Effects of Habitat Fragmentation on Biodiversity

Author(s): Lenore Fahrig

Source: Annual Review of Ecology, Evolution, and Systematics, Vol. 34 (2003), pp. 487-515
Published by: Annual Reviews
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 Annu. Rev. Ecol. Evol. Syst. 2003. 34:487-515
doi: 10.1146/132419
Copyright@ 2003 by AnnualReviews. All rights reserved
First publishedonline as a Review in Advance on August 14, 2003

OFHABITAT
EFFECTS FRAGMENTATION
ON
BIODIVERSITY

LenoreFahrig
Instituteof Biology,CarletonUniversity,
Ottawa-Carleton Ottawa,Ontario,
CanadaK1S5B6;email:LenorelFahrig @carleton.ca

Key Words habitatloss, landscapescale,habitatconfiguration,

patchsize, patch
isolation,extinctionthreshold,landscapecomplementation
0 Abstract The literatureon effects of habitatfragmentationon biodiversityis
huge. It is also very diverse,with differentauthorsmeasuringfragmentation in dif-
ferentways and,as a consequence,drawingdifferentconclusionsregardingboththe
magnitudeand directionof its effects. Habitatfragmentation is usuallydefinedas a
landscape-scaleprocessinvolvingboth habitatloss and the breakingapartof habi-
tat. Resultsof empiricalstudiesof habitatfragmentationare often difficultto inter-
pretbecause(a) manyresearchersmeasurefragmentation at the patchscale, not the
landscape scale and (b) most researchersmeasure fragmentation in ways thatdo not
distinguishbetweenhabitatloss and habitatfragmentation per se, i.e., the breaking
apartof habitataftercontrollingfor habitatloss. Empiricalstudiesto date suggest
thathabitatloss has large,consistentlynegativeeffects on biodiversity.Habitatfrag-
mentationper se has muchweakereffects on biodiversitythat are at least as likely
to be positive as negative.Therefore,to correctlyinterpretthe influenceof habitat
fragmentation on biodiversity,the effects of these two componentsof fragmentation
mustbe measuredindependently.More studiesof the independenteffects of habitat
loss andfragmentation per se areneededto determinethe factorsthatlead to positive
versus negativeeffects of fragmentationper se. I suggest that the term "fragmen-
tation"shouldbe reservedfor the breakingapartof habitat,independentof habitat
loss.

INTRODUCTION

A recentsearchof the CambridgeScientificAbstractsdatabaserevealedover 1600

articlescontainingthe phrase"habitatfragmentation."The task of reviewingthis
literatureis dauntingnot only because of its size but also because differentauthors
use differentdefinitionsof habitatfragmentation,andthey measurefragmentation
in differentways and at differentspatialscales.
This diversity of definitions of habitatfragmentationcan be readily seen in
the titles of some articles. For example, "Impactsof habitatfragmentationand

1543-592X/03/1215-0487$14.00 487

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488 FAHRIG

patchsize..." (Collingham& Huntly2000) suggeststhat habitatfragmenta-

tionandpatchsize aretwo differentthings.However,otherauthorsactuallyuse
patchsize to measurehabitatfragmentation (e.g.,Golden& Crist2000,Hovel&
Lipicus2001). "The effects of forest fragmentation and isolation..." (Good-
man& Rakotodravony 2000)suggests that forest fragmentation andisolationare
in
different, contrastto authors who use forest isolationas a measureof for-
est fragmentation(e.g., Mossman & Waser 2001, Rukke 2000). "Effectof land
cover,habitatfragmentation,and.. ." (Laakkonenet al. 2001) contrastswith many
authorswho equatelandscapefragmentation with landcover(e.g., Carlson&
Hartman2001; Fuller2001; Gibbs 1998, 2001; Golden& Crist2000; Hargis
et al. 1999;Robinsonet al. 1995;Summerville & Crist2001;Virg6s2001)."The
influenceof forestfragmentation andlandscapepattern..."(Hargiset al. 1999)
contrastswith researchers who definefragmentation as an aspectof landscape
pattern(e.g.,Wolff et al. 1997, Trzcinski et al. 1999). As a finalexample,"Effects
of experimental habitatfragmentation andconnectivity..."(Ims& Andreassen
1999)suggeststhathabitatfragmentation andconnectivitycanbe examinedin-
dependently, whereas some researchers actuallydefinefragmentation as "a dis-
in
ruption landscapeconnectivity" (With et al. 1997; see also Young& Jarvis
2001).
My goal in this reviewis to discussthe information availableon the effects
of habitatfragmentation on biodiversity.TomeetthisobjectiveI firstneedto ex-
aminethe differentways in whichhabitatfragmentation is conceptualized and
measured.Of course,the conceptof biodiversityis probablyat least as wide-
rangingas the conceptof habitatfragmentation. However,I do not deal with
the issues surrounding the conceptof biodiversity. Instead,I includeany eco-
logical responsevariablethatis or can be relatedto biologicaldiversity(see
Table1).
To determinecurrentusage of the termhabitatfragmentation, I conducted
a searchof the CambridgeScientificAbstracts(BiologicalSciences)database
on 11 April2002 for paperscontainingeither"habitatfragmentation," "forest
or
fragmentation,""landscape fragmentation" in the titleof the paper.I reviewed
in detailthemostrecent100resultingpapers,irrespective of thejournalin which
theyappeared. I limited this search to paperscontaining"fragmentation" in the
titleto ensurethatmy sampleincludedonlypapersthataredirectlyon thesubject
of habitatfragmentation. Theresultsaresummarized in Table1.
I then surveyedthe broaderecologicalliteratureto ask the following:How
strongaretheeffectsof habitatfragmentation on biodiversity, andaretheeffects
negativeor positive?Habitatfragmentation is generallythoughtto havea large,
negativeeffectonbiodiversity andis therefore widelyviewedasanaspectof habitat
degradation (Haila2002).However,as I show,thisconclusionis generallyvalid
onlyforconceptualizations of fragmentation thatareinseparable fromhabitatloss.
Otherways of conceptualizinghabitatfragmentationlead to otherconclusions. I
end the paperwith recommendations.

