Sei sulla pagina 1di 15
Society for Conservation Biology Disturbance, Diversity, and Invasion: Implications for Conservation Author(s): Richard J.

Society for Conservation Biology

Disturbance, Diversity, and Invasion: Implications for Conservation Author(s): Richard J. Hobbs and Laura F. Huenneke Source: Conservation Biology, Vol. 6, No. 3 (Sep., 1992), pp. 324-337 Published by: Blackwell Publishing for Society for Conservation Biology

Accessed: 26/01/2011 13:02

Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless

you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use.

Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at .

Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission.

JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org.

information about JSTOR, please contact support@jstor.org. Blackwell Publishing and Society for Conservation Biology

Blackwell Publishing and Society for Conservation Biology are collaborating with JSTOR to digitize, preserve and extend access to Conservation Biology.

http://www.jstor.org

Review

Disturbance,Diversity,and Invasion:

ImplicationsforConservation

RICHARDJ. HOBBS

CSIRO DivisionofWildlife& Ecology LMB 4, PO Midland WesternAustralia6056, Australia

LAURAF. HUENNEKE

DepartmentofBiology New Mexico StateUniversity Box 30001/Dept3AF Las Cruces,New Mexico 88003, U.S.A.

Abstract:Disturbance is an importantcomponentof many ecosystems,and variationsin disturbanceregimecan affect ecosystemand communitystructureand functioning.The "intermediatedisturbancehypothesis"suggeststhatspecies diversityshould be highestat moderatelevelsofdisturbance. However,disturbanceis also known to increase theinvasi- bilityof communities.Disturbance thereforeposes an im- portantproblemfor conservationmanagementHere,we re- view the effectsof disturbancessuch as fire,grazing soil disturbance,and nutrientaddition onplant speciesdiversity and invasion, withparticular emphasis on grassland vege- tation.Individual componentsofthedisturbanceregimecan

Introduction

Preservationof naturalcommunitieshas historically consistedof measuresprotectingthemfromphysical disturbance.Timberharvestsand livestockgrazingare usuallyexcluded frompreserves,and firesuppression has been practiced-withinthe U.S. systemofnational parks,forexample.Ecologistsand conservationistshave come to recognize,however,thatmanyformsofdistur- bance are importantcomponentsof naturalsystems. Manyplantcommunitiesand species are dependenton

Addressall correspondencetoRichardJ Hobbs. Paper submittedJune 19, 1991; revisedmanuscriptacceptedFebru- ary 14, 1992.

324

ConservationBiology Volume6, No. 3, September1992

have markedeffectson speciesdiversity,but it is oftenmod- ificationsof theexistingregimethathave thelargestinflu- ence.Similarly,disturbancecan enhance invasion of natu- ral communities,butfrequentlyit is theinteractionbetween different disturbancesthathas thelargesteffectThenatural disturbanceregimeis now unlikelytopersistwithinconser- vation areas, sincefragmentationand human intervention have usually modifiedphysicaland bioticconditions.Active managementdecisions must now be made on what distur- bance regimeis required,and thisrequiresdecisionson what speciesare to be encouragedor discouraged.

disturbance,especially for regeneration(Pickett & White1985). Preservesshouldbe largeenoughto allow the naturaldisturbanceregimeto operate and to sup-

port a mosaic of patches in differentstages of distur- bance, successionalrecovery,and communitymatura- tion (Pickett & Thompson 1978). In addition,both theory(the intermediatedisturbancehypothesis,Con- nell 1978) and growingempiricalevidencesuggestthat moderatefrequenciesor intensitiesof disturbancefos- termaximumspeciesrichness.To preservebioticdiver-

sityandfunctioningnaturalecosystems,then,conserva-

tion effortsmust include explicit consideration of disturbanceprocesses. Disturbanceacts in plant communitiesin another way, however,by promotinginvasionsby non-native

Hobbs& Huenneke

and weedy plant species (Ewel 1986; Hobbs 1989, 1991; Rejmanek1989). Invasivespecies have recently gained notorietyas major conservationand manage- mentconcernsinnaturalecosystems(see MacDonaldet al. 1989; Soule 1990; Westman1990). The controlof non-nativeplantshasbecome one ofthemostexpensive and urgenttasks of managersin several U.S. national parks,in island preservessuch as the Galapagos,and elsewhere.Invasiveplantscan reduce or displacenative species,bothplantand animal,and mayeven altereco- systemfunction(Vitousek1986; Schofield1989). They have become recognized,therefore,as significantcon- servationconcerns. Disturbancethuspresentsa conundrumto conserva- tionmanagement:thecontinuedexistenceofparticular species or communitiesoftenrequiresdisturbanceof some type-and hence disturbanceregimesmustbe in- tegratedwithmanagementplans-but disturbancemay simultaneouslylead to the degradationofnaturalcom- munitiesbypromotinginvasions.Herewe examinethis problemby discussingthe typesofdisturbanceimpor- tantinmaintainingplantspecies diversityandthosethat encourageinvasions.We identifyparticularcases where conflictsare most likelyto arise. Our examples are drawnprimarilyfromgrasslandvegetation,althoughwe discuss otherecosystemtypessuch as shrublandsand woodlands.We close by suggestingguidelinesforeval- uatingthe proper role of disturbancein the manage- mentofa naturalarea or preserve.

TheoreticalBackground

Therehas been considerabledebateon thedefinitionof disturbance,and on what does and does not constitute

a disturbanceto any given communityor ecosystem (see Rykiel1985; van Andel & van den Berg 1987). Definitionsof disturbancevary,fromGrime's(1979) view ofdisturbanceas a processremovingor damaging biomass,to White and Pickett's(1985) definitionof "anyrelativelydiscreteeventin timethatdisruptseco- system,communityorpopulationstructureandchanges resources,substrateavailability,or thephysicalenviron- ment."Petraitiset al. (1989) expand thedefinitionfur- therto include any "process thataltersthe birthand death ratesof individualspresentin the patch" by di- rectlykillingindividualsor by affectingresourcelevels, naturalenemies,or competitorsin ways thataltersur- vival and fecundity.Temporal and spatial scale are clearlyimportantin our recognitionof the "discrete- ness"ofa disturbanceevent,as nearlyanyecologicalor biogeochemicalprocess mightfallunderthe last,most inclusivedefinition.Pickettet al. (1989) definea dis- turbanceas a changein structurecaused byfactorsex- ternalto thehierarchicallevel ofthesystemofinterest; thisis necessaryto distinguishdisturbancefromother

Disturbance,Diversity,andInvasion 325

changesin thesystem.In our discussionbelow,we will include both direct disturbances(those affectingthe survivorshipofindividualsdirectly)and indirectdistur- bance (those affectingresourcelevels or othercondi- tionsthattheninfluenceindividualsin thepatch). Dis- turbances to plant communities thus include such events as fires,storms,and floods;but other changes suchas alteredgrazingregimesor nutrientinputswould also be classed as disturbanceiftheyaffectedresource levels and demographicprocesses. Withina givenpatch,theresponseofanycommunity to a disturbance(or to the disturbanceregime,charac- terisedby the naturaldistributionofdisturbancesizes, frequencies,intensities,and timing)is determinedby the attributesof componentspecies. Disturbancefre- quencyis also important;thetimeintervalbetweensuc- cessivedisturbancescan havesignificanteffectson com- munityresponse.This is because species composition changeswithtimesince disturbance,and manyspecies requiresome timeafterdisturbanceto reachreproduc- tivematurity.Ifa second disturbanceoccursbeforethey reachthatstage,therewill notbe anypropagulesavail- able to recolonizethepatch.The responseofa commu- nityto disturbanceis thenpredictedon thebasisofthe lifehistoryresponsesof those species availableforre- cruitmentor invasion(Noble & Slatyer1980; Moore & Noble 1990). Therehasbeen increasingrecognitionamongecolog- ical researchersof the importanceof naturaldistur- bance in the functionof terrestrialecosystems.Pickett andWhite(1985) provideda comprehensivereviewof therole ofdisturbancein thedynamicsofmanyecosys- tem types.They distinguishedseveral componentsof

naturaldisturbanceregimes,includingfrequency,inten-

sity,and size of disturbance(White & Pickett1985), each actingin a distinctiveway on communitiesand populations.Petraitiset al. (1989) presenteda more detailedanalysisof these components,and recognized thathypothesesabout the relationshipof disturbance andcommunityresponsecan be sortedintotwogroups:

thosepostulatingselectivemortalityor actionfora spe- cifictargetgroup,and those dealingwith randomor catastrophicmortality.Petraitiset al. (1989) suggested thatselectivemortalitycould maintainspecies diversity

or richnessat some equilibriumlevel, while random mortalitywould preventthe establishmentof commu- nityequilibrium(for example, preventingthe domi- nance ofone superiorcompetitorand the exclusionof otherspecies). Theypointedout thatbothequilibrium

and nonequilibriummodels of communitiespredict greatestspecies richnessat intermediatelevels of dis-

turbance.Variousversionsofthis"intermediatedistur-

bance hypothesis"thuspredicta similarresult-highest

species numberswhen disturbancesoccur at interme- diatefrequenciesor withintermediateintensities(Fig. 1)-despite differentunderlyingtheoriesofcommunity

ConservationBiology Volume6, No. 3, September1992

326 Disturbance,Diversity,and Invasion

C,)

a2)

a1)

U)

Low

Disturbancefrequency/intensity High

Figure 1. The intermediatedisturbancehypothesis, whichindicatesthatspeciesdiversitywithina given patch should be highestat intermediatefrequencies or intensitiesof disturbance(afterConnell 1978).

