commonpresumptionthat widercorridorsnecessarilyare better
corridors(Noss 1987; Harrison 1992; Merriam &
Saunders 1993).

Methods
STUDY

PLOT

AND

MANIPULATIONS

The studytookplace at EvenstadField Station,southeastNorwayduringAugust-October1992.The movement 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,homogeneous meadow vegetation,knownto be preferred
habitatby root voles (Ims et al. 1993). An artificial
runwaysystem(Fig. 1) was establishedas 0 1 m wide
vegetation-free
pathsalong and acrossthecorridorto
in theotherwiseverydensegrass
facilitatemovements
carpet: one 310m long mid-runwayand cross-runwaysat 10-mintervals.Theserunwaystendedto direct

15m 30m

60m

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
in thecorridorat thestartof theexperienvironment
to thesituationlateron,
mentwas not verydifferent
some adult males were temporarilyreleased in the
corridora fewdays beforethefirstexperimental
trial
was executed.
We testedthreedifferent
corridorwidthsin comofpredatorsand combinationwithabsence/presence
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
in Table 1).
varioustreatments
Presenceof competitorsand predatorsweresimulated 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 thedecreasingwidthofthecorridor.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

lOOm

60m

i
Fence

1996 British
Ecological Society,
JournalofApplied
Ecology,33, 63-70

30m 15m

Iiu

Barriercage

Vegetation - - - Runwaysystem
Release point A Trackingplate

meadow
vegetation
ofdense,homogeneous
consisted
patches
andthecorridor
plot.Thetwohabitat
Fig.1. Designofthestudy
sectionof
ofbareground.Theenlarged
(whiteareainsidethefence)consisted
(shadedarea).Theareaoutsidethecorridor
platesinthe
oftracking
system,
releasepointandthepositioning
runway
thecorridor
(inset)showsthedesignoftheartificial
(crossandacrossthecorridor
pathsalong(mid-runway)
as 0 1mvegetation-free
wereestablished
3 mwidecorridor.
Runways
forthe0-4m
(exceptin thecross-runways
Tracking
plateswereplacedin bothtypesofrunways
runways
at 10-mintervals).
corridor)
as wellas alongtheedgesofthecorridor.

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Table 1. Experimental
protocol:Competitor,otheradultmalesin thebarriercages (Fig. 1); Predator,foxscatsin thebarrier
65
male rootvoles wereused in each of the9 treatment
types
H. P. Andreassen, cages; Control,emptybarriercages. Ten different
S. Halle &
Numberof samplerecords
R. A. Ims
Treatment

Period*

Radiotrackingt Footprintst

Width,3 m
Control
Predator
Competitor

1-9 Aug
10-16 Aug
18-26 Aug

10
10
9

5
2
5

Width,1m
Competitor
Control
Predator

26 Aug-i Sep
2-7 Sep
8-15 Sep

10
10
10

5
5
4

Width,0-4m
Predator
Competitor
Control

19-23 Sep
24-8 Sep
29 Sept-3 Oct

10
9
10

5
4
4

* The variablelengthsof the experimental

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

to affectspace use in microtines(Jedrzejewski

et al.
1993). For the 'control'treatment,
the barriercages
were leftempty.Beforethe onset of each treatment,
the barriercages were carefullywashed to remove
remainingscent fromthe previoustreatment.Furthermore,the cages were always placed on plastic
sheetsto preventthesoil beneaththecages becoming
contaminatedby scent.
TREATMENT
STUDY

1996British
Ecological Society,
JournalofApplied
Ecology,33, 63-70

AND

MONITORING

OF THE

ANIMALS

On each day,tworadiocollared(Biotrack,SS-2 transmitters),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.
inthecorridorweremoniLongitudinalmovements
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-tracking
positionsobtainedpermale wereused
to estimatethe distancemoved,the speed of movementand thetimespentin thecorridorforeach male
(forfurther
oflongitudinal
description
movement
parameters,see footnotesto Table 2).
use of thecorridorwas registered
Cross-directional
bythefootprint
recordingtechniqueofKing & Edgar

(1977). Devices for recordingfootprints(tracking

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.Radiotrackingshowedthatsuchcrossing occurredin at least 29% of the trials.Due to
the generallow sample size of footprintrecordsand
missingdata, leadingto unbalanceddesigns(Table 1),
statisticaltestsofcompetitor/predator
treatment
were
not appliedto thefootprint
data.
The males used in thisstudywerelaboratoryborn
F2-F4 generationdescendantsof animalscaughtin
Pasvik, north-eastNorway. We chose to use laboratory-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 shorttermbehaviouralchanges due to reproductivehormone 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
Optimalwidthof
movement
corridors