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 EFFECTSOF HABITATFRAGMENTATION 489

the in
in more

Numbers
(3) 3
Population
growth 1 0 1 1 1 0 0 1 1 1 1 patches contain
all may
variable.
across study
(4) 3
Movement/
dispersal 2 1 0 0 1 0 0 0 1 3 0
each
use summed
(biodiversity)
or
because
(5) 3
Individual
habitat 3 0 2 1 0 2 0 3 2 2 1 100
response
to
(averaged
andadd
scale not
scale
(8) 3
Extinction/
turnover 5 0 1 3 1 0 0 1 0 8 0 or do

variables landscape rows

variable
the and
at
(10) 7 5
Species
interactions 3 4 2 2 0 1 0 4 3 3 or
(response)
Columns
patch)
(fragmentation)
(12) 3 8 each variable.
Genetic
variability 6 0 0 0 2 0 4 7 3 2
Biodiversity for
that
predictor
of
using
1
(15) 11 9
measures
Fitness
2 2 2 2 0 10 6 4 5
(individually landscape.
patch. 100)
studies* (of variable.
scale single combination
single
1 1 2 7 a a
(26) 20 13 11 3 0
Presence/
absence 16 4 6 papers
patch given
fromfrom of
the the
biodiversity
fragmentation 1 1 2 1 9 studies. one
(28) 21
Richness/
diversity 17 7 5 14 7 6 number
either studied
at information
information than
thattotal
recent corridor
the more
and
100 papers
areand
2 3 3 0 13 17 7 10 measured represents
represents
(35) 26 21 14 11
Abundance/
density of
of be
studies
names
(8) can
analysis variable
analysis
(60) numbers
(10) that the the
Summary(predictor) (35) (28) (37) in in thevariable
(7) connectivityare
(4)
1 (63) patches only(42)scaled variables point after
point
connectivityb landscape both
loss/amountof fragmentation
quality (21)
(22)
sizea isolationa shapea scalec and datadataentries
one
scales
TABLE Fragmentation
variables
PatchHabitat
PatchEdgeaNumber Patch
Structural
Matrix Qualitative
Patch Patch aPredictor
Landscape landscape).
bIncludes
cEach *Table
dEach than
parentheses

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490 FAHRIG

AND MEASUREMENT
CONCEPTUALIZATION OF
HABITATFRAGMENTATION

Fragmentationas Process
Habitat fragmentation is oftendefined asaprocessduring which"alargeexpanse
of habitatis transformed intoa number of smallerpatchesof smaller totalarea,
isolatedfromeachotherbyamatrix ofhabitats unliketheoriginal"(Wilcove etal.
1986)(Figure1).By thisdefinition, a landscape canbequalitatively categorized
as eithercontinuous (containing continuous habitat)or fragmented, wherethe
fragmented landscape represents the endpoint of the of
process fragmentation.
Manystudiesof theeffectof habitat fragmentationon biodiversity conform
to thisdefinition some of
bycomparing aspect(s) biodiversity at"reference" sites
withina continuous landscape to the same of
aspect(s) biodiversity atsiteswithin
a fragmented landscape (e.g.,Bowers&Dooley1999,Cascante et al.2002,Diaz
et al.2000,Groppe et al.2001,Laurance et al.2001,MacNally&Brown2001,
Mahan& Yahner 1999,Morato2001,Mossman & Waser2001,Renjifo1999,
Walters et al. 1999).Frommysampleof 100recentstudies, 28%conducted such
comparisons of continuous versusfragmented landscapes(Table1).Inthesestud-
ies, thecontinuous landscape represents a landscape beforefragmentation (time
1 in Figure1) andthefragmented landscape representsa landscape following
fragmentation (time2 ortime3 inFigure1).
Although thisapproach conforms tothedefinitionoffragmentation asaprocess,
it has two inherentweaknesses.First,becausehabitatfragmentationis a landscape-
scale process (McGarigal& Cushman2002), the sample size in such studies, for
questions about the effects of habitatfragmentationon biodiversity,is typically

1 2 3

time
Figure 1 The processof habitatfragmentation, where"alargeexpanseof habitatis
transformedinto a numberof smallerpatches of smallertotalarea,isolatedfromeach
otherby a matrixof habitatsunlikethe original"(Wilcoveet al. 1986). Black areas
representhabitatandwhiteareasrepresentmatrix.

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 OFHABITAT
EFFECTS FRAGMENTATION 491

onlytwo,i.e.,onecontinuous landscape andonefragmented landscape. Withsucha

design,inferencesabout the effectsof fragmentationareweak. Apparent effectsof
could
fragmentation easily be due to other differences
between the landscapes.For
Mac
example, Nally et al. (2000) found consistent
vegetation differencesbetween
fragments andreferencesitesandconcludedthatapparent effectsof fragmentation
onbirdscouldbeduetopreexisting habitatdifferencesbetweenthetwolandscapes.
Second,thischaracterization of habitatfragmentation is strictlyqualitative,
i.e.,
eachlandscapecanbe in only one of two states,continuousor fragmented. This
designdoesnotpermitoneto studytherelationship betweenthedegreeof habitat
fragmentation andthe magnitudeof the biodiversityresponse.Quantifyingthe
degreeof fragmentation requiresmeasuringthe patternof habitaton the land-
scape.The diversityof approaches in the fragmentation literaturearisesmainly
fromdifferencesamongresearchers in how theyquantifyhabitatfragmentation.
Thesedifferenceshavesignificantimplicationsfor conclusionsaboutthe effects
of fragmentationon biodiversity.

Fragmentationas Pattern:QuantitativeConceptualizations
Thedefinitionof habitatfragmentation aboveimpliesfoureffectsof theprocessof
fragmentation on habitatpattern:(a) reductionin habitatamount,(b) increasein
numberof habitatpatches,(c) decreasein sizesof habitatpatches,and(d)increase
in isolationof patches.These foureffects formthe basis of most quantitative
measuresof habitatfragmentation. However,fragmentation measuresvarywidely;
someincludeonlyoneeffect(e.g.,reducedhabitatamountorreducedpatchsizes),
whereasothersincludetwoorthreeeffectsbutnotall four.
Does it matterwhichfragmentation measurea researcher uses?The answer
depends on whetherthe differenteffectsof the of
process fragmentation onhabitat
pattern have the same effects on If
biodiversity. they do, we can draw general
conclusionsaboutthe effectsof fragmentation on biodiversityeven thoughthe
differentstudiesmakingup the fragmentation literaturemeasurefragmentation
in differentways.As I showin Effectsof HabitatFragmentation on Biodiversity,
thedifferenteffectsof theprocessof fragmentation onhabitatpatterndonotaffect
biodiversityin thesameway.Thishasledto apparently contradictoryconclusions
abouttheeffectsof fragmentation on biodiversity.
Inthissection,I reviewquanti-
tativeconceptualizationsof habitatfragmentation. Thisis animportant steptoward
reconcilingtheseapparently contradictory results.