function(see Fox 1979; Huston 1979; Sousa 1984). These argumentsare oftenbased on thefactthatonlya few species (ruderals) can persistin the face of fre- quent,severedisturbance;onlya fewspecies (the long- est-lived,bestcompetitors,and thoseable to regenerate withoutdisturbance)can persistover the long termin theabsence ofdisturbance;butmanyspecies(including some representativesof each of these,plus intermedi- ates) can findsome place to survivein a regioncom- prisingpatches in variousstagesofrecovery,arisingat some intermediatefrequency. How is "intermediate"defined?Itisperhapseasiestto relatethefrequencyofdiscreteeventsto thelongevity ofmajorspecies in the system.Approximatelyhalfthe lifespanof the dominantspecies has been used as one estimation of intermediate disturbance frequency (Hobbs et al. 1984). Definitionsof intermediateinten- sitymayhave fewerexternalreferents,however;inten- sitycan be evaluatedin termsofpercentageofindivid- uals killed, or the degree of structuralor resource alterationcaused. The above discussionconcentrateson within-patch diversity(alpha diversity),but disturbanceis also im- portantforcreatingor maintainingdiversitybetween patches or at the landscape level (beta diversity).By creating patches of differentages and successional stages,disturbanceaffectsstructuraland habitatdiver- sityas well as overallspecies diversity.Whilewe con- centrateon within-patchdiversityin this review,the roleofdisturbanceincreatinglandscapemosaicsshould also be noted (see Turner1987). Whiledisturbanceis importantformaintainingdiver- sitybothwithincommunitiesandat a landscapelevel,it has become increasinglyrecognized that disturbance mayalso haveundesirableeffects.Particularlyimportant is the recognitionthatdisturbancemayact to increase

ConservationBiology Volume6, No. 3, September1992

Hobbs& Huenneke

thelikelihoodofinvasionofa community.For invasion to occur theremustbe availablepropagulesofan inva- sive species capable of dispersinginto a given plant community,and therethenhas to be a suitablemicro- siteforgerminationand establishmentto occur.Thatis, therehasto be a suitableinvasion"window"(Johnstone 1986). Disturbanceusuallyacts primarilyby affecting the availabilityof suitable microsites,althoughsome formsofdisturbancemayaffectthe availabilityofinva- sive propagules. For instance,non-nativeherbivores maybringseed intoan area eitheron theircoats or in feces.Here we will discussprimarilythe effectof dis- turbanceon micrositeavailability. The spatialand temporaldistributionofdisturbance ina regionor an ecosystemgivesriseto thedisturbance mosaic of an area. Pickett and Thompson (1978) pointedout thatthe recurrenceofdisturbancesneces-

sitatesthe preservationof a or an area large enough to

patches in various stages of disturbanceor recovery such thatinternalrecolonizationcan contributeto the maintenanceoftheoverallecosystem.The dynamicsof

patch disturbance and of biotic exchanges among patches,whichdeterminethepatternofrecovery,areof majorconcernin definingthe minimumcriticalsize of

ecosystems(see Lewin1984), thesize requiredtomain-

taincharacteristicspeciescompositionandsystemfunc-

tion.Withincreasingfragmentationofnaturalareas,itis likelythatthese minimumareas are now to be found onlywithinthe largestconservationunits,and distur- bance regimesand biotic exchangesbetween patches are liable to be significantlyalteredin smallerremnant areas(Hobbs 1987; Saunderset al. 1991). In particular, invasionsare likelyto become more important.How shall managers respond to or compensate for the changednatureofdisturbance?We approachthisques- tion by surveyingthe major typesof disturbanceand reviewingtheireffectson plant species diversityand invasions.We mostlyconsidergrasslands,but we also include illustrativeexamples fromother vegetation types.

"minimumdynamicarea," containwithinit multiple

EmpiricalEvidence

1. Fire

The centralroleoffireinmaintainingtheopen natureof thevegetationhas been acknowledgedformanygrass- lands,particularlyin mesic regions.Further,research has documentedthatfirecan stimulateor maintainhigh primaryproductivity.In tall-grassprairies of North America,fireenhances productivityby removingthe thicklitterlayerand alteringthe microclimateand nu- trientcontentof surfacesoil (see Knapp & Seastedt 1986). Fire also influencesspecies diversityand the characteristicstructureof these prairiecommunities. Classical work on fireecology of prairies(Kucera &

Hobbs& Huenneke

Koelling 1964; Abramset al. 1986) foundthatannual burningfavoredtallwarm-seasongrassesandresultedin low abundanceoftypicalprairieforbsafter5-10 years. Biennial burning resulted in the highest commu- nitydiversitywithmixed grassesand forbs.Areaswith longfire-freeperiodsresembledunburnedareasintheir heavylitteraccumulationand decline in grasses. Firesmayfavorthedominant"matrix"prairiegrasses and thuscan actuallydecrease diversity(Collins 1987). Apparentlymost prairie fires stimulate individual grassesand do not killthem;fewopeningsare created forthe establishmentofnew individualsor species.As

we have noted previously,however,species diversity comprisestwo maincomponents:species densityor al- pha diversitywithina patch,and patchdiversityor the numberof typesof differentpatches or microhabitats. Glenn-Lewinand ver Hoef (1988) reportedthatgrass- landsvaryin thedegreeto whichthesetwo contribute to overalldiversity.In threegrasslands,patch diversity ratherthanspecies densitywas themajorcontributorto overall communitydiversity.Fire (and other distur- bances) may create a heterogeneouspatch structure, evenifwithinpatchesitservesto decreasespecies den- sity. Lifehistory,of course, determinesthe vulnerability and response of plantsto fire.In annual grasslandsin California,firehad onlytemporaryeffectson botanical composition(forbsincreasedand grassdominancede- creasedfora brieftime). Here the restructuringofthe communityeach autumn with germinationquickly swampsany temporaryeffecton the seed bank or on germinationconditions(Heady 1972). Suppressionoffiresin ecosystemsdominatedbyfire- adaptedspecies can cause severedisruptionofcommu- nityand ecosystemprocesses,whichmayhave implica- tionsfortheconservationofnative,fire-tolerantspecies. Forexample,Cowlinget al. (1986) foundthatfiresup- pressionhas been responsibleforthe conversionof a South Africanopen, grassyveld to a vegetationnow

dominatedbyundesirablenon-nativeshrubs.Theysug-

gestedfrequentprescribedfiresas thebest mechanism forrestoringthe originalopen natureofthevegetation andformaintainingpopulationsoftheregion'sendemic geophytes.Strang(1973) similarlysuggestedthatfire was an expensive but necessarypart of reversingthe conversionofmoistgrasslandin south-centralAfricato brush.Firecan also be used morepreciselyto favorthe performanceofone species over another.For example, in an attemptto restoreprairieon the site of an aban- doned agriculturalsite,firewas used successfullyto cre- ate openingsin a turfofnon-nativePoa species and to enhancethe colonizationand expansionoftrueprairie species (Curtis& Partch1948). As earlyworkin tall-grassprairieconfirmed,theover- all fireregimeratherthananysinglefireis the critical

Disturbance,Diversity, andInvasion 327

factorin determiningcommunityresponse.Firesofdif- feringintensityor occurringin differentseasons are

likelyto affectspecies diversityin a varietyofways by alteringthepotentialofindividualspeciesto regenerate. Hobbs et al. (1984) provide an example of how fire

intervalscan

to regeneratein heathland,and hence affectoverall

communitydiversity.Anintermediatefirefrequencyre-

sultedin thehighestspecies diversity. Firehas been discussedas a factorthatcan increase the likelihood of invasions (Christensen & Burrows 1986). Fire acts to removemuch of the plant canopy andusuallyhas a short-termfertilizingeffecton thesoil; hence both lightand nutrientavailabilitycan be in- creased temporarily.Zedler and Scheid (1988) discuss theinvasionofcoastalchaparralby Carpobrotusedulis

followingfire.There is clear evidence,however,that not all firesresultin increasedinvasionand thatvaria- tions in fireregimecan affectthe extentof invasion. Hobbs and Atkins(1990) have illustratedhow invasion ofBanksia woodlands differsbetween firesburnedin springversusautumn.In somecases,fireper se does not affectthe degree of invasion,or will do so onlywhen combinedwithsome othertypeofdisturbance,such as mechanicaldisturbanceofthesoil or nutrientinput.For instance,Hesterand Hobbs (1992) studiedburnedand unburnedshrublandpatcheswithinan area of remant vegetationin the WesternAustralianwheatbelt and foundthatinvasionbynon-nativeannualswas restricted to theremantedges,evenfollowingburning.In adjacent woodland,theabundanceofnon-nativespecies actually declinedfollowingthefire.Followinganotherfireinthe same area, thistime in heathiandvegetation,invasion increasedonlywhere thefireimpingedon a roadverge thathad been subjectto priordisturbanceduringroad grading.This interactionis importantwhen manage- ment of roadside vegetationcorridorsis considered (see Loney& Hobbs, 1991; Panetta& Hopkins1991). Because species varyin theirresponse to fires,fire mayfavorone set of species over another;these rela- tionshipscan explain the balance between nativeand non-nativespecies in some fire-impactedsystems. Wherenativespecies are sensitiveto fire(because fuel loads were such thatfiresin thenativeecosystemwere of low frequencyand intensity),firecan enhance the invasionofnon-nativefire-tolerantspecies. Whenthese fire-tolerantspecies contributeto increasedfuelloads and inflammability,the disturbance regime can be shiftedtowardmore frequentand intensefires;these firesfurtherenhancethedominanceofnon-nativeover nativespecies.Justsuch a cycle has enhancedinvasion ofwoody species in SouthAfricanMediterraneansys- tems,and of annualgrassesinto otherMediterranean- climateregions(MacDonald et al. 1989). Similarly,in- vasionoffire-tolerantgrassesin dryHawaiianlowlands

alterthe diversityofspecies thatare able

ConservationBiology Volume6, No. 3, September1992

328 Disturbance,Diversity,andInvasion

has had severe effectson nativespecies (Hughes et al.