Table 2. The effectof corridorwidthon movementand corridoruse parameters.Data are lumpedforthe treatment
factor
ofcompetitor/predator.
absence/presence
Values are presentedas mean +-standard errorof themean
Parameters

Corridorwidth
3m

Im

0 4m

Longitudinalmovements*
Maximumdistancereached(m)
Cumulativedistancetraversed(m)

75 + 16
163 + 33

205 + 22
440 + 57

35 + 15
51 + 21

Movementspeed/efficiencyt
Time spentin corridor(h)
Maximumspeed (m/h)
Averagespeed (m/h)
Progressrate(m/h)

79+
61 +
21 +
10 +

59 +
160 +
76 +
40 +

3-9 +
93 +
51 +
35 +

Cross-directional
movementst(footprint
counts)
Edge counts
ratio
Edge/mid-runway
ratio
Cross-runway/mid-runway

0't
11
4
2

1 42 + 0 51
0 23 + 0-09
1 56 + 0 31

04
19
11
10

0 50 + 0 20
0 03 + 0-01
0-36+ 0 05

11
24
13
10

0 21 + 0 11
0 09 + 0-07

* 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.
use parametersare based on footprint
countrecords(Table 1) fromthefootprint
t Cross-directional
platessituatedalong
thecorridoredges (edge counts),in themid-runway
and thecross-runways
(Fig. 1). No estimateof theratiovariable(crossis givenforthe0 4 m corridor,sincefootprints
werenot recordedin thecross-runways
forthiscorridor
runway/mid-runway),
width.

experiment.All males used were sexually mature,

adult animals(> 50 g and scrotaltestes),and thusof
adequate size forthe2 5 g radiocollars.
different
Twenty-six
individualswereavailable 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.Genermalesweretestedpertreatment
ally,10 different
combination(Table 1). Males used in morethanone treatment were released in alternatingpatches with the
longestpossible inter-trial
intervalto avoid habituation 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. scentaccumulation (see also Discussion) were importantin this
crossed
study.Males partneredin trialsinfrequently
each otherin the corridor(29% of all trials) so it
is unlikelythatdirectinterference
betweenthe freetheresults.Hence,we findit
rangingmalesinfluenced
justifiableto treateach trialpermale as thestatistical
unit(cf.Table 1) in theanalyses.

Results
? 1996British
EcologicalSociety,
JournalofApplied
Ecology,33, 63-70

Corridorwidthhad a substantialeffecton all movementparameters(Table 2) (two-wayANOVAS, all partialr2> 0 25, P < 0 001). Presence/absence
of predatorsand competitors
wererelatively
for
unimportant

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 intermediatewidthcorridorwere on average threeand
six timeslongerthan in the widerand the narrower
In the intermediate
corridor
corridors,respectively.
morethanone-thirdofthevolesreachedtheopposite
end, while this rarelyhappened in the wideror the
narrowercorridor(Fig. 2).
The low successof the narrowestcorridorin connectingthe 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 enteringthewidestcorridorspenttwiceas 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 progressrate(expressedas maximumdistancereachedin
thecorridordividedby timespentin corridor)in the
1m wide corridorwas on averagefourtimeshigher
thanin the3 m widecorridor,whereasthisparameter

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67
H. P. Andreassen,
S. Halle &
R.A.Ims

25
20
15

10

Corridorwidth:3m

25

Corridorwidth:1m

201.
15.,

10
5

F 171

251+

Corridorwidth:0-4m

20O.
15fl
101
iof

0
300
60
240
120
180
of maximumdistances
Fig.2. Frequencydistribution
reached(20-mintervals)
fromthereleasepoint(see Fig.1)
widths.
Blackbarsattheendof
forthethree
tested
corridor
oftimesmalesdid notleave
thecorridor
givethenumber
theirreleasepatchor reachedtheoppositepatch,respecArrows
ofthethree
denotethemedians
distributions.
tively.

did not differ

significantly
betweenthe 1m and 0 4 m
corridor(Table 2).
The footprint
recordsindicatedan increaseduse of
corridoredges(edge counts;Table 2) withincreasing
corridorwidth.The two ratios'edge counts/mid-runway counts' and 'cross-runwaycounts/mid-runway
counts', whichboth may serveas indicesfor crossdirectionaluse of the corridor,had highestmean
values in the 3 m wide corridor(Table 2). This indimovecates that therewere more cross-directional
mentsin thewidestcorridorthanin thetwonarrower
corridors.

