The Role of Ecology in Biological Conservation Author(s): Peter F. Brussard Source: Ecological Applications, Vol. 1, No. 1 (Feb., 1991), pp. 6-12 Published by: Ecological Society of America Stable URL: http://www.jstor.org/stable/1941843 Accessed: 15/02/2010 11:49 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 http://www.jstor.org/action/showPublisher?publisherCode=esa. 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 [email protected].

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Ecological Applications, 1(1), 1991, pp. 6-12 ? 1991 by the Ecological Society of America

THE ROLE OF ECOLOGY IN BIOLOGICAL CONSERVATION' PETER F. BRUSSARD

Departmentof Biology, Universityof Nevada-Reno,Reno, Nevada89557 USA

Abstract. The emerging science of conservation biology represents an intersection of elements of ecology, genetics, biogeography, and many traditional applied disciplines such as wildlife management and forestry. Its major concern is providing a valid scientific basis for actions that will slow or stop the accelerating loss of biological diversity worldwide. Ecology's major contributions to conservation biology so far include the equilibrium theory of island biogeography and the theoretical relationship between population size and persistence time. In the future ecologists can contribute their skills to conservation biology in numerous ways; I suggest three in particular. These are investigating the autecology and natural history of rare species, testing hypotheses concerning population viability with carefully designed laboratory and field experiments, and working to establish and implement a national policy for the protection of biological diversity on United States public lands. Key words: biologicaldiversity;conservationbiology;extinctions;populationpersistencetimes;rare YellowstoneNational Park. species;Uncompahgrefritillary;

INTRODUCTION

If currentpredictionsaretrue,and thereis littledoubt that most are, the last decade of the twentiethcentury and the firstfew decadesof the twenty-firstwill be years of ever-worseningecological crises. The driving force behind the crises is the human population,now numberingmore than five billion and growingat a rate of :2%/yr. As this population expands, its demand for food, fuel, and other resourcesincreases proportionally. In the less-developedworld human resourcedemandtendsto affectthe environmentdirectly,resulting in easily observedphenomenasuch as widespreaddesertificationand deforestation.The effects of human resourcedemandin more-developedcountriesare less direct but just as destructiveto the environment;examples include acid precipitation,the ozone hole, and global warming. One of the upcoming ecological crises is a major extinctionevent. Within the next centurythousandssome say millions-of species will disappearforever. Prime candidates for extinction include all the large carnivores,many largeherbivores,most primates,and countless other plants and animals that have the misfortune to be rare or highly specialized, or to have restrictedhabitat requirementsand limited distributions. The prospectof human-causedextinctionsofthis magnitudeshouldhorrifyany thinkingpersonfor moral and aestheticreasonsalone, but if it does not, there are excellent practicalreasonsfor alarm.Not only are many of these species direct sources of hundreds of 1 Manuscriptreceived 17 November 1989;revised27 April 1990; accepted 1 May 1990.

useful products such as drugs, building materials, chemicals, and food, but they are also the underpinnings of the naturalecosystems that currentlyprovide free environmentalservices-waste detoxification,pest control, climate amelioration,and flood preventionto name but a few-that will be extremely costly, if not impossible, to replace. The root causes of the pending extinction crisis are economic, social, and political, and until human institutions and values change it is unlikely that major losses of biological diversity can be prevented. However, the timely application of information from the biological sciences can help ameliorate the problem. Conservationbiology is a syntheticdiscipline that focuses on the applicationof biologicalprinciplesto the preservationof biodiversity;it representsa fusion of relevant ideas from ecology, genetics, biogeography, behavior, reproductivebiology, and a number of applied disciplinessuch as wildlife managementand forestry. In fact, virtuallyall of the subdisciplinesof biology have somethingworthwhileto contributeto the conservationchallenge,and conservationbiology actively seeks to incorporaterelevant informationfrom all of them. Conservationbiology emergedas a nomothetic discipline nearly a decade ago with the publication of Soule and Wilcox's (1980) book Conservationbiology: an evolutionary-ecologicalperspective.When I made this claim a few years ago (Brussard1985) I did not mean to imply that there were no biologists working in the field of conservation or that conservationists ignored biology prior to 1980. Rather, the Soule and Wilcox volume breached traditional disciplinary boundaries, brought together a great deal of diverse