FRAGMENTATION LOSS The most obvious effect of the process of

AS HABITAT
fragmentationis theremovalof habitat(Figure1). Thishasled manyresearchers
to measurethe degreeof habitatfragmentation as simplythe amountof habitat
remainingon the landscape(e.g., Carlson
& Hartman 2001,Fuller2001, Golden
& Crist2000,Hargiset al. 1999,Robinsonet al. 1995,Summerville& Crist2001,
Virg6s 2001). If we can measurethe level of fragmentationas the amountof habi-
tat, why do we call it "fragmentation"?Why not simply call it habitatloss? The

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492 FAHRIG

reasonis thatwhenecologiststhinkof fragmentation, thewordinvokesmorethan

habitatremoval:"fragmentation ... not only causesloss of the amountof habi-
tat,butby creatingsmall,isolatedpatchesit also changesthe propertiesof the
remaining habitat"(vandenBerget al. 2001).
Habitatcanbe removedfroma landscapein manydifferentways,resultingin
manydifferentspatialpatterns(Figure2). Do somepatternsrepresenta higher
degreeof fragmentation thanothers,anddoes this haveimplicationsfor biodi-
versity?If the answer to eitherof thesequestionsis "no,"thenthe conceptof
fragmentation is redundant withhabitatloss. Theassertionthathabitatfragmen-
tationmeanssomethingmorethanhabitatloss dependson theexistenceof effects
of fragmentation on biodiversitythatcanbe attributedto changesin the pattern
of habitatthatareindependent manyresearchers
of habitatloss.Therefore, define
habitatfragmentation as anaspectof habitatconfiguration.

AS A CHANGE IN HABITATCONFIGURATION In addition to loss

FRAGMENTATION
of habitat,the processof habitatfragmentation resultsin threeothereffects:in-
creasein numberof patches,decreasein patchsizes, andincreasein isolation
of patches.Measuresof fragmentation thatgo beyondsimplyhabitatamountare
generally derived from these or other stronglyrelatedmeasures(e.g., amountof
edge). There are at least 40 such measures of fragmentation (McGarigalet al.
2002),manyof whichtypicallyhavestrongrelationships withtheamountof habi-
tat as well as witheachother(Belisleet al. 2001, Boulinieret al. 2001, Drolet
et al. 1999,Gustafson1998,Haines-Young & Chopping1996,Hargiset al. 1998,
Robinsonet al. 1995,Schumaker 1996, Trzcinskiet al. 1999,Wickham et al. 1999)
(Figure3).
Theinterrelationships amongmeasuresof fragmentation arenotwidelyrecog-
nizedin thecurrentfragmentation Mostresearchers
literature. do notseparatethe
effectsof habitatloss fromtheconfigurational effectsof fragmentation. Thisleads
to ambiguousconclusionsregardingthe effectsof habitatconfiguration on bio-
diversity(e.g., Summerville & Crist 2001, Swenson & Franklin2000). is also
It
commonfor fragmentation studiesto reportindividualeffectsof fragmentation
measureswithoutreporting therelationships amongthem,whichagainmakesthe
resultsdifficultto interpret.

PROBLEM Similarproblemsarise when fragmentationis mea-

THEPATCH-SCALE
suredat thepatchscaleratherthanthelandscapescale.Becausefragmentation is
a landscape-scale
process(Figure1), fragmentation are
measurements correctly
madeat the landscapescale (McGarigal& Cushman2002). As pointedout by
Delin & Andr6n(1999),whena studyis at the patchscale,the samplesize at
the landscapescale is only one, whichmeansthatlandscape-scaleinferenceis
not possible(Figure4; see Brennanet al. 2002, Tischendorf& Fahrig2000).
However,in approximately42% of recent fragmentationstudies, individualdata
points representmeasurementson individualpatches, not landscapes (Table 1).
Similarly,using a differentsampleof the literature,McGarigal& Cushman(2002)

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 EFFECTS
OFHABITAT
FRAGMENTATION 493

numberof patches
mean patchsize
mean isolation

A
numberof patches
mean patchsize
B mean isolation

numberof patches
C mean patchsize
mean isolation

D
numberof patches
mean patchsize
mean isolation
E

numberof patches
mean patchsize
mean isolation

Figure 2 Illustrationof habitatloss resultingin some,butnot all, of the otherthree

expectedeffects of habitatfragmentationon landscapepattern.Expectedeffects are
(a) an increasein the numberof patches,(b) a decreasein mean patch size, and
(c) anincreasein meanpatchisolation(nearestneighbordistance).Actualchangesare
indicatedby arrows.

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494 FAHRIG

A B

Size
Patches
of
Patch

Number Mean

0 HabitatAmount(%)
100 0 HabitatAmount(%) 100

C D

Distance

Edge

Neighbor Total

Nearest
0 100 0 HabitatAmount(%) 100
HabitatAmount(%)
Mean

Patch

Largest
of

Size
0 HabitatAmount(%) 100

Figure 3 Illustrationof the typicalrelationshipsbetweenhabitatamountandvariousmea-

suresof fragmentation. Individualdatapointscorrespondto individuallandscapes.Basedon
relationshipsin B61isleet al. (2001), Boulinieret al. (2001), Droletet al. (1999), Gustafson
(1998), Haines-Young& Chopping(1996), Hargiset al. (1998), Robinsonet al. (1995),
Schumaker(1996), Trzcinskiet al. (1999), andWickhamet al. (1999).

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 OFHABITAT
EFFECTS FRAGMENTATION 495

Patch-ScaleStudy Landscape-Scale
Study

Density

Patch
Population
in in

PatchSize
Density

Population
Landscape
AmountinLandscape
Habitat

Figure 4 (A)Patch-scalestudy.Eachobservationrepresentsthe informationfroma single

patch.Only one landscapeis studied,so samplesize for landscape-scaleinferencesis one.
(B) Landscape-scalestudy.Eachobservationrepresentsthe informationfroma singleland-
scape. Multiplelandscapes,with differentstructures,are studied.Here, sample size for
landscape-scaleinferencesis four.

estimatedthat more than 57% of all fragmentationstudies are at the patch scale.
Some researcherseven refer to patch-scalemeasuresas landscapefeatures(e.g.,
Fernandez-Juricic2000, Schweiger et al. 2000).

Patch size: an ambiguousmeasureof fragmentation The relationshipbetween

patch size and fragmentationis ambiguousbecause both habitatloss and habitat
fragmentationper se (i.e., the breakingapartof habitat,controlling for changes
in habitat amount) result in smaller patches (Figure 5). Using patch size as a
measureof habitatfragmentationper se implicitly assumesthatpatch size is inde-
pendentof habitatamountat the landscapescale (e.g., Niemelai2001). However,
regions where patches are large often correspondto regions where there is more
habitat(Fernandez-Juricic2000, McCoy & Mushinsky 1999) (Figure 6). Ignor-
ing potential relationshipsbetween a patch-scale measure (e.g., patch size) and
landscape-scalehabitatamountdoes not control for this relationship;it can lead
to misinterpretationof results.