1991).

2. Grazing

Grazinganimalsare conspicuousand importantfeatures ofmanygrasslands;it has long been knownthatsome plantsare tolerantofgrazingwhile othersare not,and that grazingalters the appearance,productivity,and compositionofgrasslands.Milchunaset al. (1988) have reviewedthe effectsof grazingby largeherbivoreson

differingtypesofgrasslandand relatetheseto theinter-

mediatedisturbancehypothesis.Theysuggestthatgraz-

ingconstitutesa disturbanceonlywheretheevolution- ary historyof grazing is short. This has also been discussedby Naveh and Whittaker(1980) and Peet et al. (1983). We suggest,however,thatin anysituationa significantchange in grazingregimewill constitutea disturbance.Thus,impositionofgrazinganimals(or dif- ferentherbivores)on a systemnotpreviouslysubjectto thattype or level of grazingwill constitutea distur- bance.So,too,will theremovalofgrazingfroma system with a long grazinghistory.Species diversitywill be affectedby the directionof change in grazingregime relativeto the historicalregime(Ranwell 1960; White 1961; van der Maarel 1971; Milchunaset al. 1990; Dol- man & Sutherland1991). Numerousauthorshave re- portedmaximumspecies diversityunderintermediate levelsofgrazing(Zeevalking& Fresco 1977; Milchunas et al. 1988; Puertoet al. 1990). The most detailedunderstandingof how grazingaf- fectscommunitystructurecomes fromthe chalkgrass- landsofBritainandnorthernEurope;theseinfertilesites supporta diverse mixtureof grasses and forbs,with species adaptedto openingsofdifferentkindsandscales (Grubb 1976). These communities,althoughadmittedly artifactsof humanactivity(clearing,fires,or grazing), have long been prominentfeaturesof the landscape; today they are of major conservationvalue both for theirdiversityand forparticularrare species. Repeat- edlyit has been demonstratedthatgrazingis an impor- tantfactorin themaintenanceofchalkgrasslanddiver- sity;thecessationofgrazingleads to dominanceofa few grasses,and even to incursionsby shrubs or other woody species (Wells 1969). Entirecomponentsofthe floramay be lost; for example, During and Willems (1986) blamedthe loss ofmostlichensand theimpov- erishmentof the bryophteflorain Dutch chalk grass- landson the absence ofgrazing. Grazing maintainshigh species diversityin other grasslands,as well. Grazingmanagementis an important and successfultechnique forpreservingdiversityand conservationvalue ofold grasslandsand pasturesin En- gland (Hopkins & Wainwright1989). Sykora et al. (1990) found that grasslandon embankmentsin the

ConservationBiology Volume6, No. 3, September1992

Hobbs& Huenneke

Netherlandswas convertedto woody scrub in the ab- sence of grazing;under lightgrazing,a species-poor grasslandresultedfromcompetitionfroma few com-

petitivegrasses.Under more intensivegrazing,those grassesdid notdominate,and a more diversegrassland was maintained.Mediterranean-climategrasslandsmay respondsimilarlyto grazingmanagement;ina California grasslandon serpentinesubstrate,cessationoflivestock grazingenhancedthe dominanceof non-nativeannual grassesand led to a rapiddecline in abundance of the diversenativeannualforbflora(Huennekeet al.,unpub- lisheddata). One straightforwardeffectof grazingis the elimina-

tionoftreesandshrubsinvadingmesicgrasslands.With-

out grazing,manyNorthAmericanprairiesites have been convertedto woodland. Similarly,thereare also documentedcases of grazingpreventingor reversing the successionofAfricansavannato woodland. For ex- ample,Smartet al. (1985) foundthatin Uganda the exclusion of elephantswas even more importantthan firesuppressionin encouragingacacia invasion,leading to theloss ofmanyspecies includingtheoriginalgrass- land dominants.In these regions,a long evolutionary historyofgrazinghas led to thedominanceofgrassland plantsadaptedto and tolerantofgrazingpressure.Cald- well'swork(forexample,Caldwellet al. 1981) has doc- umented the many physiological traitsthat affecta plant'stoleranceofgrazinglosses. In contrast,regionswithno recenthistoryofgrazing are oftendominatedbyplantsthatlack thesetolerance mechanisms.Extremeexamples are presentedby oce- anic islands with no native mammalianherbivores, wheretheintroductionoflivestockor othergrazershas usuallybeen catastrophicin itseffecton nativevegeta- tion-for example,the effectof feralgoats on island floras(Coblentz 1978) and ofintroducedherbivoresin the Galapagos on native vegetation(Hamann 1975, 1979). A less obvious but stillmajor impacthas been made on regionswithfewnativegrazers(at leastsince post-Pleistocenetime),such as theintermountainWest (Mack & Thompson1982). In semi-aridgrasslandsin the AmericanSouthwest, species diversityhas declined and, in manycases, the physiognomyof the vegetationhas been alteredfrom perennialgrasslandto shrub-dominateddesert scrub. The chiefquestionofrangemanagementand ecologyis the determinationof the proper utilizationrate:what levelofgrazingwillmaximizeproductivityandmaintain thegrassland'sgeneralcharacter?Unfortunately,itis not knownwhat utilizationlevel maximizesplant species diversityor productivity,or whetherthe same level maximizesboth.Westobyet al. (1989) outlinedthedif- ferencesin grazingmanagementthatwould resultfrom consideringgrazingin an equilibrial,successionalcon- text versus a nonequilibrialseries of alternatestates;

Hobbs& Huenneke

workingwiththe second mentalmodel requiresmuch moreactivemanagementto "seize opportunitiesand to evade hazards." Givengrazing'simpacton communitystructure,ithas been used as a managementtool in conservationappli- cations.One exampleis a grasslandrestorationproject, where an abandoned,species-poorpasturein Holland was beinggrazedbycattle;seed inputsfromcattlefeces (togetherwith openingscreated by grazing)contrib- uted significantlyto increasingspecies diversity(Bu- low-Olsen 1980). In anothercase, sheep were used to restoreabandonedfields(Gibson et al. 1987), againby importingseeds and creatingopeningsforrecruitment. Severalgrassesin theMiddleEast,wild cereal ancestors of conservation interest,are negativelyaffectedby heavygrazingbut also vulnerableto competitionfrom tallperennialgrasses.Thereforethe two setsofspecies alternateon lightlygrazedor protectedsites(Noy-Meir 1990). UplandBritishgrasslandspecies ofconservation valuevaryin theirresponse.Somebenefitfromremoval ofgrazing,whileothersare negativelyinfluencedbythe resultingincrease in grass (Rawes & Welch 1972). Wells (1969) commented that grazing(or mowing) duringthe season when the dominantgrassspecies is growingmostrapidlyis usuallythe mosteffectiveway to maintaindiversityin chalkgrassland.He statedthat thecessationofgrazingis themajorconservationprob- lem in those grasslands,eliminatingmanyforbsand causingincreasesoflitterandwoodyspecies.He added, however,thatgrazingshouldbe timedto avoidthesen- sitivephases in the lifecycle of species vulnerableto grazing. This raisesan importantpoint:Effectsof grazingare species-specific.That is, two species in the same com- munitymay varyin theirresponse to grazingor to a specificgrazingregime.Forexample,in an Englishhigh- elevationgrasslandon limestone,aftersheep were ex- cludedfromone site,severalrareshrubsbenefitedfrom protection,but one species declined(Elkington1981). The optimaldesignofgrazingmanagementmaythusbe difficult.Vinther(1983) foundthata mesic meadow was maintained as open meadow if it was heavily grazed-because tree seedlingswere killedby brows- ing-or ifit was not grazed at all-because seedlings couldn'testablishin the dense herblayer.Intermediate grazinglevels allowed woody regenerationand loss of the meadow's open character. Unfortunately,these same intermediatelevels ofgrazingare thosemaximiz- ingtherichnessofherbaceousspeciesintheshortterm. An alternate means of preventingwoody plant en- croachmentwould thenbe necessaryto allow contin- ued managementformaximumspecies diversity. Grazing'simpactpresentsan interestingcontrastto mowing,which is oftensuggestedas an alternativeto grazingmanagement.Mowingcan reduce thegrowthof competitivelydominantgrasses,allowing the persis-

Disturbance,Diversity,andInvasion 329

tenceofless competitivespecies,butitdoes notcreate openingsforrecruitmentof seedlingsas grazingdoes. Sykoraet al. (1990) emphasizedthe differentresultsof the two,withgrazingcreatingmore micrositesfores- tablishmentand greaterheterogeneity,while providing seed dispersalin animalfeces,hooves,and coats.As van den Bos and Bakker(1990) pointedout,grazersdo not use an entirearea evenlybut alwaysprefersome spots to others,so they create greaterheterogeneitythan does mowing.There is also a differencein the formin whichnutrientsare returnedor retainedin the system (Rizand et al. 1989). Grazingis thusan amalgamofdif- ferenteffects.Clearly,ifmowingis to be used by man- agersinpreferenceto grazing,moresophisticatedmeth- ods involvingvariationsinmowingtimeandpatternand degreeofmulchremovalshouldbe investigated. Grazinganimalsmayfrequentlybe implicatedin the invasionof naturalcommunities.Grazersmay import non-nativeplantpropagulesintonativevegetation,but theymayalso act to providemicrositesforinvasion.In particular,wheregrazingaltersthevegetationstructure or is accompaniedby soil disturbance(trampling,dig- ging,and so forth),conditionsare modifiedin such a way thatinvadingspecies can become established.For instance,Cross(1981) showed thatgrazingbythenon- nativesika deer facilitatedthe invasionof oak wood-

lands byRhododendronponticum by removing the her-

baceous understoryand providingmore safe sites for establishment.The arrivaloflargenumbersoflivestock followingEuropeansettlementhas been implicatedin thedeclineinnativeperennialgrassesand theirreplace- mentwithnon-nativeannualgrassesin severalgrassland areas in NorthAmerica and Australia(Moore 1970; Mack 1981, 1989). Braithwaiteet al. (1989) suggested thatwaterbuffaloactivitiesaid in the establishmentof Mimosa pigra in northernAustralia.Pickard(1984) im- plicatedgrazingdisturbanceas one ofthemajorfactors influencinginvasionon LordHowe Islandin the South

Pacific.