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information,and attemptedto apply this information directlyto the preservationof global biodiversity. Subsequently,I believe that the book has had at least three major impacts. First, it stimulated many basic biologiststo applytheirexpertiseto conservationproblems; second, it provided a broaderperspectiveto scientists alreadyworkingin the field. Third, it defineda new interdisciplinaryarea that is now somewhat of a growthindustry.Forexample,over a dozenbooks have appearedwith conservation biology in the title, and severalmore arein preparationor in press.The Society for ConservationBiology, barely four years old, now has >2000 members, and Volume IV of its journal ConservationBiologyis currentlybeingpublished.The EcologicalSociety of America now recognizesconservation biology as a legitimate enterpriseand is providing a publication outlet throughEcological Applications. Although conservation biology is not just applied ecology, ecology has played a centralrole in its development, and it will continue to do so in the future. Major intellectualcontributionsfrom ecology include the theory of equilibrium biogeography(MacArthur and Wilson 1968) and the relationshipbetween populationsize and persistencetime (e.g.,May 1973).These topics have recentlybeen reviewed in detail by Simberloff(1988) and Pimm and Gilpin (1989), and their papers are stronglyrecommendedto those who wish to pursuethese topics in depth. Rather than cover much of this same ground here, I addressthreeareasin whichecologistscan applytheir expertiseto currentproblemsin conservationbiology. These are (1) investigatingthe autecologyand natural history of rarespecies, (2) testing hypothesesconcerning populationpersistencethroughlaboratoryand field experimentation,and (3) workingto establishand implement a nationalpolicy for maintainingbiodiversity on United States public lands. While this is far from an exhaustivelist of whatneeds to be done to minimize extinctions in the next half-century,these three areas will provide dozens of challengingproblems for conservation-orientedecologicalresearch. THEAUTECOLOGY ANDNATURAL HIsToRYOFRARESPECIES

It is usually taken as a sine qua non that sensible managementbegins with a solid, fundamentalunderstandingof a species' ecological relationsand natural history. Unfortunately,we are woefully short on this informationfor most species of conservationconcern. While this may come as no surprisefor rareand littleknown species, it is also true for many of the more charismaticbirds and mammals. For example,tens of millions of dollars are spent annually on grizzly bear research,but thereare no reliabledata on male mating success in this species. Such informationis of critical importancefor calculatinggeneticallyeffective popu-

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lation size, a key element in population viability estimates. In the absence of such data, managers are forced either to extrapolatefrom other species or to ignorethe parameteraltogether.Neitherchoice is likely to result in sound managementplans. As another example, I describe a project recently completed by my researchgroup. The Uncompahgre fritillaryBoloriaacrocnemais probablythe rarestbutterflyin North America.It has beenproposedforlisting as endangered,and the agenciesinvolved (UnitedStates Fish and WildlifeService,United StatesForestService, United StatesBureauof Land Management,Colorado Natural Areas Survey) were anxious to have information on its statusand recommendationsfor its management and recovery. Funds were provided for a 2-yr survey of its distributionand abundance. Althoughthis specieswas describedalmost a decade ago (Gall and Sperling1980), it has not been the object of intensive study by ecologicallyminded lepidopterists. The following information was available in the literatureand agencyreports:The two known colonies are both located above 4000 m in elevation on northeast-facingslopes in the San Juan Mountains of Colorado. Evolutionarily,it representsthe southern terminus of an increasingly differentiated series of populationsof an arcticbutterfly,Boloriaimproba(Gall and Sperling 1980, Ferris 1984, 1986). Its larval food plant is snow willow, Salix nivalis,and the larvaetake two years to develop (Scott 1986). Flights were observed, and the butterflieswere abundantat both colonies for a few yearsaftertheir initial discovery.Capture-mark-recapturestudies were performed on one colony in 1979-1980 (Gall 1984a, b), but no other quantitativedata were obtained on the species' abundance after that time. In 1987, the first year of our study, we found it to be extirpatedat one of the two known localities and very rareat the other; in 1988 it was presentat both, althoughat greatlyreducednumbers from those previously reported.Althoughwe surveyedthe surrounding areaextensively,no othercolonies of any size were found even though there had been anecdotal reports of their existence in previous years. In the absenceof detailed autecologicalinformation and long-termpopulationdatawe wereforcedto make several assumptions to interpretthis precipitous decline in numbersand to recommendappropriatemanagementactions. These assumptionswere (1) that the colony locations-both on northeast-facing slopes above 4000 m-reflect the butterfly'sarctic heritage and a requirementfor the coolest, most mesic habitat availablein the San Juans;(2) that B. acrocnema,like other fritillaries,has no highly specializedhabitat requirementsotherthan an appropriatemicroclimatefor larval development, sufficientamounts of the larval food plant, and the presence of adult nectar sources; (3) that other colonies existed-at least in the recent past-and that the species was distributedhistorically