Discussion

? 1996British
Ecological Society,
JournalofApplied
Ecology,33, 63-70

Many observationalstudieshave indicatedthatlinear

structuresin fragmentedlandscapes may play an
importantrole as movementcorridorsand hence,
of
influencethe abundance and spatial structuring
fragmentedpopulations (e.g. Wegner & Merriam
1979; Middleton & Merriam 1983; Hansson 1987;
Szacki 1987;Verboom& van Apeldoorn1990;Zhang
& Usher 1991; Fitzgibbon1993). However,veryfew
studieshave assessed movementratesof individuals
as functions
ofqualitativeand quantitativeaspectsof
such corridors,whichin turncould serveas empirically based guidelinesfor design of movementcorridorsforconservationpurposes.Preferentially,
such
guidelinesshould stemfromexperimental
studiesin
which the causal relationship between corridor
characteristics
and connectivity
could be established.
The greatdifficulties
of conductingstudiesaddressing

questions about corridordesign,for which all the

requiredelementsof properexperimental
designs(i.e.
randomization
oftreatment
combinationsas to ensure
independent
measurements)
are satisfied,
has recently
been discussedby Inglis& Underwood(1992).
Our studylacks some statisticalrigourof a proper
experiment
in thesensethatwe werenot able to have
more than one corridorsystem.To obtain proper
randomizationof treatmentcombinationswith the
same sample size would have required45 corridor
systems.The present study should thereforebe
regardedas a thoroughlycontrolled,manipulative,
observationalstudyratherthana properexperiment.
However,we are not able to imagineconfounding
variables that could have produced the very pronounced patternof our results.With respectto the
most obvious confoundingfactorof the study,i.e.
time,our resultsare not compatiblewitha consistent
timebias, e.g. broughtabout by an accumulationof
scentthroughtime.Such a bias shouldhavegenerated
a generaldecreaseor increasein movementestimates
bothwithinand betweenthetreatments.
Our resultscorroboratethreeearlierstudiesshowing that structuralaspects of corridorsaffectmovementratesin smallmammals(Lorenz& Barrett1990;
Merriam& Lanoue 1990; La Polla & Barrett1993).
However,our data do not lend supportto thenotion
thatwidercorridorsnecessarilyconnecthabitatpatchesmoreefficiently
thannarrowcorridors(e.g. Harrison1992).Of threecorridorwidthstested,theintermediate width stimulatedthe highestfrequencyof
long-distance movements. Similar results have
recentlybeen obtained for meadow voles Microtus
pennsylvanicus
(La Polla & Barrett1993)undermore
natural,butlesscontrolledconditions.In themeadow
vole study,therewereless movementsin 5 m thanin
1m wide corridorsconnectinghabitatpatches with
free-ranging
subpopulationsof voles. Whereas the
meadowvole corridorswereonly10m long(La Polla
& Barrett 1993) and probably supported mainly
within-homerange movements,the long-distance
movementsin our 310m long corridorare probably
forexploratorymovementsdurmorerepresentative
ing dispersal(sensuStenseth& Lidicker1992). Typically exploratorymovements,e.g. duringdispersal
events,take place on groundwhichis novel to the
movinganimal(thisstudy),whilewithin-home-range
movements(the studyof La Polla & Barrett1993)
are normallybased on spatial memory(Ims 1994).
betweencorridorwidthand
However,therelationship
movementrate was the same for the two Microtus
species, despite the fact that theywere studied in
different
settings,presumablyperformingdifferent
movementtypes at different
spatial scales (Wiens
1989; Johnsonet al. 1992). This bears promisefor
generalizations.The potential robustness of our
resultsis further
emphasizedby thefactthatthegeneral relationshipbetweencorridorwidthand movementbehaviourin M. ceconomuswas notchangedby

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68
Optimalwidthof
movement
corridors