3. Soil Disturbances

In grasslands,as in most plant communitytypes,soil disturbancecreates openings for establishment,fre- quently of weedy or ruderal species. It is unclear whethertemporaryincreasesin nutrientsand otherre- sourcesaredirectlyresponsibleforthisenhancementof establishmentor whetherreduced competitionfrom neighboringplantcanopiesandrootsis moreimportant, and it is usuallydifficultto separate the two effects. Wheresuch disturbancehas longbeen a componentof theecosystem,thereis likelya substantialfractionofthe florathat is specialized or adapted to establishment there.Thus in theMediterraneanregion,where human agriculturalandotheractivityhas longcreatedsuch soil disturbance,there is a large and successfulgroup of

ConservationBiology Volume 6, No. 3, September1992

330 Disturbance,Diversity, andInvasion

weedyspecies.These are thecolonistsandinvadersthat have become so pervasivein disturbedsiteselsewhere in the world,where agriculturalactivityhas a much

shorter history and

adaptedto such a habitat(Naveh 1967; Hobbs & Hop-

kins,1990).

where few native species are

Plowing is said to diminishspecies richness,espe-

ciallythatofdicots,inlowlandgrasslands(Fuller 1987). Evenso, particularspecies mayrequireplowingto per- sist(Preston& Whitehouse1986). Smaller-scaledistur-

bances maybe

nitiesbothecological and evolutionary;forexample,in tall-grassprairies,mounds created by badger excava- tionssupporta distinctiveand diversefloraof"fugitive" prairieplants that live only on those mounds (Platt 1975). Thisdistinctivegroupofspeciescontributessub- stantiallyto the overalldiversityof thoseprairies,par- ticularlyto overgrazedones inwhichthebackgroundor matrixis relativelyspecies-poor.Otherdisturbancesby

prairiedogs,buffaloes,andgophersalso have significant effectson prairiediversity(Coppock et al. 1983; Collins

& Barber1985; Huntly& Inouye 1988; Whicker& Det-

ling 1988; Martensenet al. 1990). Moundsofbare soil formedby the activityofpocketgophersact in Califor- nianannualgrasslandsto providesubstratesforseedling establishmentin an environmentof lower densityand alteredmicroclimateand soil nutrientstatus(Hobbs & Mooney 1985; Koide et al. 1987). Coffinand Lauenroth (1988) used a modelingapproach and foundthatthe effectof soil disturbances(ant mounds and mammal burrows)on a shortgrasscommunitywas chieflya func- tionofdisturbancefrequencyand secondarilyofdistur- bance size. While soil disturbances,especiallyby animals,often have importanteffectson the dynamicsofnativeplant communities,therearealso numerousexamplesofsuch soil disturbancesfacilitatinginvasionbynon-nativespe- cies. Disturbanceby gopherswas foundby Hobbs and Mooney(1985, 1991) to be an importantfactorin the invasionofserpentinegrasslandbyBromus mollis and other non-nativeannual grasses followingyears of above-averagerainfall.Bromus mollis became estab- lishedin greaterabundanceon gophermoundsthanin undisturbedgrassland,andwas virtuallyabsentfromar- eas where gopherswere excluded.Bromus mollis was able to disperseseeds onto gophermoundsmoreeffec- tivelythan some of the nativespecies because of its

tallerinflorescence,and it thensurvivedbetteron the moreopen microhabitatthanin theundisturbedgrass- land. Experimentsinwhichartificialsoildisturbanceswere createdhave had mixedresults,withtheeffectsvarying among differentplant communities.Hobbs and Atkins (1988) found that some communities were more readilyinvaded than others,and thatsoil disturbance did not necessarilyincreasethe ease withwhich non-

equallyimportantin providingopportu-

ConservationBiology Volume6, No. 3, September1992

Hobbs& Huenneke

nativespeciescould become establishedor survive.Dis- turbancehad thelargesteffectin thecommunitiesthat were alreadymoresusceptibleto invasion. Whydoes soil disturbancefacilitateinvasion?Distur- bance mayact primarilybyprovidinga roughersurface on which seeds can lodge; in otherwords,the distur- bance increasesthe availabilityof safesites (Hobbs & Atkins1988). Hobbs and Mooney (1985) foundthat plantsofbothnativeand non-nativespecies grewmuch largeron gopher mound microhabitatsthan it undis- turbedgrassland,but Koide et al. (1987) found that nutrientavailabilitywas actually lower in gopher moundsoilsthanin undisturbedsoil.Hence removalof competitorsmaybe the majorfactorin thiscase.

4. NutrientInputs

Anothertypeofdisturbance,whichis oftenless obvious, is a changein the inputand cyclingof nutrientsin an

ecosystem.Inputofadditionalnutrients,particularlyni-

trogenand phosphorus,in low-fertilitysites can be as devastatingas eutrophicationin freshwaterecosystems. Fertilizationhas contributedto a markeddeclinein spe-

cies richnessin Britishand Dutch grasslands(Willis 1963; Bakker1987; Fuller1987). Grassesare oftenthe species to respondand to dominateundernutrienten- richment,to the detrimentofbroadleavedplants.Dur- ingandWillems(1986) suggestedthatcontinuedinput ofpollutantsand nitrogenwere partiallyresponsiblefor the floristicimpoverishmentof nonvascularflora in Dutch chalk grassland.Input of atmosphericnitrogen was apparentlyto blame forthe increasingdominance ofone grassspeciesandtheloss ofmanyforbsandother grasses,regardlessof management(mowing, grazing, burning)in chalkgrassland(Bobbink& Willems1987).

Certainlytheproblemofincreaseddepositionofnutri-

ents fromthe atmosphereis likelyto be chronic and widespread. Gough and Marrs(1990) suggestedthathighphos- phorus levels in the soil of abandoned pasturespre- cluded the reestablishmentof species-richgrassland there. Natural or successional losses of phosphorus were too slow froma managementperspective;incur- sion of scrub or woody species apparentlyincreased levels of extractablephosphorus.They suggestedthat managersuse cropping(cuttingand removingabove- ground biomass each season) or heavy leaching to lower soil-extractablephosphoruslevels more quickly. Marrs(1985) reporteda similareffortto reduce soil fertilityin a site where managerswere attemptingto reestablishan acid heathland.In anothertwenty-two- year experimentwith cutting,Rizand et al. (1989) foundthatretainingclippingson thesitekeptphospho- rus availabilityhigh,witha possible negativeinfluence on speciescomposition,comparedwithremovalofclip- pingsor withgrazing.Green(1972) pointedout early

Hobbs& Huenneke

on thatchalkgrassland,dune grassland,and heathwere all seral,low-fertilityecosystemswithhighconservation value. He suggestedmore studyofnutrientbudgetson thosesystemsandpointedoutthatgrazing,burning,and mowingall decreased the likelihoodof nutrientaccu- mulations. In North American old fields, nutrient- enrichedfieldssupportedlower species richnessand retaineda weedy annual,largelynon-nativeflora(Car- son & Barrett1988) ratherthanthe perennialgrasses typicaloffieldsofequivalentage. In ecosystems with predominantlynutrient-poor soils,additionofnutrientscan constitutea majordistur- bance,which has been shownin manyexamplesto fa- cilitateinvasionby non-nativespecies. Huennekeet al. (1990) have shown thata serpentinegrasslanddomi- natedby annualforbscan be transformedin two years intoone dominatedby non-nativegrassesby the addi- tionofnutrients,particularlynitrogenand phosphorus. Hobbs et al. (1988) produced similar results and showed thatsurvivalof non-nativegrasseswas signifi- cantlyenhancedon fertilizedplots,while thatofnative forbswas reduced.In boththesecases,invasionwas not relateddirectlyto soil disturbanceand,infact,Hobbs et al. (1988) foundthatsubsequent gopher disturbance actuallyreduced the dominanceof non-nativegrasses and allowed the re-establishmentofnativeforbs. Nutrientinputhas also been shownto facilitateinva- sionofAustralianplantcommunities.Heddle andSpecht (1975) reportedincreased abundances of non-native herbaceous species in areas of heathlandthathad re- ceived fertilizer.Otherstudieshave indicateda strong relationshipbetween the degree of invasionby non- nativespecies and soil nutrientlevels, particularlyof phosphorus (Cale & Hobbs 1991; Hester & Hobbs 1992). Experimentswhere nutrientswere added to plotswithina numberofdifferentplantcommunitiesin WesternAustraliashowed thatincreasednutrientsre- sulted in increased growthof non-nativespecies in some plantcommunitiesbut not others(Hobbs & At- kins1988). Ofparticularinterestwas thefindingthata combinationof soil disturbanceand nutrientaddition had the greatesteffectin enhancingthe establishment and growthofnon-nativespecies.