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EcologicalApplications I

as a metapopulationwith a dynamic equilibriumbetween the rate of colonization of new patches and the extinctionof old ones;and (4) that the species'biennial life historyhas led to both odd- and even-yearbroods, which functionessentiallyas separatepopulationswith independentdemographictrends. Since there had been several extremelyhot and dry summersin southwesternColoradoduringthe past few years, and since both colonies apparentlydeclined simultaneouslyduringthis period, we proposedthat climatic stresswas responsiblefor the observeddeclines in the known colonies and the apparentextirpationof the others that were rumoredto exist. Duringour survey we identified>50 sites at the appropriateelevation and exposure that supportedthriving populations of snow willow and had suitableadult nectarsources.We quantified29 variablesrelatedto these habitatparametersand searchedforboth univariateand multivariate correlationsthat might separatethe two occupiedsites from the rest. None was found, consistent with our assumption of ample habitat availability. However, there will always be the nagging doubt that we had overlooked the importance of some critical habitat component,and that unoccupiedsites were vacant for reasons other than a long run of bad weather. We interpretedthe lack of a flightat one known site in 1987 as the extirpationof the odd-yearbroodtherein other words, the loss of one quarterof the species' entire population. However, there is a possiility that B. acrocnemamay actuallyhave only one brood, with individual larvae showing a large variance in developmental rates, some maturingin 2 yr, others in ?3 yr, dependingon ecologicalconditions. Sucha life history patternis known in other butterflies,and if true for the Uncompahgrefritillary,it would make the absence of the 1987 flight at one colony much less ominous for the species' continued existence. Clearly, adequate ecological data on this species would have provided us with a much more firm foundation on which to base interpretationof the results of our survey and to prescribeappropriatemanagement strategies.It is importantto note thatsuchstudies need not have been mindless exercises in data gathering;rather,the four assumptionslisted above could have been used as testablehypothesesto drive several researchprojects. During the early 1980s the species was abundantenough to produce respectablesample sizes, and the two colonies are located within 160 km of the Rocky Mountain Biological Laboratory,a field station well known for its research on lepidopteran populations. However, for whatever reasons-reluctanceto workon rarespecies,lackof interestor funding opportunities,logistical problems, etc.-the excellent early work on B. acrocnemawas not followed by detailed ecological studies. As a result we had to make our recommendationson the basis of seemingly reasonable,but largelyuntested,assumptions.If these assumptionsarevalid, the returnof morenormalweather

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conditions should eventuallyresult in the recoveryto early 1980's levels of both broods at one colony and of the even-yearbrood at the other. After this level of recoveryhas occurred,artificaltranslocationto other apparently suitable, but currently unoccupied, sites could providea greatermarginof safetyfor the species in the future. If our assumptionsare not valid, however, B. acrocnema may be the first full species of butterflyto go extinct in North America. EMPIRICAL VERIFICATION OFPOPULATION CONCEPTS VIABILITY One of the major tenets of conservationbiology is that stochastic, rather than deterministic, processes govern the dynamics of small populations.Four types of stochasticityarerecognized,two of which have been modeledfairlyextensively(see Soule [1987]fora recent review and appropriatereferences).Accordingto these models, demographicstochasticity-the randomvariationin individualbirthanddeathrates-has littleeffect on the persistenceof any but the smallest populations (i.e., those with <20 breedingindividuals).On the other hand, environmental stochasticity-environmentally driven variability in birth and death rates that affect the population as a whole-has a much more importanteffecton persistenceprobabilities,and much larger numbers are required for a population to be immune from its effects.Catastrophe,the thirdtype of random occurrencethat can affect persistencetimes, may be consideredas the endpoint of a continuum of environmentalstochasticity.If the catastropheis sufficientlysevereand widespread,no populationsize will be largeenough to guaranteethe avoidance of extinction-witness the events at the Cretaceous/Tertiary boundary. The fourth type of stochasticity involves random genetic processesthat occur in small populations:specifically, increased inbreedingand loss of variability by drift. The influenceof genetic stochasticityon population persistence has proved to be particularlyrefractoryto predictionfor several reasons. First, its effects dependnot on the census size of a populationbut on its geneticallyeffective size, a number notoriously difficult to estimate. Second, past population history enters into the picture,and this is hardlyever known. Third, genetic events often interactwith demographic ones in complex ways, and it is very difficultto sort out cause from effect. Although modeling the four apocalyptichorsemen of persistencetimes has sharpenedour perceptionsof the kinds and potentialseverityof events that can happen to small populations,none of these ideas has been adequately tested in the field. However, there is no reasonwhy appropriateexperimentalprogramsutilizing laboratory, natural, and semi-natural situations cannot be establishedto test the predictionsof these models. For example,Goodman (1987a, b, c) has concluded that under readily recognizablecircumstances