? 1996British
EcologicalSociety,
JournalofApplied
Ecology,33, 63-70

circumstances
suchas simulatedpresenceofpredators
or competitorsin the corridor.The apparentunimportanceof thesetwo potentiallymodifyingfactors
mighthave been due to a generalfailureto simulate
predators and competitorsadequately. However,
similarmethodsused in earlierstudies(althoughnot
conductedincorridors)producedclearresponses(Ims
et al. 1993;Nelson 1994).
1988,1990;Jedrzejewski
The detailedobservationsmade duringthepresent
studysuggestseveral behaviouralmechanismsthat
with
may account for the variationin connectivity
of the narrowest
corridorwidth.The low efficiency
volesbetweenhabitatpatches
corridorin transferring
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
widthcorricorridor(Table 2). For the intermediate
dor, however,the short residencytime probably
reflects
thehighlinearmovementrateratherthanany
aversionbehaviour(cf. 100% entranceratein the 1m
reached
wide corridor;Fig. 2). The voles frequently
theoppositeend quicklywhentestedin the 1m wide
corridor.The low linearmovementspeedin thewidest
highentrancerate(83%),
corridor,despitea relatively
may have been caused by a highfrequencyof crossdirectional(zig-zag) movements.This was indicated
and
by the highproportionateuse of cross-runways
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
Whileanimalsmovingin nardirectionalmovements.
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 behaviouralmechanismsinvolvedin 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
whethercorridorsfuncimportantfactordetermining
tion as links or sinks (sensu Pulliam 1988) in the
dynamicsof fragmented
populations(Fahrig& Merriam1985;Henein& Merriam1990).Still,onepremise
underlying
the presumptionthatwidercorridorsare
bettercorridorsis an inverserelationshipbetween

mortality
rateand corridorwidth.Movementsin narrowcorridorsarebelievedto be moreriskythanmovementsin 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 (McNamara & Houston 1987). However, our resultsalso
pointto thepossibilityof a higheraccumulatedmortalityrisk in wide corridorssince voles spentmore
timein the widestcorridor,and since theyused the
(cf. numberof edge
corridoredges more intensively
designedto meacounts;Table 2). Studiesspecifically
surehow mortality
rateschangewithcorridorwidth,
will be an importantnextstep towarda knowledge
may determinewhether
about whichcharacteristics
corridorsfunctionas links or sinks in a landscape
context.

Conclusion
We have shownthatthe efficiency
of linearhabitats
as corridorsconnectingisolated habitat patches is
manner
dependenton theirwidth,but in a different
thanoftensupposed.Furthermore,
we have identified
what may be importantbehavioural mechanisms
movementpatternsin linearhabitats.
underlying

Acknowledgments
OttarBj0rnstad,JorunFauske, Barbara Halle, Edda
Johannesen,Piotr Pawlak, Gytis Racius and AnnaBarbaraUtelliparticipated
duringthefieldwork.Barbara Halle computerizedthe data. Ottar Bj0rnstad,
Gary Fry,LennartHansson,Karine Hertzberg,Harald Steen and Nils Chr. Stensethprovidedvaluable
commentsto draftsof the manuscript.This studyis
part of the project 'The effectof habitatcorridors
on biodiversityin agriculturallandscapes' which is
supportedfinancially
bytheResearchCouncilofNorway (NFR).

References
Andreassen.,Ims, R.A, Stenseth,N.C. & Yoccoz, N.G.
(1993) Investigating
space use by meansof radiotelemetry
and othermethods:a methodologicalguide. The Biology
of Lemmings(eds N. C. Stenseth& R. A. Ims), pp. 589618. AcademicPress,London.
Andren,H. & Angelstam,P. (1988) Elevatedpredationrates
in habitatislands:experimental
as an edgeeffect
evidence.
Ecology,69,344-547.
Bell, W.J. (1991) SearchingBehaviour.Chapman & Hall,
London.

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

Bennett,A.F. (1990) Habitatcorridorsand theconservation

69
of small mammalsin a fragmented
forestenvironment.
H. P. Andreassen,
LandscapeEcology,4, 109-122.
S. Halle &
Boyce, M.S. (1992) Population viabilityanalysis. Annual
R.A.Ims
Reviewsin Ecologyand Systematics,
23, 481-506.