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496 FAHRIG

Habitat Habitat
loss fragmentation
perse

Figure 5 Bothhabitatloss andhabitatfragmentation per se (independentof habitat

loss) resultin smallerpatches.Therefore,patch size itself is ambiguousas a mea-
sureof eitherhabitatamountor habitatfragmentation per se. Note also thathabitat
fragmentation per se leads to reducedpatchisolation.

Patch isolation: a measure of habitat amount In the fragmentationliterature,

patch isolation is almost universallyinterpretedas a measureof habitatconfigu-
ration. However, patch isolation is more accuratelyviewed as a measure of the
lack of habitatin the landscapesurroundingthe patch. The more isolated a patch
is, generally speaking, the less habitatthere is in the landscape that surrounds
it (Figure 7). Therefore,when translatedto the landscape scale, isolation of a
patch is a measure of habitatamountin the landscape,not configurationof the
landscape.
Bender et al. (2003) reviewed measures of patch isolation. All measures are
stronglynegatively related to habitatamountin the surroundinglandscape.The
most common measureof patchisolationis the distanceto the next-nearestpatch,

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 EFFECTS
OFHABITAT
FRAGMENTATION 497

1km
Figure 6 Landscapein southernOntario(fromTischendorf2001) showingthatregions
whereforestpatches(blackareas)are smalltypicallycorrespondto regionswherethereis
little forest.Compare(A) and (B), where(A) has smallpatchesandless than5%forestand
(B) has largerpatchesandapproximately 50%forest.

or "nearest-neighbor distance"(e.g., Delin & Andr6n1999, Haig et al. 2000, Hargis

et al. 1999). Patcheswith small nearest-neighbordistancesaretypicallysituatedin
landscapescontainingmore habitatthan are patches with large nearest-neighbor
distances (Figure 7), so in most situationsthis measureof isolation is relatedto
habitatamountin the landscape.Anothercommon measureof patch isolation is
the inverseof the amountof habitatwithin some distanceof the patchin question
(e.g., Kinnunenet al. 1996, Maguraet al. 2001, Miyashitaet al. 1998). In other
words, patch isolation is measuredas habitatamountat the landscapescale. All
othermeasuresof patch isolation are a combinationof distancesto otherpatches
and sizes of those patches (or the populationsthey contain) in the surrounding
landscape(reviewedin Bender et al. 2003). As such they are all measuresof the
amountof habitatin the surroundinglandscape.

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498 FAHRIG

I
jlN

I
I
I

1 km
Figure 7 Illustrationof the relationshipbetweenpatchisolationandamountof habitatin
the landscapeimmediatelysurroundingthe patch.Grayareasare forest.Isolatedpatches
(blackpatcheslabeled"I")aresituatedin landscapes(circles)containingless forestthanare
less isolatedpatches(blackpatcheslabeled "N").

MEASURING HABITAT FRAGMENTATION PERSE How can we measurehabitatfrag-

mentation independentof habitat amount? Some researchershave constructed
landscapes in which they experimentallycontrolledhabitatamountwhile vary-
ing habitatfragmentationper se (e.g., Caley et al. 2001, Collins & Barrett1997).
Researchersstudyingreal landscapeshave used statisticalmethodsto controlfor
habitatamount.For example,McGarigalandMcComb (1995) measured25 land-
scape indices for each of 30 landscapes.They statisticallycorrectedeach index
for its relationshipto habitat amount and then entered the corrected variables
into a PCA. Each axis of the resultingPCA representeda differentcomponentof

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 OFHABITAT
EFFECTS FRAGMENTATION 499

landscapeconfiguration.In a similarapproach,Villardet al. (1999) measuredthe

numberof forest patches, total length of edge, mean nearest-neighbordistance,
and percent of forest cover on each of 33 landscapes.They used the residualsof
the statistical models relating each of the first three variables to forest amount
as measuresof fragmentationthat have been controlledfor their relationshipsto
habitatamount.

EFFECTSOF HABITATFRAGMENTATION
ON BIODIVERSITY

InthissectionI reviewtheempiricalevidenceforeffectsof habitatfragmentation

on biodiversity.This review is not limited to the 100 paperssummarizedin Table
1. The fragmentationliteraturecan be distilled into two majoreffects: the gener-
ally strong negative effect of habitatloss on biodiversity,and the much weaker,
positive or negative effect of fragmentationper se on biodiversity.Because the
effect of fragmentationper se is weakerthanthe effect of habitatloss, to detectthe
effect of fragmentationper se, the effect of habitatloss must be experimentallyor
statisticallycontrolled.

Effectsof HabitatLoss on Biodiversity

Habitatloss haslarge,consistentlynegativeeffectson biodiversity,so researchers
who conceptualize andmeasurefragmentation as equivalentto habitatloss typ-
ically concludethatfragmentation has largenegativeeffects.The negativeef-
fects of habitatloss applynot only to directmeasuresof biodiversitysuch as
species richness(Findlay& Houlahan1997, Gurdet al. 2001, Schmiegelow
& Minkktinen2002, Steffan-Dewenter et al. 2002, Wettstein& Schmid1999),
population abundance and distribution(Best et al. 2001, Gibbs1998, Guthery
et al. 2001, Hanski et al. 1996, Hargis et al. 1999, Hinsley et al. 1995, Lande
& Calvo1999,Venier& Fahrig1996)andgeneticdiversity
1987,Sainchez-Zapata
(Gibbs2001),butalso to indirectmeasuresof biodiversityandfactorsaffecting
biodiversity.A model by Bascompte et al. (2002) predicts a negative effect of
habitatloss on populationgrowthrate. This is supportedby Donovan & Flather
(2002),whofoundthatspeciesshowingdecliningtrendsin globalabundance are
morelikelyto occurin areaswithhighhabitatloss thanarespecieswithincreas-
ing or stable trends.Habitatloss has been shown to reduce trophicchain length
(Komonen et al. 2000), to alter species interactions(Taylor & Merriam 1995),
and to reduce the numberof specialist, large-bodiedspecies (Gibbs & Stanton
2001). Habitatloss also negativelyaffectsbreedingsuccess(Kurkiet al. 2000),
dispersalsuccess(B61isleet al. 2001, Pither& Taylor1998,With& Crist1995,
With & King 1999), predationrate (Bergin et al. 2000, Hartley& Hunter 1998),
andaspectsof animalbehaviorthataffectforagingsuccessrate(Mahan& Yahner
1999).