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populations are liable to local extinction in relatively shorttimes, and that the variancein populationgrowth rates is a robust predictor of persistence times. Furthermore,Goodman suggestsprotocols for testing his model by accumulatingdata on the spectrum of accessible sizes in a number of carefullychosen populations. I takeGoodman'srecommendationsone step further and point out that the requirementsfor an appropriate test organismwould be a species that is rathersedentary and inhabits well-definedpatches of habitat. Individualsmust be easy to census so that samplingerror is a minor component of population size estimation. The species must have a relatively high intrinsic rate of increase (r), and its reproductivesuccess must be dependent to some extent on easily monitored environmentalvariation(e.g.,the extentand timing of rainfall,temperatureat criticaltimes,etc.).Generationtimes must be shortenoughso that usefuldata, includingthe observation of several extinctions, can be obtained within reasonabletime periods. If the populationsare located within normal dispersaldistances for the species it will also be possible to use the experimental system to test several aspects of metapopulationtheory and to relate metapopulation dynamics to the structureof the environmentand to the size and isolation of habitat patches (e.g., Hanski 1989). Habitat patches arrayed along an environmental gradient of known effect on the species would be ideal for this. If tissue samples can be obtained nondestructively,or if individualscan be sampledafterreproductionwith no effecton subsequentdemographictrends,then gel electrophoresiscan furnishinformationon changesoccurring in genetic parameterswithin the populations as well. Manyinsects or otherinvertebrates,many plants, and some small vertebrateshave naturalhistorycharacteristics that would make them amenable to such studies. Severalexamples have appearedin the recent literature.Forney and Gilpin (1989) used laboratory populations of Drosophilato test the relationshipbetween population size and persistence times, and to test the efficacyof corridorsin preventingextinctions. A semi-naturalfield experimenton populationpersistence using marine snails is describedby Quinn et al. (1989), and McCauley's(1989) paperon milkweedbeetles should convince the skepticalthat naturalextinctions ofpopulationsofthis speciesarecommon enough to merit detailed study. It must be noted that studies like these increase in value considerablyif they are carriedout for long periods of time. Although some meaningful investigations of population persistence may be reasonable graduatestudentprojects,the most significantpayoffs will come from long-termcommitments. Perhapsthe ideal situationwould be that every ecologist,upon taking a permanentposition somewhere, choose an appropriatesystem and work on it faithfully-even at a low level-for at least 15 yr.

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PROTECTIONAND MANAGEMENTOF BIOLoGIcAL DIvERsITY