? 1996British
Ecological Society,
JournalofApplied
Ecology,33, 63-70

Burkey,T. (1989) Extinctionin naturereserves.The effect

of
fragmentation
and the importanceof migrationbetween
reservefragments.
Oikos,55, 75-81.
Diamond, J.M. (1976) Island biogeography and conservation:Strategyand limitations.Science, 193, 10271029.
Fahrig,L. & Merriam,G. (1985) Habitatpatchconnectivity
and populationsurvival.Ecology,66, 1762-1768.
Fahrig,L. & Paleheimo,J. (1988) Effectof spatial arrangementofhabitatpatcheson local populationsize. Ecology,
69, 468-475.
Fitzgibbon,C.D. (1993) The distributionof greysquirrel
dreysin farmwoodland: the influenceof wood area, isolation and management.Journalof AppliedEcology,30,
736-742.
Gilpin,M.E. & Diamond, J. (1980) Subdivisionof nature
reservesand themaintenanceof speciesdiversity.
Nature,
285, 567-568.
Hansson,L. (1987) Dispersalroutesofsmallmammalsat an
abandonedfieldin centralSweden.HolarcticEcology,10,
154-159.
Harris, L.D. (1984) The FragmentedForest: Island BiographicTheoryAnd The Preservation
Of BioticDiversity.
of Chicago Press,Chicago.
University
corHarrison,R.L. (1992) Towards a theoryof inter-refuge
ridordesign.Conservation
Biology,6, 293-295.
Henein,K.M. & Merriam,G. (1990) The elementof connectivitywhere corridorquality is variable. Landscape
Ecology,4, 157-170.
Higgs,A.J.& Usher,M.B. (1980) Should naturereservesbe
largeor small?Nature,285, 568-569.
Hobbs, R.J. (1992) The role of corridorsin conservation:
Solutionor bandwagon?TrendsinEcologyand Evolution,
7, 389-392.
Ims, R.A. (1988) Spatial clumpingof sexually receptive
femalesinducesspace sharingamongmale voles. Nature,
335, 541-543.
on dispersaland
Ims,R.A. (1989) Kinshipand origineffects
space sharingin Clethrionomys
rufocanus.Ecology,70,
607-616.
Ims, R.A. (1990) Mate detection success of male Clein relationto thespatialdistribution
thrionomys
rufocanus
of sexuallyreceptivefemales.Evolutionary
Ecology,4, 5761.
Ims, R.A. (1995) Movementpatternsin relationto spatial
structures.Mosaic Landscapes and Ecological Processes
(eds L. Hansson, L. Fahrig& G. Merriam).Chapman &
Hall, London.
fall.
Ims,R.A. & Stenseth,N.C. (1989) Divided thefruitflies
Nature,342, 21-22.
Ims, R.A., Rolstad,J. & Wegge,P. (1993) Predictingspace
use responsesto habitatfragmentation:
can volesMicrotus
oeconomusserveas an experimental
model system(EMS)
forcapercailliegrouse Tetraourogallusin boreal forest.?
BiologicalConservation,
63, 261-268.
Inglis,G. & Underwood,A.J. (1992) Commentson some
on thebiologicalimportdesignsproposedforexperiments
ance ofcorridors.Conservation
Biology,6, 581-586.
Jedrzejewski,
W., Rychlic,L. & Jedrzelewska,B. (1993)
Responses of bank voles to odours of seven species of
data and theirrelevanceto natural
predators,experimental
predator-volerelationships.
Qikos,68, 251-257.
Johnson,A.R., Milne, B.T. & Wiens,J.A. (1992) Diffusion
in fractallandscapes:simulationsand experimental
studies
of tenebrionid
beetlemovements.Ecology,73, 1968-1983.
Kawata, M. (1989) Growthand dispersaltimingin malered-