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500 FAHRIG

INDIRECT EVIDENCEOF EFFECTSOF HABITATLOSS Negative effects of habitat

loss on biodiversity are also evident from studies that measure habitatamount
indirectly,using measuresthat are highly correlatedwith habitatamount.For ex-
ample, Robinson et al. (1995) found that reproductivesuccess of forest nesting
bird species was positively correlatedwith percentageof forest cover, percentage
of forest interior,and average patch size in a landscape.Because the latter two
variableswere highly correlatedwith percentageof forest cover, these all repre-
sent positive effects of habitatamounton reproductivesuccess. Boulinier et al.
(2001) found effects of mean patch size on species richness,local extinctionrate,
and turnoverrateof forestbirdsin 214 landscapes.Because meanpatchsize had a
0.94 correlationwith forestamountin theirstudy,this resultmost likely represents
an effect of habitatamount.

Patch isolation effects Patch isolation is a measureof the lack of habitatin the
landscapesurroundingthe patch(Figure7). Therefore,the many studiesthathave
shown negativeeffects of patchisolation on species richnessor presence/absence
representfurtherevidence for the strongnegativeeffect of landscape-scalehabitat
loss on biodiversity(e.g., McCoy & Mushinsky1999, Rukke2000, Virg6s 2001).
Benderet al. (2003) andTischendorfet al. (2003) conductedsimulationanalyses
to determinewhichpatchisolationmeasuresaremost stronglyrelatedto movement
of animalsbetween patches. They found that the "buffer"measures,i.e., amount
of habitatwithin a given bufferaroundthe patch,were best. This suggests a strong
effect of habitat amount on interpatchmovement. It also suggests, again, that
effects of patch isolation and landscape-scalehabitatamountare equivalent.

Patchsize effects Individualspecies have minimumpatchsize requirements(e.g.,

Diaz et al. 2000). Therefore,smallerpatchesgenerallycontainfewer species than
largerpatches (Debinski & Holt 2000), and the set of species on smallerpatches
is often a more-or-lesspredictablesubset of the species on largerpatches (e.g.,
Ganzhorn& EisenbeiB2001, Kolozsvary& Swihart1999, Vallan2000). Similarly,
the amountof habitaton a landscaperequiredfor species occurrencetherediffers
among species (Gibbs 1998, Vance et al. 2003), so landscapeswith less habitat
should containa subset of the species found in landscapeswith more habitat.
Despite this apparentcorrespondencebetween patch- and landscape-scaleef-
fects, the landscape-scaleinterpretationof patch size effects dependson the land-
scape context of the patch. For example, Donovan et al. (1995) found that forest
birds had lower reproductiverates in small patchesthanin large patches.If small
patches occur in areas with less forest, the reducedreproductiverate may not be
the result of patch size, but may result from largerpopulationsof nest predators
and brood parasitesthat occur in landscapeswith more open habitat(Hartley&
Hunter1998, Robinsonet al. 1995, Schmiegelow & Monkkinen 2002).

EXTINCTIONTHRESHOLD The numberof individualsof any species that a land-

scape can should
support be a positivefunctionof the amountof habitatavailableto

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 EFFECTS
OFHABITAT
FRAGMENTATION501

size area
Proportional
hypothesis

Extinction threshold
Population hypothesis

0
HabitatAmount
Figure8 Illustration
of the extinctionthreshold in comparison
hypothesis to the
area
proportional hypothesis.

thatspeciesin thelandscape.However,severaltheoretical studiessuggestthatthe

relationship is notproportional;theypredicta threshold habitatlevelbelowwhich
the populationcannotsustainitself,termedthe extinctionthreshold(Bascompte
& Sole 1996,Boswellet al. 1998,Fahrig2001, Flather& Bevers2002, Hill &
Caswell1999,Lande1987,With& King1999;Figure8). Therehavebeenvery
few directempiricaltestsof the extinctionthresholdhypothesis(butsee Jansson
& Angelstam1999).
Notethatthepredictedoccurrence of theextinctionthreshold resultsfromhabi-
tatloss, nothabitatfragmentation perse. Theoretical studiessuggestthathabitat
fragmentation perse canaffectwheretheextinctionthresholdoccurson thehabi-
tatamountaxis.Also, the effectsof habitatfragmentation perse arepredictedto
increasebelow some level of habitatloss (see The 20-30%Threshold,below).
However,the occurrenceof the extinctionthresholdis a responseto habitatloss,
notfragmentation perse. Thishasledto someambiguity in interpretation
of empir-
ical literature.Forexample,Virg6s(2001)foundthatpatchisolationaffectsbad-
ger densityonlyforpatchesin landscapeswith<20%forestcover.As explained
above,patchisolationis typicallyanindexof habitatamountatthelandscapescale.
Therefore, thisresultprobablysuggestsa thresholdeffectof forestloss on badger
density.Thisconclusionis differentfromthatof the author,who interpreted the
isolationeffectas aneffectof habitatconfiguration. Theinterpretation is ambigu-
ous becausetherelationship betweenhabitatamountandpatchisolationwasnot
statisticallycontrolledin thisstudy.Similarly,Andr6n(1994)reviewedpatchsize
andpatchisolationeffectson population densityandconcludedthattheseeffects

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502 FAHRIG

increasebelow a thresholdamountof habitatin the landscape.Because patchsize

and isolation can be indicatorsof habitatamountat a landscapescale (see Patch
size: An Ambiguous Measureof Fragmentationand Patch Isolation:A Measure
of HabitatAmount, above), this result could be interpretedas an intensification
of the effects of habitatloss at low habitatlevels, i.e., it supportsthe extinction
thresholdhypothesis (Figure8). This resulthas also been viewed as evidence for
configurationeffects below a thresholdhabitatlevel (e.g., Flather& Bevers 2002,
Villardet al. 1999). Again, the interpretationis ambiguousbecause the relation-
ships between patch size and isolation and amountof habitatsurroundingeach
patchwere not controlledfor.