Ecologists study populations of vertebrates,invertebrates, plants, and microbes, and they seek to understandthe patternsand linkagesamong these populations when they assemble into communities. Thus biological diversity truly is the ecologists' bread and butter.Yet anyone who has done field work for more than a decade knows that biodiversityis disappearing, not only in the tropics, but in North America as well. Favorite field sites have vanished under condominiums or parking lots; species once common are now rareor extirpated;unusualcommunitytypes no longer exist locally. To be blunt, preservingbiodiversityis clearlyin every ecologist's own self-interest,and he or she should pledge to spend a measurableportion of time working to accomplishthis goal. The firststep is a commitment to the politicalactionnecessaryto enactan appropriate nationalpolicy on biodiversity.Such a policy requires threemajorelements.First,it mustexplicitlyrecognize that biodiversityincludesall speciesof plants,animals, and microbiallife, the genetic variation within them, and the full variety of communities that result from their aggregations.Second, it must contain provisions for the inventory and continued monitoring of these elements. Third, it must mandatethat the diversity of species, genetic variation, and community types now found in public lands will be maintained over an acceptable period of time with an acceptabledegree of risk. Environmentalgroups often need ecological expertiseto articulatethese requirementsadequately,and legislatorsneed biologicallysophisticatedadvice when drafting appropriatelegislation. Ecologists can contribute much in either arena. HR 1268, the Scheuer biodiversity bill currently pending in Congress (see Blockstein[1989] for details),is not perfect,but it has many good features.Every ecologist should lobby for its passage. I specifypublic lands in particularbecausethese are the majorreservoirsof biologicaldiversityin the United States.At present,however, the Forest Service and the Bureauof LandManagementlargelymanagelands undertheir control for the productionof certaincommodities (e.g., timber, cattle, or minerals)without too much regardfor the effectsthese activities might have on othercomponentsofthe system,althoughmanaging for "wildlife"(i.e., species subjectto sport harvest)or "wildlife habitat"may be one of several multiple-use managementgoals. Lands managed by the Fish and WildlifeServiceand the National ParkServicearegenerally oriented toward the protection of charismatic, recreationallyimportant,or officiallylisted endangered or threatenedspecies,while otherelementsofbiodiversity are usually ignored. It is often assumed that the biological diversity of an area,if given some degreeof protectionfrom human

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activities and left to its own devices, will be self-perpetuating.This is probablytrue(at least on a time scale of decades) if the area is large enough to support a naturaldisturbanceregime of sufficientmagnitudeto ensurecontinuedsuccessionand viable populationsof the species with the largestspace requirements.However, these conditions are rarely met on any of our publiclands. Eitherpreserveareasarenot largeenough to supportviable populationsof the largestspecies, or naturaldisturbancesare not allowed to proceed without major human intervention,or both. Yellowstone National Park is an excellent case in point. About 20 yr ago Yellowstone chose to follow a policy of "naturalregulation."The philosophybehind this decision was that if left alone, or at least managed minimally, Yellowstone would eventually revert to somethingresemblingits pre-settlementstate. Among otherthings,naturalregulationmeantthat the elk population would no longerbe controlledby parkpersonnel; rather,their numberswould be governedby their food supply. Bears would no longer be fed at garbage dumps;instead,they would foragecompletelyon their own. Fires, if ignited by lightning,would be allowed to burnas long as they did not threatenlife or property. The successof these policies has been mixed at best. Aftercontrol of elk stopped,the animals soon reached very high numbers. Although the effects of this population explosion are still a matter of debate, many biologists feel that they caused a significantreduction of aspen communities and riparianvegetation, with attendant negative impacts on other animal species. The effectsof the naturalregulationpolicy on the grizzly seem to be moreclear-cut.Afterthe garbagesubsidy stopped, the bears expanded their foragingareas and increasinglycame into serious conflicts with human activities both on and off park land-resulting in a substantialpopulationdecline.The naturalburnpolicy had little effect in reducingthe extent and intensity of the major fires of 1988, which, at least in retrospect, could have been predicted from the ecology and fire history of the region. During years with normal precipitationfew naturalignitionsoccur,and the firesthat do startdo not usuallyburnverywell. However,during severe droughtyears such as 1988, fires characteristically sweep throughthe Yellowstonearea,clearingout thousandsof acres of aged, insect-damagedlodgepole forests and setting the stage for massive renewaland regeneration(Christensenet al. 1989). The major reason that naturalregulationdoes not work very well in Yellowstone is because of a scaling incompatibility;the park is too small to contain selfregulatingpopulationsof largemammalsor to support a naturaldisturbanceregime of the size characteristic of the northernRocky Mountainregion.When events suchas the 1988 firesor a severewinterkillof ungulates occur, they are perceivedas disastersbecausethey are concentratedin a relativelysmall, highly visible, area. And if Yellowstone, the largest national park in the