5. Trampling

Like the other disturbanceswe have discussed,tram- pling can create openings in vegetationthatprovide opportunitiesfor new individualsto become estab- lished,and it can slow the growthofdominantspecies sufficientlyto allow thepersistenceoflessvigorousspe- cies. Again,intermediatelevels oftramplingseem most effectiveatmaintaininghighspeciesrichnessbecause of the suppression of competitive dominants (Liddle 1975). The season or timingoftramplinghas a signifi- canteffecton thechance,rate,and species composition

Disturbance, Diversity, and Invasion

331

ofrecovery(Harrison1981). Thereare species-specific responsesto trampling:in one studymost but not all species were negativelyaffected(Crawford& Liddle 1977): invertebratesseemfarmoresensitivethanplants (Duffey1975). We have encounteredlittleinformation on theeffectsoftramplingon invasions,althoughtram- pling effectsare frequentlyconsidered togetherwith thoseofgrazing.

6. Fragmentation

The fragmentationand insularizationof ecosystemsis nota disturbancewithinan individualsystembuta land- scape-leveldisturbanceresultingin the rearrangement ofthelandscapematrix.Byinfluencingedge effectsand

the likelihoodof movementof nutrients,propagules, and faunafromadjacentpatches,fragmentationaffects disturbanceregimesin individualpatches of remnant vegetation(Hobbs 1987; Saunderset al. 1991). How does fragmentationaffectthe species compositionand richnessof grasslands?Simberloffand Gotelli (1984) surveyedpatches of prairieand foundthat"archipela- goes" ofsmallgrasslandpatchessupportedmorespecies thandid singlelarge patches of equivalenttotal area. Thus smallpatch size does not constraintotalspecies richness.QuinnandRobinson(1987) andRobinsonand Quinn (1988) used an experimentalapproach to this question, subdividing annual grassland into fenced patchesseparatedbyheavilygrazedzones; species rich- ness was substantiallyhigherin the more subdivided treatments.Singlespecies frequentlycame to dominate single plots, so a region with a greater number of patchessupportedboth more dominantspecies (alter- nate dominantsin differentplots) and more edge spe- cies (growing along the greaterperimeter).Murphy

(1989) has pointed out

(1988) studywas carriedout at an inappropriatescale and in a grasslandthatis dominatedby non-nativean- nuals.However,thepointthatfragmentationwilllead to an increasein edge species is important.Froma conser- vation managementperspective,one would want to knowjustwhich species are beingfavoredby edge ef- fects.A highertotalspecies richnesscould be primarily due to an increasednumberofruderalorweedyspecies of low conservationvalue (as found,forexample for invertebratesby Webb & Hopkins [1984]), or to a highernumberoflegitimatecommunitymembers.

that Robinson and Quinn's

7. InteractionofDisturbances

Ofcourse,mostecosystemsexperiencemultipledistur-

bances and are shaped by multiplefactors.In many cases the resultsare not merelyadditive,and distur- bances can act synergistically.For example,grazingre- duced fuelloads, reduced firefrequency,and allowed the invasionof woody species into many regions of semi-aridgrassland(such as the historicalexpansionof

ConservationBiology Volume 6, No. 3, September1992

332 Disturbance,DiversityandInvasion

pinyon-junipervegetationintowesternU.S. grasslands; Wrightet al. 1979). In an experimentalstudy,Collins (1987) foundthatfiresignificantlyincreased species diversityin grazedtallgrassprairiebutnoton ungrazed grassland;insomerespectstheeffectsofgrazingandfire were additive.Collinsand Gibson(1990) have further illustratedhow grazing,fire,and small-scalesoil distur- bance all affectthematrixstructureofthesegrasslands differently,and hence can interactto increasecommu- nitydiversity.Leighet al. (1987) foundthatrabbitpop- ulationsincreasedon burnedareas ofsubalpinevegeta- tion, while Noy-Meir (1988) found that elevated populationsof voles had the greatesteffectson grass- landswhere othergrazingwas minimized.Sykoraet al. (1990) suggestedthatfireinDutchgrasslandsincreased nutrientsand thusincreasedthe likelihoodof"ruderal- ization"-increasingdominancebya fewgrassesleading to a decline in diversity.Hodgkin(1984) foundthat woody encroachment increased soil fertilityand changed the natureof Britishdune grassland.It was suggestedthatthe myxomatosis-causeddecline in rab- bitpopulationshad resultedin the increasedestablish- mentofwoody vegetation,and thattheresultingscrub had increased soil nutrientsto the point thatweedy plantspecies were favored. Invasion by the nitrogen-fixingMyrica faya onto younglava flowsin Hawaii has been shownto alterthe natureof ecosystemdevelopmentfollowingvolcanic eruptions(Vitousek et al. 1987; Vitousek & Walker 1989). In thisand othercases, such as thatofMimosa pigra inAustralia(Braithwaiteet al. 1989), theinvading plantsthemselvesconstitutea majordisturbanceto the systemstheyare invading.

GoodDisturbancesTurnedBad:Conflicts

Aretherecases where disturbanceis a necessarycom- ponentofecosystemand communitydynamics,butalso enhancesthelikelihoodofinvasion?Fromtheforegoing, itwould seem thatvirtuallyanytypeofdisturbancecan

facilitateinvasionundercertaincircumstances.Invasion is,afterall,simplya subsetofthepossiblerecolonization

response to disturbance.As an example,Griffinet

al.

(1989) have shownhow periodicfloodingcan lead to theinvasionofaridzone riversystemsbyTamarixaph- ylla It is not the typeofdisturbancebutrathercertain aspectsofitsactionin a particularsystemthatshiftthe resulttowardenhancementofinvasionsat theexpense ofnatives(see McIntyreet al. 1988). For example,it is not fireper se but the combinationof firewith other disturbance,or the adoptionof a fireregimeinappro- priateto the lifehistoriesof nativeplants,thatfavors non-nativefire-tolerantspecies at the expense of na- tives.Aprimaryconsideration,then,mustbe thesuiteof adaptationsand lifehistoriesfoundin the nativeplants, particularlythose ofconservationvalue.

ConservationBiology Volume6, No. 3, September1992

Hobbs& Huenneke

The relationshipbetween soil disturbanceand inva- sionis also complex,and mechanicaldisturbancein the absence ofnutrientadditionmaynotnecessarilylead to enhancedinvasion(see Hobbs 1989). Frequently,how- ever,physicaldisturbanceand nutrientenrichmentco- incide,as when rabbitsscrape the soil and defecateat thesame time,or when disturbanceenhancesnitrogen mineralization.Animportantproblemforsystemswitha naturallylow nutrientstatusis the gradualnutrienten- richmentthatcan occur via atmosphericinput,wind- blown fertilizer,or inputfromlivestockfeces (Lands- berg et al. 1990; Cale & Hobbs 1991). An increased baselinenutrientstatuswillhaveimportantimplications forthewhole ecosystem,but in the shorttermit may exacerbatethe likelihoodof invasionsby weedy pest species.

Conclusions

No systemcan remain immune fromcertain distur- bances(such as nutrientinputfromtheatmosphere);in thefuture,fewareaswill even be protectedfromdirect humanactivity.Somedisturbancetypescan be modified byon-sitemanagement(fireandgrazingregimes)while

otherscannot(floods,storms).Human-induceddistur-

bancessuchas roadconstructioncan also be minimized.

"Naturaldisturbanceregimes"maybe desirablebut are oftenimpracticablein thealteredsettingsofcontempo- raryreserves.We need to acknowledgethe actual dis- turbanceregimeoperatingcurrentlyin a reserve,and the currentpropagulerain,which determinesthe im-

portanceofcopingwithlikelyinvasions.Further,man-

agersneed to take an active role in designingthe dis-

turbanceregime,tailoringitto thelandscape,thebiotic community,and theirspecificconservationgoals. Denslow (1980) hypothesizedthatanynaturalcom- munitywould be richestin species adaptedto establish- mentin the typeofpatch mostcommonlycreatedby disturbance.For example, where large scale distur- bances are the norm,mostspecies will establishthere and species richnesswill decline throughtimeand suc- cession.In contrast,in an ecosystemwhere small-scale disturbancesare normal,mostspecies will establishin smallscale gaps or in undisturbedsites,and diversity will increasewithtimeaftera largedisturbance.Total diversityofnativespecies at thelandscapelevelwill be greatestwhen disturbanceoccurs at its historicalfre- quencyandin thehistoricalpattern(Fig. 2). Changesin thesize ofthefrequency,as well as the type,ofdistur- bance willmeanthatmostnativespecieswill no longer be well adaptedforrecruitmentor establishment. In addition,evenwhen disturbanceregimeshave not been significantlyaltered,the availabilityof weedy or invasivespecies may alter systemresponse to distur- bance. Managementmustconsidernot onlyalterations

Hobbs& Huenneke

Natural Disturbance Regime maintains native species diversity

(historical

decrease

ina

/

frequency/

type, frequency, intensity of disturbance)

chafrge

i'r

ty, se

increase

\

ina

frequency/

intensity

of disturbance

intensity

Decreased

Elimination

Elimination

diversity

of natives;

of natives;

of natives

Enhancement

Enhancement

(dominance

of

of invasions

of invasions

competitively

(direct damage to

(direct damage to

superior species)

natives: creation of new microsites)

natives; creation of new microsites)

Figure2. Anychange in thehistoricaldisturbance regimeof an ecosystemmay alterspecies composi- tion by reducingtheimportanceof nativespecies,by creatingopportunitiesfor invasivespecies,or both.

oftheoriginaldisturbanceregime,butalso alterationin the pool of potential responding species (in other words,the availabilityofcolonistsor pests). The responseofinvadersto disturbanceis an extreme case ofan underlying,unavoidableconflict-anydistur- bance and any managementregimewill be good for some species and bad forothers.The decision maybe easy(althoughthetechniquesformanagementmaynot be) when thechoice is betweennativesand non-native pest species.The dilemmais thornierwhen non-natives have some appeal oftheirown (forinstancein termsof grazingvalue), and it is stillmore difflcultwhen the choice is betweenone setofnativespecies andanother. In theend,thewisestchoice maybe to use a diversity of managementstrategies,to encourage differentspe- cies in differentpartsof the reserveor in differentre- serveswithina region.There is no singleoptimalstrat- egy;managersmustmakedecisionsbased on thelikely costs and benefitsin termsofmaintenanceofdiversity versusinvasionby non-natives. We drawthefollowingconclusionsfromthisreview of scientificresearchon disturbanceand species com- positionand diversityin grasslands.