Effectsof HabitatFragmentationper se on Biodiversity

In this section I reviewthe empiricalevidence for fragmentationeffects per se, i.e.,
for effects of "breakingapart"of habitaton biodiversity,thatareindependentof or
in additionto the effects of habitatloss. The 17 studiesin Table2 representall of the
empiricalstudies of fragmentationper se of which I am aware.Some theoretical
studiessuggest thatthe effect of habitatfragmentationper se is weak relativeto the
effect of habitatloss (Collingham& Huntley2000, Fahrig1997, Flather& Bevers
2002, Henein et al. 1998), althoughother modeling studies predict much larger
effects of fragmentationper se (Boswell et al. 1998, Burkey 1999, Hill & Caswell
1999, Urban& Keitt2001, With& King 1999;reviewedin Fahrig2002). All these
recent models predictnegativeeffects of habitatfragmentationper se, in contrast
with some earliertheoreticalwork (see Reasonsfor PositiveEffects of Fragmenta-
tion, below). The empiricalevidence to date suggests thatthe effects of fragmen-
tationper se are generallymuch weakerthanthe effects of habitatloss. Unlike the
effects of habitatloss, and in contrastto currenttheory,empiricalstudies suggest
thatthe effects of fragmentationperse areatleast as likely to be positiveas negative.
The 17 empiricalstudieson the effects of habitatfragmentationper se (Table2)
range from small-scale experimentalstudies to continental-scaleanalyses. They
cover a range of response variables, including abundance,density, distribution,
reproduction,movement, and species richness. About half of the studies are on
forest birds; other taxa include insects, small mammals,plants, aquaticinverte-
brates, and a virus, and other habitatsinclude grasslands,cropland,a coral reef,
and an estuary.
The 17 studies used a variety of approachesfor estimatingthe effect of frag-
mentationper se. In five of them, experimentallandscapeswere constructedto
independentlycontrolthe levels of habitatamountand fragmentationper se. Four
of these variedboth habitatamountand fragmentationper se, and one variedonly
fragmentation,holding the amountof habitatconstant.Threeof the 12 studies in
real landscapescomparedthe response variablein one large patch versus several
smallpatches(i.e., holdinghabitatamountconstant).In the remainingnine studies
in real landscapes,the effect of fragmentationper se was estimatedby statistically
controllingfor the effect of habitatamount.

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 EFFECTSOF HABITATFRAGMENTATION 503

of on
se
habitat per
of effect(s) negative negative negative (Continued)
of 1 4 2
effects
for effect effect effect
positive, negative positive, positive,
Direction
fragmentation
biodiversity
No 6 No 2 2 No 4 Positive
Positive

controlling
i.e., versus
se, lossse
per per
habitat
of

fragmentation fragmentation
fragmentation fragmentation
fragmentation fragmentation
fragmentation
fragmentation
fragmentation
effects
fragmentation
> > > > > - > >

habitat
of habitat n.a.a Amount
Relative Amount Amount Amount
Amount
Amount
Amount
Amount

effects
the

variable(s) homing
bird and success
examined owl:
(various):
forest presence/
reproductionspecies:
species:
species:
that incidence
species:
responsetaxa abundance
spotted species bird bird bird homing
bird
and andvirus:
studies forest
late-seral forest forest forest
Tanager forest
Taxa 15 species:
111distribution
Northern
absence
presence/absence,
persistence,
6 populations:
31 presence/absence
14 presence/absence
15 presence/absence
3 timeHanta

empirical
of
1999
1999 2001
McComb
Merriam
landscapes al.
&
1998 al. 1999 1999 2001 al.
Summary & et
biodiversity real al. et al. al. al. et
2 on in et et et et

1983 1995
Middleton
McGarigal
Meyer Rosenberg
Trzcinski
Drolet VillardBdlisleLanglois
TABLE
amount StudyStudies

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504 FAHRIG

of on
se
per
effect(s) (adult
of (juvenile (total
richness)
endangered
negative effect
positive positive
1 density)
andspecies positive
1 survival),
fragmentation
biodiversity
Direction 2 No Positive 1
Positive Positive n.a.b

versus
lossse
per
habitat
of amount
>
fragmentation fragmentation
fragmentation
effects
fragmentation
> > >>
stated
Relativen.a.a
habitat n.a.a Amountn.a.a
Fragmentation Amount
Not Amount

aggregation
rate,
variable(s) richness species
survival, richness, abundance,
density abundance,
density spatial
abundancevaried.
speciesabundance
species vole: was
juvenile
response insects: and
vole: reproductive
richness
(predator) birds: commensals:
insects: insects.
and crab:
the
adult endangered density, coral
species
recruitment richness of
Taxa Blue Butterflies: Gray-tailed
Forest Meadow 8
Grassland Clover
fragmentation

only

landscapes: distribution
1998
2001 2002 1997 constant;
spatial
al. 1999 held
was
et 1997 2001
Forman 2002 was
(Continued) al. Barrett &
Lipcius & al. al. al.
& et experimental et
2 in et et variable
amount

Hovel Flather Collins

Tscharntke Wolff Caley With
Collinge
TABLE Study Studies aHabitat
bResponse

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 EFFECTSOF HABITATFRAGMENTATION 505

The overall result from these studies is that habitat loss has a much larger
effect thanhabitatfragmentationper se on biodiversitymeasures(Table2). When
fragmentationper se did have an effect, it was at least as likely to be positive
as negative (Table2). Given the relatively small numberof studies and the large
variationin conditions among studies, it is not possible to tease apartthe factors
that lead to positive versus negative effects of fragmentationper se. However,
the positive effects of fragmentationcan not be explained as merely responses
by "weedy,"habitat generalist species. For example, the results reportedfrom
McGarigal& McComb (1995) are specificallylimited to late-seralforest species,
and Tscharntkeet al. (2002) found a positive effect of fragmentationper se on
butterfly species richness, even when they only included endangeredbutterfly
species.

THE 20-30% THRESHOLD Some theoretical studies suggest that the effects of frag-
mentationper se should become apparentonly at low levels of habitatamount,
below approximately20-30% habitaton the landscape (Fahrig 1998, Flather&
Bevers2002). Todate,thereis no convincingempiricalevidencefor thisprediction.
If the thresholddoes occur, it should result in a statisticalinteractioneffect be-
tween habitatamountand habitatfragmentationper se; such an interactionwould
indicate that the effect of fragmentationper se depends on the amountof habitat
in the landscape.Trzcinskiet al. (1999) tested for this interactioneffect but found
no evidence for it. The hypothesis that fragmentationeffects increase below a
thresholdof habitatamounthas not yet been adequatelytested.

REASONSFOR NEGATIVEEFFECTSOF FRAGMENTATION

PER SE Negative effects
of fragmentationare likely due to two main causes. First, fragmentationper se
implies a largernumberof smaller patches. At some point, each patch of habi-
tat will be too small to sustain a local populationor perhapseven an individual
territory.Species that are unable to cross the nonhabitatportionof the landscape
(the "matrix")will be confinedto a large numberof too-small patches,ultimately
reducingthe overallpopulationsize and probabilityof persistence.
The second main cause of negative effects of fragmentationper se is negative
edge effects; more fragmentedlandscapescontainmore edge for a given amount
of habitat.This can increasethe probabilityof individualsleaving the habitatand
enteringthe matrix.Overallthe amountof time spent in the matrixwill be larger
in a more fragmentedlandscape,which may increase overall mortalityrate and
reduceoverallreproductiverateof the population(Fahrig2002). In addition,there
arenegativeedge effects due to species interactions.Probablythe most extensively
studied of these is increasedpredationon forest birds at forest edges (Chalfoun
et al. 2002).