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lower 48 states,with an areaof 900 000 ha, is too small for naturalregulationto succeed, such a management philosophy is surelydoomed to fail in the smallernational parksas well. If our major naturereservesare too small to maintain biodiversityover the long term,whatcan be done? One possibility might be to increasethe preservearea. Managingthe entire 5 200 000 ha of public land in the "GreaterYellowstoneEcosystem"as a wildernesspreserve has receiveda greatdeal of discussion(e.g.,Clark and Harvey 1989). This land area probably is large enoughto supportviable populationsof all native species along with the local disturbanceregimeon an appropriate scale. Unfortunately, much of this land is enormouslyvaluable for other uses, and political and economicrealitiesindicatethatcommodityproduction and multiple-usephilosophies will prevail over preservationistgoals for a long time to come. The second, and more realistic, possibility, is to managepublic lands actively to protecttheir biodiversity. Managementactionsto accomplishthis goalwould rangefrom largelylaissez-faireto highly manipulative depending upon the circumstancesand the degree of risk involved, and the policy would applyto both multiple-use (e.g., Forest Service, Bureau of Land Management)and ParkServicelands.The majoradvantage of this approachon multiple-use lands is that under an active biodiversity management policy relatively little additionalacreagewould have to be "lockedup" in preserves, and traditional activities (e.g., mining, logging,grazing,recreation)would not usuallyhave to be curtailed. Rather, the effects that these activities mighthave on biodiversitywould need to be evaluated in the same manner as their effects on watershedsor "wildlife" are evaluated now. Any negative impacts these activitiesmight have on biodiversitywould have to be mitigated; only if no mitigation were possible would the activities not be allowed. The majoradvantageof active managementfor biodiversity in the national parks is simply that it will work better than the currentpolicy of naturalregulation. Species are being lost from Park Service lands because of their small size and insularity(e.g., Newmark 1987); communities are disappearingbecause natural disturbanceregimes are interrupted.If these trendsare to be reversed,managementactions such as speciesreintroductions,habitatrestoration,population enhancement,control of overabundantcommon species, and restrictionof human visitation will have to be implemented on Park Service lands if and when circumstancesdictate."Ecosystemmanagement,"a new conceptual approachfor managingparks and wilderness areasthat has supportfrom some agencyscientists (Ageeand Johnson1988),may be ableto accommodate most of the manipulationslisted above. If so, ecosystem managementmay provide the Park Service with a viable substitute for natural regulation.While this degreeof interferencemay sound to some like treating

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nationalparksas backcountryzoos, it is far preferable to just wishfully thinking that natural regulationwill result in their retaininga full complement of species and communities a centuryfrom now. In orderfor active managementto work successfully on either multiple-use or wilderness lands, well-formulated strategicand tactical plans must be designed and implemented. Unfortunately,the currentphilosophies of the land managementagenciesand the skills of most of their personnelare not up to this task. For example,the idea that biodiversityincludesmore than just "wildlife"sensu strictois virtuallyforeignto agency thinking. Likewise, comprehensive approachesto management do not characterizethe strategies currentlyemployedby land-managementagencies;rather, compartmentalizationand specializationhave become theirprimaryfoci. Finally,the solid, underlyingscience necessaryfor developingappropriateconceptualmodels for biodiversity managementis generallylacking; even worse, what is known is rarelyapplied. For these reasons the active participationof ecologists in the development and implementationof biodiversitymanagementis extremelyimportant.Appropriate managementactions will often require a wide variety of interventions at the population to wholeecosystem levels of organization.This means that unanticipatedand unwantedside effectswill most likely accompanymany of these actions unless a substantial amountof modelingandexperimentationhas preceded them. The models will have to be based on current theory in ecology, biogeography,genetics, and evolution, and the experimentsmust be well thought out, rigorous,adequatelyreplicatedif possible, and genuinely relevantto the contemplatedactions. Even then, unwelcome surprises will doubtless occur, and the managementmust be flexibleenoughto accommodate changesand reversalswhen they are called for. This interplay among modeling, experimentation, and managementaction will requirenot only increased communicationbetween resourcemanagersand ecologists in generalbut also active involvement of nonagencyscientists in the entire process-from planning through implementation to evaluation. Such an involvement is a two-way street. First, it will require overcoming the prevailing apathy of most ecologists towardconservationissues. Second,it will requirethat agency biologists and managersseek out non-agency scientistsand encouragethem to utilize their expertise in solving these criticallyimportantproblems.