* Disturbanceplays an integralrole in structuring plantcommunities,butsome typesor combinations ofdisturbancecan increasethepotentialofinvasion by non-nativespecies.

* Backgroundlevelsofdisturbance,resourceavailabil- ity,and thepool ofpotentialspecies in anyecosys- temall differnow fromprimevalcondition.This is true even in the largestparks and reserves (see Chase 1987). Itis notenoughto saythattheoriginal disturbanceregimeis the desiredstate.

* Speciesvaryintheirresponseto disturbance,requir-

Disturbance,Diversity,andInvasion 333

ing managersto make deliberatechoices ofwhich taxa to favor.

* Managersmayhaveto choose betweenspecificcon- servationtargets,such as preventingthe spread of invasivespecies,and themoregeneralgoal ofmain- tainingoverallspecies diversity.

* Nearlyall systemsare likelyto be nonequilibrialin the future;we must be activistsin determining which species to encourageand which to discour- age. We cannotjust managepassively,or formaxi- mal diversity,but mustbe selectiveand tailorman- agementto specificgoals.

Acknowledgments

We thankJuliArmstrong,RichardGroves,and two ref- erees for constructivecommentson the draftmanu- script.

LiteratureCited

Abrams,M.D., A.K Knapp,and L.C. Hulbert.1986. A ten-year record of abovegroundbiomass in a Kansas taligrassprairie:

effectsoffireand topographicposition.AmericanJournalof

Botany73:1509-1515.

Bakker,J.P. 1987. Restorationofspecies-richgrasslandaftera periodoffertiliserapplication.Pages 185-200 inJ.vanAndel, J.P. Bakker,and R.W. Snaydon,editors.Disturbancein grass- lands:causes,effectsand processes.Junk,Dordrecht.

Bobbink,R.,andJ.H. Willems.1987. Increasingdominanceof Brachypodiumpinnatum in chalk grasslands:a threatto a

species-richecosystem.BiologicalConservation40:301-314.

Braithwaite,R.W., W.M. Lonsdale,and J.A. Estbergs.1989. Alienvegetationand nativebiota in tropicalAustralia:theim- pact ofMimosa pigra BiologicalConservation48:189-210.

Bulow-Olsen,A. 1980. Changesin thespecies compositionin an area dominatedby Deschampsia flexuosa as a resultof

cattlegrazing.BiologicalConservation18:257-270.

Caldwell,M.M.,J.H. Richards,D. A.Johnson,R.S. Nowak,and R.S. Dzurec. 1981. Copingwithherbivory:photosyntheticca- pacity and resource allocation in two semiaridAgropyron bunchgrasses.Oecologia 50:14-24.

Cale, P., and R.J.Hobbs. 1991. Conditionofroadsidevegeta- tioninrelationto nutrientstatus.Pages 353-362 inD. A.Saun- dersandR.J.Hobbs,editors.Natureconservation2: theroleof corridors.Surrey-Beatty,ChippingNorton,Australia.

Carson,W. P.,and G.W. Barrett.1988. Successionin old-field

plantcommunities:effectsofcontrastingtypesofnutrienten-

richment.Ecology69:984-994.

Chase,A. 1987. Playinggod inYellowstone.The destructionof America'sfirstnationalpark.HarcourtBraceJovanovich,New York.

ConservationBiology Volume 6, No. 3, September1992

334 Disturbance,Diversity, andInvasion

Christensen,P. E., and N. D. Burrows.1986. Fire:an old tool witha new use. Pages 57-66 in R.H. GrovesandJ.J.Burdon,

editors.Ecologyofbiologicalinvasions:anAustralianperspec-

tive.AustralianAcademyofScience,Canberra,Australia.

Coblentz,B. E. 1978. The effectsofferalgoats(Capra hircus) on islandecosystems.BiologicalConservation13:279-286.

Coffin,D. P., and W. K Lauenroth.1988.The effectofdistur- bance size and frequencyon a shortgrassplant community.

Ecology69:1609-1617.

Collins,S. L. 1987. Interactionofdisturbancesintallgrassprai- rie:a fieldexperiment.Ecology68:1243-1250.

Collins,S. L.,and S. C. Barber.1985. Effectsofdisturbanceon diversityin mixed-grassprairie.Vegetatio64:87-94.

Collins,S. L.,and D. J.Gibson.1990. Effectsoffireon commu- nitystructurein tallgrassandmixed-grassprairie.Pages 81-98 in S. L. Collinsand L.L. Wallace,editors.Firein NorthAmeri- can tallgrassprairies.Universityof OklahomaPress,Norman, Oklahoma.

Connell,J.H. 1978. Diversityin tropicalrainforestsand coral reefs.Science 199:1302-1310.

Coppock, D. L.,J.K Detling,J.E. Ellis,and M. I. Dyer. 1983. Plant-herbivoreinteractionsin a NorthAmericanmixed-grass prairie1. Effectsofblack-tailedprairiedogs on intraseasonal abovegroundplantbiomassandnutrientdynamicsandspecies diversity.Oecologia (Berlin) 56:1-9.

Cowling,R.M.,S.M. Pierce,and E.J.Moll. 1986. Conservation and utilizationof South Coast Renosterveld,an endangered South Africanvegetationtype. Biological Conservation37:

363-377.

Crawford,A.K, and M.J.Liddle.1977. The effectoftrampling on naturalgrassland.BiologicalConservation12:135-142.

Cross,J.R. 1981. The establishmentofRhododendronponti- cum in the Killarnyoakwoods,S.W. Ireland.JournalofEcol-

ogy69:807-824.

Curtis,J.T., and M.L. Partch.1948. Effectoffireon the com- petitionbetween blue grassand certainprairieplants.Amer- ican MidlandNaturalist39:437-443.

Denslow,J.S. 1980. Patternsofplantspecies diversityduring successionunderdifferentdisturbanceregimes.Oecologia 46:

18-21.

Dolman,P., and W. Sutherland.1991. Historicalclues to con- servation.New Scientist1749:22-25.

Duffey,E. 1975. The effectsofhumantramplingon thefauna

ofgrasslandlitter.BiologicalConservation7:255-274.

During,H.J.,andJ.H. Willems.1986. The impoverishmentof thebryophyteandlichenfloraoftheDutchchalkgrasslandsin

thethirtyyears1953-1983. BiologicalConservation36:143-

158.

Elkington,T. T. 1981. Effectsof excluding grazinganimals fromgrasslandon sugarlimestonein Teesdale, England.Bio- logical Conservation20:25-35.

ConservationBiology Volume6, No. 3, September1992

Hobbs& Huenneke

Ewel,J.1986. Invasibility:lessons fromSouth Florida.Pages 214-230 in H.A. MooneyandJ.A. Drake,editors.Ecologyof biological invasionsof NorthAmericaand Hawaii. Springer- Verlag,New York.

Fox,J.F. 1979. Intermediate-disturbancehypothesis.Science

204:1344-1345.

Fuller,R.M. 1987. The changingextentand conservationin- terestoflowlandgrasslandsinEnglandand Wales:a reviewof grasslandsurveys1930-84. BiologicalConservation40:281-

300.

Gibson,C. W. D., T. A.Watt,andV.K Brown.1987. The use of sheep grazingto recreatespecies-richgrasslandfromaban- doned arableland.BiologicalConservation42:165-183.

Glenn-Lewin,D. C., and J.M. ver Hoef. 1988. Scale, pattern analysis,and species diversityin grasslands.Pages 115-129 in H.J.During,M.J.A. Werger,andJ.H. Willems,editors.Diver- sityandpatternin plantcommunities.SPB AcademicPublish- ing,The Hague,The Netherlands.

Gough,M.W., and R.H. Marrs.1990. A comparisonof soil fertilitybetween semi-naturaland agriculturalplantcommu- nities:implicationsforthecreationofspecies-richgrasslandor abandonedagriculturalland. Biological Conservation51:83-

96.

Green,B. H. 1972. The relevanceof seral eutrophicationand plantcompetitionto themanagementofsuccessionalcommu-

nities.BiologicalConservation4:378-384.

Griffin,G.F.,D. M. StaffordSmith,S. R.Morton,G.E. Allan,and

K A. Masters.1989. Statusand implicationsofthe invasionof

tamarisk(Tamarix aphylla) on theFinkeRiver,NorthernTer- ritory,Australia.Journalof EnvironmentalManagement29:

297-315.