REASONSFOR POSITIVEEFFECTSOF FRAGMENTATION

PER SE More than half of
the effects of fragmentationperse thathavebeen documentedarepositive(Table2).
Some readerswill findthis surprising,probablybecausehabitatloss is inextricably

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506 FAHRIG

includedwithin theirconceptualizationof habitatfragmentation.In this case even

if fragmentationper se has a positive effect on biodiversity,this effect will be
maskedby the large negativeeffect of habitatloss.
Haila (2002) describeshow the currentconcept of habitatfragmentationemer-
ged fromthe theoryof islandbiogeography(MacArthur& Wilson 1967). The two
predictorvariablesin this theory are island size and island isolation, or distance
of the island from the mainland.When this theory was conceptually extended
from island archipelagosto terrestrialsystems of habitat patches, the concept
of isolation changed; isolation was now the result of habitatloss, and it repre-
sentedthe distancefrom a patchto its neighbor(s),not the distanceto a mainland.
Because of its roots in island biogeography,isolation was viewed as represent-
ing habitat subdivision even though it was inextricably linked to habitat
loss.
However,a parallelresearchstream,which arose independentlyof the theory
of island biogeography,suggested thathabitatfragmentationcould have positive
effects on biodiversity.Huffaker's(1958) experimentsuggestedthatsubdivisionof
the sameamountof habitatinto manysmallerpieces can enhancethepersistenceof
a predator-preysystem. He hypothesizedthathabitatsubdivisionprovidestempo-
raryrefugiafor the prey species, where they can increasein numbersand disperse
elsewherebeforethepredatororparasitefindsthem.Theplausibilityof thismecha-
nism was supportedby earlytheoreticalstudies(Hastings1977,Vandermeer1973).
Early theoreticalstudies also suggested that habitatfragmentationenhances the
stabilityof two-species competition(Levin 1974, Shmida& Ellner 1984, Slatkin
1974), and in an empiricalstudy,Atkinson& Shorrocks(1981) foundthatcoexis-
tenceof two competingspecies couldbe extendedby dividingthehabitatintomore,
smallerpatches.Enhancedcoexistenceresultedfroma trade-offbetweendispersal
rateand competitiveability.This trade-off,along with asynchronousdisturbances
thatlocally removedthe superiorcompetitor,allowed the inferiorcompetitor(but
superiordisperser)to colonize the emptypatchesfirst,beforebeing laterdisplaced
by the superiorcompetitor(Chesson 1985). Otherresearcherssuggestedthathabi-
tat subdivisioncould even stabilizesingle-speciespopulationdynamicswhen local
disturbancesareasynchronousby reducingthe probabilityof simultaneousextinc-
tion of the whole population(den Boer 1981; Reddingius& den Boer 1970; Roff
1974a,b).
Why has this earlywork,suggestingpositiveeffects of habitatfragmentationper
se, been largely ignored in the more recent habitatfragmentationliterature?One
reason is that later theoreticaland empirical studies (reviewedin Kareiva 1990)
demonstratedthat the predictedpositive effects of fragmentationper se depend
stronglyon particularassumptionsaboutthe relativemovementrates of predator
versusprey (or host versusparasite),the trade-offbetween competitiveabilityand
movementrate,andthe asynchronyof disturbances.It seems thatthe sensitivityto
these assumptions,alongwith the misrepresentationof patchisolationas a measure
of habitatsubdivision,led researchersto ignore the possibility thatfragmentation
per se could have a positive effect on biodiversity.

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 EFFECTS
OFHABITAT
FRAGMENTATION507

Thereareatleastfouradditional possiblereasonsforpositiveeffectsof habitat

fragmentation perse on biodiversity. First,Bowmanet al. (2002)arguedthat,for
manyspecies,immigration rateis a functionof the lineardimensionof a habitat
patchratherthantheareaof thepatch.Forthesespecies,overallimmigration rate
shouldbe higherwhenthe landscapeis comprisedof a largernumberof smaller
patches(higherfragmentation perse) thanwhenit is comprisedof a smallernum-
berof largerpatches.In situationswhereimmigration is animportantdeterminant
of population density, this could resultin a positive effectof fragmentation perse
on density.
Second,if habitatamountis held constant,increasingfragmentation per se
actuallyimplies smaller distances between a
patches(Figure5).Therefore,positive
effectof fragmentation perse couldbe dueto a reductionin patchisolation.
Third,manyspeciesrequiremorethanone kindof habitat(Law& Dickman
1998).Forexample,immature insectsandamphibians oftenusedifferenthabitats
thanthosetheyuse as adults.A successfullife cyclerequiresthatthe adultscan
moveawayfromthehabitatwheretheywererearedto theiradulthabitatsandthen
backto theimmature habitatto layeggs.Theproximityof differentrequired habi-
tattypeswill determinethe ease withwhichindividualscanmoveamongthem.
Forexample,Popeet al. (2000)showedthatthe proximityof feedinghabitatto
breedingpondsaffectedthe abundance of leopardfrogpopulations. Pedlaret al.
(1997)foundthatraccoonabundance washighestin landscapeswithintermediate
amountsof forest.Theysuggestedthatthislevel of forestmaximizedthe acces-
sibilityto the raccoonsof bothfeedingareas(grainfields)anddenningsites in
forest.
Thedegreeto whichlandscapestructure facilitatesmovementamongdifferent
required habitattypes was labeled "landscape complementation" byDunninget al.
(1992). For the same amount of habitat, a more fragmentedlandscape(more,
smallerpatches,and moreedge) will have a higherlevel of interdigitation of
differenthabitattypes.This shouldincreaselandscapecomplementation, which
hasapositiveeffectonbiodiversity (Law&Dickman1998,Tscharntke et al.2002).
Finally,it seemslikely thatpositiveedge effectsare a factor.Some species
do showpositiveedge effects(Carlson& Hartman2001, Kremsater & Bunnell
1999,Laurance et al. 2001).Fora givenamountof habitat,morefragmented land-
scapescontainmoreedge.Therefore,positiveedge effectscouldbe responsible
forpositiveeffectsof fragmentation perse on abundance or distributionof some
species.