backed vole Clethrionomys

rufocanusbedfordiae.Oikos,
54, 220-226.
King, C.M. & Edgar, R.I. (1977) Techniquesfortrapping
and trackingstoats (Mustela erminea);a review,and a
new system.New Zealand Journalof Zoology, 4, 193212.
La Polla, V.N. & Barrett,G.W. (1993) Effectsof corridor
widthand presenceon the populationdynamicsof the
meadow vole (Microtuspennsylvanicus).
LandscapeEcology,8, 25-37.
Lande, R. (1988) Genetics and demographyin biological
conservation.Science,241, 1455-1460.
D.B. & Nix, H.A. (1993) Ecologicalprinciples
Lindenmayer,
forthedesignof corridors.Conservation
Biology,7, 627630.
Lorenz,G.C. & Barrett,G.W. (1990) Influenceof simulated
landscapecorridorson house mouse (Mus musculus)dispersal.AmericanMidlandNaturalist,123,348-356.
Mann, C.C. & Plummer,M. (1993) The highcosts of biodiversity.Science,160, 1868-1871.
McNamara, J.M. & Houston, A.I. (1987) Starvationand
predationas a factorslimitingpopulationsize. Ecology,
68, 1515-1519.
Merriam,G. & Lanoue, A. (1990) Corridoruse by small
mammals:fieldmeasurements
forthreeexperimental
types
of Peromyscusleucopus.LandscapeEcology,4, 123-131.
Merriam,G. & Saunders,D. (1993) Corridorsin restoration
of fragmented
3: Reconlandscapes.NatureConservation
struction
Of Fragmented
Ecosystems(eds D. Saunders,R.
J.Hobbs & P. Erlich).SurreyBeatty& Sons, Australia.
Middleton,J. & Merriam,G. (1983) Distributionof woodland speciesin farmlandwoods. Journalof AppliedEcology,20, 625-644.
Nelson,J.(1994) Spacingbehaviour,
matingsystem,
and indisuccessin malefieldvoles.PhD thesis,
vidualreproductive
of Lund, Sweden.
University
Noss, R. (1987) Corridorsin real landscapes: a replyto
and Cox. Conservation
Simberloff
Biology,1, 159-164.
Paton,P.W. (1994) The effect
ofedgeon aviannestsuccesses:
How strongis theevidence?Conservation
Biology,8, 1726.
Pulliam,H.R. (1988) Sources,sinks and population regulation.AmericanNaturalist,132,652-661.
Saunders, D.A. & Hobbs, R.J. (eds)(1991) Nature Conservation2: The Role Of Corridors.SurreyBeattyand
Sons, Australia.
Simberloff,
D., Farr,J.A.,Cox, J.& Mehlman,D.W. (1992)
Movement corridors: conservationbargains or poor
investment.
Conservation
Biology,6, 493-504.
D. & Cox, J. (1987) Consequencesand costs of
Simberloff,
conservationcorridors.Conservation
Biology,1, 63-71.
Soule, M. (1986) ConservationBiology. Sinauer, Massachusetts.
Stamps,J.A, Buechner,M & Krishnan,V.V. (1987a) The
effects
of edge permeability
and habitatgeometry
on emigrationfrompatchesofhabitat.AmericanNaturalist,129,
533-552.
Stamps,J.A, Buechner,M & Krishnan,V.V. (1987b) The
effectof habitat geometryon territorialdefencecosts,
intruderpressure in bounded populations. American
Zoologist,27, 307-325.
Stenseth,N.C. & Lidicker,W.Z. (1992) AnimalDispersal.
Small Mammalsas a Model. Chapman& Hall, London.
Szacki,J.(1987) Ecologicalcorridorsas a factordetermining
thestructure
and organisationof a bank vole population.
Acta theriologica,
32, 31-44.
van Apeldoorn,R., Hollander,H., Nieuwenhuizen,W. &
Vliet,F. van der(1992) De Noordsewoelmuisin hetDeltagebied.Is er een relatietussenhabitatfragmentatie
en concurrentieop de schaal van het landschap?Landschap,9,
189-202.

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

70

Optimalwidthof
movement
corridors

Verboom,B. & van Apeldoorn,R. (1990) Effectof habitat

fragmentation
on the red squirrel,Sciurus vulgarisL.
LandscapeEcology,4, 171-176.
Wegner,J.F. & Merriam,G. (1979) Movementof birdsand
small mammalsbetweenwood and adjoining farmland
habitats.JournalofAppliedEcology,16, 349-358.
Wiens,J.A.(1989) Spatialscalinginlandscapeecology.FunctionalEcology,3, 386-397.
Wiens, J.A., Stenseth,N.C., Horne, B. Van & Ims, R.A.
(1993) Ecological mechanismsand landscape ecology.
Oikos,66, 369-380.

Wilson,E.O. & Willis,E.O. (1975) Applied biogeogral

Ecologyand theEvolutionofCommunities
(eds M. L. C
& J.M. Diamond),pp. 522-536.HarvardUniversity
PI
Massachusetts.
Zhang,Z. & Usher,M.B. (1991) Dispersalofwood mice,
bank volesin an agricultural
landscape.Acta theriolog
36, 239-245.

Received27 April1994; revisionreceived21 October199<

? 1996British
Ecological Society,
JournalofApplied
Ecology,33, 63-70

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