Grime,J.P. 1979. Plant strategiesand vegetationprocesses. Wiley,New York.

Grubb,P.J. 1976. A theoreticalbackgroundto the conserva- tionofecologicallydistinctgroupsofannualsand biennialsin

thechalkgrasslandecosystem.BiologicalConservation10:53-

76.

Hamann,0. 1975. Vegetationalchangesin the Galapagos Is- lands duringthe period 1966-73. BiologicalConservation7:

37-59.

Hamann,0. 1979. Regenerationofvegetationon SantaFe and

PintaIslands,Galapagos,aftertheeradicationofgoats.Biolog-

ical Conservation15:215.

Harrison,C. 1981. Recoveryof lowlandgrasslandand heath- land in southernEnglandfromdisturbanceby seasonal tram-

pling.BiologicalConservation19:119-130.

Heady,H. F. 1972. Burningand the grasslandsin California. ProceedingsoftheTwelfthAnnualTall TimbersFireEcology Conference.

Heddle, E. M., and R. L. Specht. 1975. Dark Island Heath (Ninety-MilePlain,SouthAustralia).VIII. The effectsoffertil- izerson compositionandgrowth.AustralianJournalofBotany

23:151-164.

Hobbs& Huenneke

Hester,A.J.,and R.J.Hobbs. 1992. Influenceoffireand soil nutrientson nativeand non-nativeannualsat remnantvege- tationedges in the WesternAustralianwheatbelt.Journalof VegetationScience. 3:101-108.

Hobbs,R.J.1987. Disturbanceregimesin remnantsofnatural vegetation.Pages 233-240 in D.A. Saunders,G.W. Arnold,

A.A. Burbidge,and A.J.M. Hopkins,editors.Natureconserva- tion:the role ofremnantsofnativevegetation.SurreyBeatty,

ChippingNorton,Australia.

Hobbs,R.J. 1989. The natureand effectsofdisturbancerela- tiveto invasions.Pages 389-405 inJ.A. Drake,H.A. Mooney,

F. di Castri,R.H. Groves,F.J. Kruger,M. Rejmanek,and M.

Williamson,editors.Biologicalinvasions.A globalperspective.

Wiley,Chichester,England.

Hobbs,R.J.1991. Disturbanceas a precursortoweed invasion in nativevegetation.PlantProtectionQuarterly6:99-104.

Hobbs,R.J.,and L.Atkins.1988. The effectofdisturbanceand nutrientadditionon native and introducedannuals in the WesternAustralianwheatbelt.AustralianJournalof Ecology

13:171-9.

Hobbs, R.J.,and L. Atkins.1990. Fire-relateddynamicsof a Banksia woodlandinsouth-westWesternAustralia.Australian

JournalofBotany38:97-110.

Hobbs,R.J.,and A.J.M. Hopkins.1990. Fromfrontierto frag- ments:Europeanimpacton Australia'svegetation.Proceedings ofthe Ecological SocietyofAustralia16:93-114.

Hobbs,R.J.,and H.A. Mooney. 1985. Communityand popu- lationdynamicsofserpentinegrasslandannualsin relationto gopherdisturbances.Oecologia (Berlin) 67:342-351.

Hobbs, R.J.,and H.A. Mooney. 1991. Effectsof rainfallvari- abilityand gopherdisturbanceon serpentineannualgrassland dynamicsin N. California.Ecology72:59-68.

Hobbs,R.J.,S. L.Gulmon,V.J.Hobbs,and H.A.Mooney.1988. Effectsof fertilizeradditionand subsequent gopher distur- bance on a serpentineannualgrasslandcommunity.Oecologia (Berlin) 75:29 1-295.

Hobbs,R.J.,A.U. Mallik,and C. H. Gimingham.1984. Studies on firein Scottishheathlandcommunities.III. Vitalattributes ofthe species.JournalofEcology72:963-976.

Hodgkin,S. E. 1984. Scrubencroachmentanditseffectson soil fertilityon NewboroughWarren,Anglesey,Wales. Biological

Conservation29:99-119.

Hopkins,A., and J. Wainwright.1989. Changes in botanical compositionand agriculturalmanagementof enclosed grass- landinuplandareasofEnglandandWales,1970-86, andsome conservationimplications.Biological Conservation47:219-

235.

Huenneke,L.F., S.P. Hamburg,R. Koide, H.A. Mooney,and

P. M.Vitousek.1990. Effectsofsoilresourceson plantinvasion

and communitystructurein Californianserpentinegrassland.

Ecology71:478-491.

Disturbance,Diversity, andInvasion 335

Hughes,R.F.,P. M.Vitousek,andJ.T. Tunison.1991. Effectsof invasionbyfire-enhancingC4 grasseson nativeshrubsin Ha- waii Volcanoes NationalPark.Ecology72:743-746.

Huntly,N., and R. Inouye. 1988. Pocket gophersin ecosys- tems:patternsand mechanisms.BioScience 38:786-793.

Huston,M. 1979. A generalhypothesisof species diversity.

AmericanNaturalist113:81-101.

Johnstone,I. M. 1986. Plantinvasionwindows: a time-based

classificationofinvasionpotential.BiologicalReviews61:369-

394.

Knapp,A.K, and T. R. Seastedt.1986. Detritusaccumulation

limitsproductivityoftallgrassprairie.BioScience36:662-668.

Koide, R., L.F. Huenneke,and H.A. Mooney. 1987. Gopher moundsoil reduces growthand affectsion uptakeoftwo an- nual grasslandspecies. Oecologia (Berlin) 72:284-290.

Kucera,C. L.,and M. Koelling.1964. The influenceoffireon compositionof centralMissouriprairie.AmericanMidland

Naturalist72:142-147.

Landsberg,J.,J.Morse,and P. Khanna.1990. Tree diebackand insect dynamicsin remnantsof nativewoodlands on farms.

ProceedingsoftheEcologicalSocietyofAustralia16:149-165.

Leigh,J.H., D. J.Wimbush,D. H. Wood, M.D. Holgate,A.V.

Slee, M.G. Stanger,and R.I. Forrester.1987. Effectsofrabbit grazingand fireon a subalpineenvironment.I. Herbaceous and shrubbyvegetation.AustralianJournalofBotany35:433-

464.

Lewin,R. 1984. Parks:how big is big enough?Science 225:

611-612.

Liddle,M.J.1975. A selectivereviewofthe ecological effects ofhumantramplingon naturalecosystems.BiologicalConser-

vation7:17-36.

Loney,B., and R.J.Hobbs. 1991. Establishment,maintenance and rehabilitationof vegetationcorridors.Pages 299-311 in D. A. Saundersand R.J.Hobbs,editors.Natureconservation2:

therole ofcorridors.Surrey-Beatty,ChippingNorton,Austra- lia.

MacDonald,I.A.W.,L.L. Loope, M.B. Usher,and 0. Hamann.

1989. Wildlifeconservationand the invasionof nature re-

servesbyintroducedspecies:a globalperspective.Pages 215-

255 inJ.A. Drake,H.A. Mooney,F. di Castri,R.H. Groves,F.J.

Kruger,M. Rejm'anek,and M. Williamson,editors.Biological invasions:a globalperspective.Wiley,Chichester,England.

Mack,R.N. 1981. InvasionofBromus tectorumL. intowest- ernNorthAmerica:an ecological chronicle.Agro-Ecosystems

7:145-165.

Mack,R.N. 1989. Temperategrasslandsvulnerableto plant invasions:characteristicsandconsequences.Pages 155-179 in J.A.Drake,H.A.Mooney,F.di Castri,R.H. Groves,F.J.Kruger, M. Rejmanek,and M. Williamson,editors.Biologicalinvasions:

a globalperspective:Wiley,Chichester,England.

ConservationBiology Volume6, No. 3, September1992

336 Disturbance,Diversity,andInvasion

Mack,R.N., and J.N. Thompson. 1982. Evolutionin steppe with few large,hooved mammals.AmericanNaturalist119:

757-773.

Marrs,R.H. 1985. Techniques forreducingsoil fertilityfor natureconservationpurposes:a reviewinrelationto research

atRoper'sHeath,Suffolk,England.BiologicalConservation34:

307-332.

Martensen,G. D.,J.H. Cushman,andT. G. Whitham.1990. Im- pactofpocketgopherdisturbanceon plantspeciesdiversityin a shortgrassprairiecommunity.Oecologia (Berlin) 83:132-

138.

McIntyre,S.,P.Y. Ladiges,and G. Adams.1988. Plantspecies- richnessand invasionby exoticsin relationto disturbanceof wetlandcommunitieson the RiverinePlain,NSW.Australian JournalofEcology 13:361-373.

Milchunas,D. G., W. K Lauenroth,P. L. Chapman,and M.K Kazempour.1990. Communityattributesalonga perturbation gradientin a shortgrasssteppe.JournalofVegetationScience

1:375-384.

Milchunas,D. G.,0. E. Sala,and W.K Lauenroth.1988. A gen- eralizedmodel oftheeffectsofgrazingbylargeherbivoreson

grasslandcommunitystructure.AmericanNaturalist132:87-

106.

Moore,A.D., and I. R.Noble. 1990. Anindividualisticmodelof

vegetationstanddynamics.JournalofEnvironmentalManage-

ment31:61-81.

Moore,R.M. 1970. Australiangrasslands.AustralianNational UniversityPress,Canberra,Australia.

Murphy,D. D. 1989. Conservationand confusion:wrongspe- cies, wrong scale, wrong conclusions.ConservationBiology

3:82-84.

Naveh, Z. 1967. Mediterraneanecosystemsand vegetation typesin Californiaand Israel.Ecology48:445-459.