CONCLUSIONSAND FUTUREDIRECTIONS

Habitat Loss Versus Fragmentation

Mostresearchersviewhabitatfragmentation asa processinvolvingboththelossof
habitatandthebreakingapartof habitat.Thefactthatmostfragmentationresearch
does not differentiatebetween these two effects has led to severalproblems.First,

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508 FAHRIG

the apparentinconsistencyin the effects of a single process (fragmentation)gives

the impressionthat fragmentationeffects are difficultto generalize.In fact, gen-
eralizationis possible, but only for the separatecomponentsof fragmentation,not
for the combinedconceptof loss andbreakingapartof habitat.Empiricalevidence
to date suggests thatthe loss of habitathas large negative effects on biodiversity.
On the otherhand,the breakingapartof habitat,independentof habitatloss, has
ratherweak effects on biodiversity,which are as likely to be positive as negative.
Second, the merging of these two aspects of fragmentationhas obscuredthe
fact thatthe effects of habitatloss outweighthe effects of habitatfragmentationper
se. In fact, the effects of fragmentationper se areabsentor too small to be detected
in most empiricaltests to date.This is in contrastto severaltheoreticalpredictions
(Burkey1999, Hill & Caswell 1999, Urban& Keitt2001, With & King 1999) and
has importantimplicationsfor conservation.It suggests that conservationefforts
shouldfocus on habitatpreservationandrestoration.It also suggests thatresearch
in supportof particularconservationproblems should focus on determiningthe
amount of habitatrequiredfor conservationof the species of concern. The fact
that effects of fragmentationper se are usually small and at least as likely to be
positive as negative suggests that conservationactions that attemptto minimize
fragmentation(for a given habitatamount)may often be ineffectual.
Note, however,thatthis conclusion is preliminarybecause thereare still only a
small numberof relevantempiricalstudies.To my knowledgethereare,to date,no
studiesin tropicalregions of the effects of forestfragmentationper se (controlling
for habitatloss). Lauranceet al. (2002) concludedthatin Braziliantropicalforest
there are strong negative effects of forest edge on several taxa. These effects
are apparentlymuch strongerthan negative edge effects in temperatesystems
(Kremsater& Bunnell 1999). Negative edge effects could translateinto a negative
effect of fragmentationper se at the landscape scale because fragmentationper
se increases the amountof edge on the landscape.This suggests that effects of
fragmentationper se may be greaterin tropicalsystemsthanin temperatesystems.
This predictionremainsto be tested.
Third,ambiguousempiricalresultscould lead to errorsin modelingstudies.For
example,Donovan& Lamberson(2001) constructeda model to look at the effects
of habitatfragmentationon populationgrowthrate. They held amountof habitat
constantandvariedmeanpatchsize. Forinputparameterstheyused empiricalwork
suggestingthatreproductivesuccess increaseswithincreasingpatchsize. However,
as they point out, in these empiricalstudiespatch size was highly correlatedwith
habitatamountin the surroundinglandscape.It is not knownwhetherreproductive
success increaseswith increasingpatchsize when habitatamountin the landscape
is held constant.It could be that reproductivesuccess increases with amountof
habitaton the landscape,independentof habitatfragmentationper se. If this is
true,the resultsof the simulationmay be misleading.
These conclusions are based on the relatively small, but growing, numberof
empiricalstudiesthatseparatethe effects of habitatloss andfragmentationper se.
So far these studieshave been conductedon a limited set of taxa primarilywithin

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 EFFECTS
OFHABITAT
FRAGMENTATION509

NorthAmerica.Moreresearchis neededto determine

howgeneraltheconclusions
are(Harrison
& Bruna1999).

A USEFUL
IS "FRAGMENTATION" TERM?The term"fragmentation"
is quickly los-
ing its usefulnessas moreandmoreeffectsof humanactivitiesareincorporated
intothissingleterm.Someauthorshaveevensuggestedthatsomespeciesare"in-
dicatorsof fragmentation"(e.g.,Hager1998,Niemeli2001).Theimplication that
fragmentation canbe indicatedby the declineof somespeciesor speciesgroup
suggeststhatthe termis becominga catchallfor human-caused habitatchanges
thathavenegativeeffectson biodiversity. As questionedby Haila(2002),"Isa
conceptuallyambiguousandempiricallymultifaceted termfruitfulas a generic
description of humaneffectson landscapes?"
I suggestthattheterm"fragmentation" shouldbe limitedto thebreakingapart
of habitat.Habitatloss shouldbe calledhabitatloss; it has importanteffectson
biodiversity thatare of
independent any effectsof habitatfragmentation per se.
Habitatfragmentation shouldbe reservedforchangesin habitatconfiguration that
resultfromthebreakingapartof habitat,independent of habitatloss.

Implications for Biodiversity Conservation

Does ourknowledgeaboutfragmentation effectshavegeneralimplicationsfor
conservation of biodiversity,
particularlysimultaneousconservation
of multiple
species?The fragmentation literatureprovidesstrongevidencethathabitatloss
haslarge,consistentlynegativeeffectson biodiversity.Thisimpliesthatthemost
important questionforbiodiversityconservation is probably"Howmuchhabitat
is enough?"Differentspeciesuse differentkindsof habitat,anddifferentspecies
requiredifferentamountsof habitatfor persistence.Therefore,conservation of
all speciesin a givenregionrequiresidentifyingwhichspeciesin thatregionare
mostvulnerableto habitatloss (Fahrig2001, With& King1999)andestimating
the minimumhabitatrequiredfor persistenceof each of thesemostvulnerable
species.Thisdeterminesthe minimumhabitatamountsfor eachkindof habitat
in the region.In addition,manyspeciesrequiremorethanone kindof habitat
withina life cycle.Therefore,landscapepatternsthatmaintaintherequiredhabitat
amounts,butintersperse the differenthabitattypesas muchas possible,should
producethelargestpositivebiodiversity response(Law& Dickman1998).

ACKNOWLEDGMENTS
I thankJeffBowman,JulieBrennan, DanBert,Tormod Burkey,NeilCharbonneau,
KathrynFreemark, Jeff
AudreyGrez, Houlahan, Jochen Jaeger,MaximLarriv6e,
LutzTischendorf,RebeccaTittler,PaulZorn,andmembersof theLandscape Ecol-
ogy Laboratoryat Carleton
forcomments on anearlier
versionof thismanuscript.
Thisworkwas supported by a grantfromthe NaturalSciencesandEngineering
ResearchCouncil of Canada.

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510 FAHRIG

The AnnualReviewof Ecology,Evolution,and Systematicsis online at

http://ecolsys.annualreviews.org

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