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with my views, I appreciatetheirinput.Finally,I thankSimon Levin for inviting me to participatein this symposiumand to reviewersfor useful comments. LITERATURECITED

Agee, J. K., and D. R. Johnson, editors. 1988. Ecosystem managementfor parksand wildernessareas.University of WashingtonPress, Seattle,Washington,USA. Blockstein,D. E. 1989. Biodiversitybill update.BioScience 37:677. Brussard,P. F. 1985. The currentstatus of conservation biology. Bulletinof the EcologicalSociety of America66: 9-11. Christensen,N. L., J. K. Agee, P. F. Brussard,J. Hughes,D. H. Knight, G. W. Minshall, J. M. Peek, S. J. Pyne, F. J. Swanson,J. WardThomas, S. Wells, S. E. Williams, and H. A. Wright. 1989. Interpretingthe Yellowstonefiresof 1988. BioScience39:678-685. Clark,T. W., and A. H. Harvey. 1989. Managementof the GreaterYellowstone Ecosystem:an annotated bibliography. Northern Rockies Conservation Cooperative, Box 2705, Jackson,Wyoming,USA. Ferris,C. D. 1984. Overviewof Clossianaimproba(Butler) in North Americawith a descriptionof a new subspecies fromWyoming(Nymphalidae:Argynninae).Bulletinof the Allyn Museum number89. . 1986. Fieldnotes of ClossianaimprobaharryiFerris (Lepidoptera:Nymphalidae).Journalof Researchon the Lepidoptera25:71-72. Forney, K. A., and M. E. Gilpin. 1989. Spatial structure and population extinction: a study with Drosophilaflies. ConservationBiology 3:45-51. Gall, L. F. 1984a. Populationstructureand recommendations for conservationof the narrowlyendemicalpine butterflyBoloria acrocnema(Lepidoptera:Nymphalidae).BiologicalConservation28:111-138. . 1984b. The effects of capturingand marking on subsequent activity in Boloria acrocnema (Lepidoptera: Nymphalidae)with a comparison of differentnumerical models that estimate population size. Biological Conservation 28:139-154. Gall, L. F., and F. A. H. Sperling. 1980. A new high-altitude species of Boloriafrom southwesternColorado(Nymphalidae) with a discussion of pheneticsand hierarchicaldecisions. Journalof the Lepidopterists'Society 34:230-252. Goodman,D. 1987a. Thedemographyofchanceextinction. Pages 11-34 in M. E. Soule, editor. Viable populationsfor conservation.CambridgeUniversityPress,Cambridge,England. . 1987b. How do any species persist?Lessons for conservationbiology. ConservationBiology 1:59-62. . 1987c. Considerationof stochasticdemographyin the designand managementof biologicalreserves.Natural ResourceModeling1:205-234. Hanski, I. 1989. Metapopulationdynamics:does it help to have more of the same?Trendsin Ecologyand Evolution 4:113-114. MacArthur,R. H., and E. 0. Wilson. 1968. The theory of island biogeography.Princeton University Press, Princeton, New Jersey,USA. May, R. M. 1973. Stabilityand complexityin model ecosystems. PrincetonUniversity Press, Princeton,New JerACKNOWLEDGMENTS sey, USA. Workon the Uncompahgrefritillarywas supportedby the McCauley,D. E. 1989. Extinction,colonization, and popU.S. Forest Service, the Bureauof Land Management,and ulation structure:a study of a milkweedbeetle. American the U.S. Fish and WildlifeService.David Kuntz of the ColNaturalist134:365-376. orado NaturalAreas Programwas instrumentalin securing Newmark,W. D. 1987. A land-bridgeperspectiveon mamthis fundingand in encouragingme to become involved with malian extinctionsin westernNorth Americanparks.Nathe project.I have discussedmy ideas on biodiversitymanture 325:430-432. agementwith many colleaguesboth in academiaand in the Pimm, S. L., and M. E. Gilpin. 1989. Theoreticalissues in conservationbiology. Pages 287-305 in J. Roughgarden, land-managementagencies.While hardlyany of them agree

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R. M. May, and S. A. Levin, editors. Perspectives in ecological theory. Princeton University Press, Princeton, New Jersey, USA. Quinn, J. F., C. L. Wolin, and M. Judge. 1989. An experimental analysis of patch size, habitat subdivision, and extinction in a marine intertidal snail. Conservation Biology 3:242-251. Scott, J. A. 1986. The butterflies of North America. Stanford University Press, Stanford, Califomia, USA.

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Simberloff,D. 1988. The contribution of population and community biology to conservation science. Annual Review of Ecologyand Systematics19:473-51 1. Soule, M. E., editor. 1987. Viable populationsfor conservation. CambridgeUniversity Press, Cambridge,England. Soule, M. E., and B. A. Wilcox,editors. 1980. Conservation biology. An evolutionary-ecologicalperspective. Sinauer, Sunderland,Massachusetts,USA.