Naveh,Z., and R.H. Whittaker.1980. Structuraland floristic diversityofshrublandsand woodlands in northernIsrael and

otherMediterraneanareas.Vegetatio41:171-190.

Noble,I. R.,and R.0. Slatyer.1980. The use ofvitalattributes to predictsuccessionalchangesin plantcommunitiessubject to recurrentdisturbances.Vegetatio43:5-21.

Noy-Meir,I. 1988. Dominantgrassesreplacedbyruderalforbs in a vole yearin undergrazedMediterraneangrasslands.Jour- nal ofBiogeography15:579-587.

Noy-Meir,I. 1990. The effectofgrazingon the abundanceof wild wheat,barleyand oat in Israel.BiologicalConservation

51:299-310.

Panetta,F.D., and A.J.M. Hopkins.1991. Weeds in corridors:

invasionand management.Pages 341-351 in D.A. Saunders and R.J. Hobbs, editors.Natureconservation2: the role of corridors.Surrey-Beatty,ChippingNorton,Australia.

Peet,R.K, D. C. Glenn-Lewin,andJ.WalkerWolf.1983. Pre- dictionofman'simpacton plantspecies diversity.Pages 41-

ConservationBiology Volume6, No. 3, September1992

Hobbs& Huenneke

54 inW. Holzner,M.J.A.Werger,and I. Ikusima,editors.Man's impacton vegetation.Junk,The Hague,The Netherlands.

Petraitis,P. S.,R.E. Latham,and R.A. Niesenbaum.1989. The

maintenanceofspeciesdiversitybydisturbance.QuarterlyRe-

view ofBiology64:393-418.

Pickard,J.1984. Exoticplantson LordHowe Island:distribu- tionin space and time.JournalofBiogeography11:181-208.

Pickett,S.T. A.,andJ.N. Thompson.1978. Patchdynamicsand

thedesignofnaturereserves.BiologicalConservation13:27-

37.

Pickett,S.T. A.,and P. S. White,editors.1985. The ecologyof naturaldisturbanceand patchdynamics.AcademicPress,Or- lando,Florida.

Pickett,S.T. A.,J.Kolasa,J.J.Armesto,and S.L. Collins.1989. The ecological concept of disturbanceand its expressionat varioushierarchicallevels.Oikos 54:129-136.

Platt,W.J. 1975. The colonisationand formationof equilib- riumplant species associationson badger disturbancesin a tall-grassprairie.Ecological Monographs45:285-305.

Preston,C. D., and H. L.K Whitehouse.1986. The habitatof Lythrumhyssopifoliumin Cambridgeshire,itsonlysurviving

Englishlocality.BiologicalConservation35:41-62.

Puerto,A.,M. Rico,M.D. Matias,andJ.A. Garcia. 1990. Vari- ation in structureand diversityin Mediterraneangrasslands relatedto trophicstatusand grazingintensity.JournalofVeg- etationScience 1:445-452.

Quinn,J.F., and G.R. Robinson.1987. The effectsof experi- mentalsubdivisionon floweringplantdiversityin a California

annualgrassland.JournalofEcology75:837-855.

Ranwell, D. S. 1960. Newborough Warren,Anglesey. III. Changesin vegetationon partsof the dune systemafterthe loss of rabbitsby myxomatosis.Journalof Ecology 48:385-

397.

Rawes,M.,and D. Welch. 1972. Trialsto recreatefloristically rich vegetationby plant introductionin the NorthernPen-

nines,England.BiologicalConservation4:135-140.

Rejm'anek,M. 1989. Invasibilityof plant communities.Pages 369-388 inJ.A.Drake,H.A.Mooney,F.di Castri,R.H. Groves, F.J.Kruger,M. Rejm'anek,and M. Williamson,editors.Biolog- ical invasions:a global perspective.Wiley,Chichester,En- gland.

Rizand,A.,R.H. Marrs,M.W. Gough,and T. C. E. Wells. 1989.

Long-termeffectsofvariousconservationmanagementtreat-

mentson selectedsoilpropertiesofchalkgrassland.Biological

Conservation49:105-112.

Robinson,G.R., and J.F. Quinn. 1988. Extinction,turnover and species diversityin an experimentallyfragmentedCalifor- nia annualgrassland.Oecologia (Berlin) 76:71-82.

Rykiel,E.J. 1985. Towards a definitionof ecological distur- bance. AustralianJournalofEcology10:361-365.

Hobbs& Huenneke

Saunders,D. A.,R.J.Hobbs, and C. R. Margules.1991. Biolog- ical consequences of ecosystemfragmentation.Conservation

Biology5:18-32.

Schofield,E. K 1989. Effectsofintroducedplantsand animals on islandvegetation:examplesfromthe GalapagosArchipel- ago. ConservationBiology3:227-238.

Simberloff,D., andN. Gotelli.1984. Effectsofinsularizationon species richnessin theprairie-forestecotone. BiologicalCon-

servation29:27-46.

Smart,N. 0. E., J.C. Hatton,and D. H. N. Spence. 1985. The effectoflong-termexclusionoflargeherbivoreson vegetation in MurchisonFalls NationalPark,Uganda.BiologicalConser-

vation33:229-245.

Soule, M.E. 1990. The onslaughtof alien species, and other challengesin the coming decades. ConservationBiology4:

233-239.

Sousa,W.P. 1984. The role ofdisturbancein naturalcommu-

nities.AnnualReviewofEcologyandSystematics15:353-391.

Strang,R.M. 1973. Bushencroachmentand veld management in south-centralAfrica:the need fora reappraisal.Biological

Conservation5:96-104.

Sykora,K V.,G. van der Krogt,andJ.Rademakers.1990. Veg- etationchangeon embankmentsin the southwesternpartof theNetherlandsundertheinfluenceofdifferentmanagement practices(in particularsheep grazing).Biological Conserva-

tion52:49-81.

Turner,M. G. editor. 1987. Landscape heterogenityand dis- turbance.Springer,New York.

van Andel,J.,and J.P. van den Bergh. 1987. Disturbanceof grasslands.Outlineoftheme.Pages 3-13 inJ.vanAndel,J.P. Bakker,and R.W. Snaydon,editors.Disturbancein grasslands:

causes,effectsand processes.Junk,Dordrecht.

van den Bos, J.,and J.P. Bakker.1990. The developmentof vegetationpatternsbycattlegrazingatlow stockingdensityin

theNetherlands.BiologicalConservation51:263-272.

van der Maarel,E. 1971. Plantspecies diversityin relationto management.Pages 45-63 in E. DuffeyandA.S. Watt,editors. The scientificmanagementof animaland plantcommunities

forconservation.BlackwellScientificPublications,Oxford,En-

gland.

Vinther,E. 1983. Invasion of Alnus glutinosa in a former grazedmeadow in relationto differentgrazingintensities.Bi- ological Conservation25:75-89.

Disturbance,Diversity,andInvasion

337

Vitousek,P. M. 1986. Biologicalinvasionsandecosystemprop- erties:can species makea difference?Pages 163-176 in H.A. Mooney and J.A. Drake,editors.Ecology of biological inva- sionsofNorthAmericaand Hawaii.Springer,New York.

Vitousek,P. M.,and L.R. Walker.1989. Biologicalinvasionby Myricafaya in Hawaii:plantdemography,nitrogenfixation, ecosystemeffects.Ecological Monographs59:247-265.

Vitousek,P. M., L.R. Walker,L. D. Whiteaker,D. Mueller- Dombois, and P. A. Matson. 1987. Biological invasion by Myricafayaaltersecosystemdevelopmentin Hawaii.Science

238:802-804.

Webb,N. R.,and P.J.Hopkins.1984. Invertebratediversityon fragmentedCalluna heathland.JournalofAppliedEcology21:

921-933.

Wells,T. C. E. 1969. Botanicalaspectsofconservationmanage-

mentofchalkgrasslands.BiologicalConservation2:36-44.

Westman,W.E. 1990. Parkmanagementof exotic plantspe- cies: problemsand issues.ConservationBiology4:251-259.

Westoby,M.,B. Walker,and I. Noy-Meir.1989. Rangemanage- menton thebasisofa modelwhichdoes notseek to establish

equilibrium.JournalofAridEnvironments17:235-240.

Whicker,A.D., and J.K Detling. 1988. Ecological conse- quences ofprairiedog disturbances.BioScience 38:778-784.

White,D. J.B. 1961. Some observationson the vegetationof BlakeneyPoint,Norfolk,followingthe disappearanceof rab- bitsin 1954. JournalofEcology49:113-118.

White,P. S.,and S.T. A. Pickett.1985. Naturaldisturbanceand patchdynamics:an introduction.Pages 3-13 in S.T. A. Pickett and P. S. White,editors.The ecology of naturaldisturbance and patchdynamics.AcademicPress,Orlando,Florida.

Willis,A.J. 1963. BrauntonBurrows:the effectson the vege- tationofthe additionofmineralnutrientsto dune soils.Jour- nal ofEcology51:353-374.

Wright,H.A.,L.F. Neuenschwander,and C. M. Britton.1979. The role and use offirein sagebrush-grassandpinyon-juniper plant communities.USDA ForestService General Technical

ReportINT-58.

Zedler,P. H.,and G.A. Scheid. 1988. InvasionofCarpobrotus edulis andSalix lasiolepis afterfirein a coastalchaparralsite in SantaBarbaraCounty,California.Madronio35:196-201.

Zeevalking,H.J.,and L.F.M. Fresco. 1977. Rabbitgrazingand species diversityin a dune area.Vegetatio35:193-196.

ConservationBiology Volume6, No. 3, September1992