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    November 1999 1189HISTORICAL VARIABILITY

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    Ecological Applications, 9(4), 1999, pp. 118912061999 by the Ecological Society of America

    APPLIED HISTORICAL ECOLOGY:USING THE PAST TO MANAGE FOR THE FUTURE

    T HOMAS W. S WETNAM ,1,4 C RAIG D. A LLEN ,2 AND JULIO L. B ETANCOURT 3

    1 Laboratory of Tree-Ring Research, The University of Arizona, Tucson, Arizona 85721 USA2U.S. Geological Survey, Jemez Mountains Field Station, Los Alamos, New Mexico 87544 USA

    3U.S. Geological Survey, Desert Laboratory, 1675 West Anklam Road, Tucson, Arizona 85745 USA

    Abstract. Applied historical ecology is the use of historical knowledge in the man-agement of ecosystems. Historical perspectives increase our understanding of the dynamicnature of landscapes and provide a frame of reference for assessing modern patterns andprocesses. Historical records, however, are often too brief or fragmentary to be useful, orthey are not obtainable for the process or structure of interest. Even where long historicaltime series can be assembled, selection of appropriate reference conditions may be com-

    plicated by the past inuence of humans and the many potential reference conditionsencompassed by nonequilibrium dynamics. These complications, however, do not lessenthe value of history; rather they underscore the need for multiple, comparative historiesfrom many locations for evaluating both cultural and natural causes of variability, as wellas for characterizing the overall dynamical properties of ecosystems. Historical knowledgemay not simplify the task of setting management goals and making decisions, but 20thcentury trends, such as increasingly severe wildres, suggest that disregarding history canbe perilous.

    We describe examples from our research in the southwestern United States to illustratesome of the values and limitations of applied historical ecology. Paleoecological data frompackrat middens and other natural archives have been useful for dening baseline conditionsof vegetation communities, determining histories and rates of species range expansions andcontractions, and discriminating between natural and cultural causes of environmentalchange. We describe a montane grassland restoration project in northern New Mexico thatwas justied and guided by an historical sequence of aerial photographs showing progressivetree invasion during the 20th century. Likewise, re scar chronologies have been widelyused to justify and guide fuel reduction and natural re reintroduction in forests. A south-western network of re histories illustrates the power of aggregating historical time seriesacross spatial scales. Regional re patterns evident in these aggregations point to the keyrole of interannual lags in responses of fuels and re regimes to the El NinoSouthernOscillation (wet/dry cycles), with important implications for long-range re hazard fore-casting. These examples of applied historical ecology emphasize that detection and expla-nation of historical trends and variability are essential to informed management.

    Key words: climate change; disturbance; re history; historical ecology; packrat middens; pa-leoecology; range of natural variation; repeat photography; restoration; southwestern United States;tree rings; vegetation change.

    INTRODUCTION

    A science of land health needs, rst of all, a base

    datum of normality, a picture of how healthy land maintains itself as a organism.Aldo Leopold (1941)

    Scientists and managers alike are increasingly usingenvironmental history as a base datum for under-standing and managing ecosystems. This expanding in-

    Manuscript received 18 May 1998; revised 11 January1999; accepted 10 February 1999; nal version received 4March 1999. For reprints of this Invited Feature, see footnote1, p. 1177.

    4 E-mail: [email protected]

    terest in applying history is evident in a recent prolif-eration of land and water management agency reportsthat outline planning stages dependent upon the gath-ering and interpretation of historical ecology infor-mation. These documents recommend the use of historyto determine the range and variability of ecologicalprocesses and structures during times when ecosystemswere less affected by humans (e.g., Brenner et al. 1993,Kaufmann et al. 1994, Manley et al. 1995, Dahm andGeils 1997). Reference conditions may be used, alongwith current condition assessments, social and eco-nomic considerations, and other practical constraints,for the setting of achievable and sustainable manage-ment goals. Several recent regional assessments have

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    also extensively utilized historical information as basedata for identifying spatiotemporal trends and vari-ability (e.g., Quigley et al. 1996, Sierra Nevada Eco-system Project 1996, Kaufmann et al. 1998). Moreover,a central tenet of the expanding eld of restorationecology is that reference conditions are useful for de-ciding upon ecologically justiable goals for restora-tion programs (Allen et al. 1996, Covington et al. 1997,Fule et al. 1997, White and Walker 1997, Moore et al.1999). Holling and Meffe (1996) dene a GoldenRule of natural resource management that asserts:management should strive to retain critical types andranges of natural variation in resource systems in orderto maintain their resiliency.

    There remains, however, much debate about how,where, and to what extent natural variations or refer-ence conditions derived from historical ecology shouldbe applied to land management (Landres et al. 1999,

    Millar and Wolfenden 1999). Although applied histor-ical ecology is an evolving eld, there appears to be abuilding consensus that, at a minimum, it is very usefulto know and understand the past to properly manageecosystems for the future.

    Recent reviews of the use of reference conditionsand the concepts of historical (or natural) range of variability have carefully dened terminology andsome of the theoretical and practical constraints in ap-plying history to restoration and management (Allen1994, Kaufmann et al. 1994, Morgan et al. 1994, Swan-son et al. 1994, Foster et al. 1996, Jackson et al. 1997,Millar 1997, White and Walker 1997, Landres et al.1999). As might be expected in the development of a

    relatively new concept, the early stages have beenmarked by some confusion and missed opportunitiesfor better communication between scientists and man-agers. Initial confusion about seemingly vague termi-nology, and perhaps a lack of familiarity with resourcemanagers information needs, may have led some sci-entists to dismiss the concept of range of natural vari-ability as misguided. Despite the lack of involvementby a broad spectrum of ecologists, (i.e., before the late1990s), some land management agencies forged aheadwith implementation of some of the basic ideas (e.g.,Manley et al. 1995).

    Many ecologists have recognized the importance of historical knowledge for understanding and managing

    ecosystems (Christensen et al. 1996), but with notableexceptions (e.g., Foster et al. 1996, Hunter et al. 1988,Tausch 1996, Delcourt and Delcourt 1998), relativelyfew paleoecologists have addressed the topic directly.In contrast, paleolimnologists and paleohydrologistshave been more directly involved in using historicalperspectives in aquatic restoration and managementprograms (Smol 1992, Brenner et al. 1993, Schmidt etal. 1998; and see Carltons [1998] plea for more his-torical oceanographic research). Historical trends inwater quality, for example, are determined by chemical

    analyses of dated sediment cores from lakes, reservoirs,and estuaries (e.g., Van Metre et al. 1997). Undesirabletrends (e.g., increasing acidity from pollution) havebeen used to justify restoration treatments (Brenner etal. 1993). Similarly, there are many examples of theapplications of paleohydrology for assessment of oodhazards (e.g, Kochel and Baker 1982, Ely et al. 1991),and historical perspectives have been useful in evalu-ating restoration and management alternatives in reg-ulated river systems (e.g., the Colorado River; Schmidtet al. 1998).

    The objective of this paper is to provide an overviewof some key applications of historical ecology for in-forming land management. We briey review the pri-mary historical ecology data types and methods, as wellas their temporal and spatial scope, resolutions, andgeneral limitations. We use examples of our own re-search in the southwestern United States to illustrate

    specic insights and limitations of applied historicalecology. We conclude by discussing some general is-sues raised by our examples. Overall, our emphasis ison the importance of knowing the history of land-scapes.

    H ISTORICAL E COLOGY : SCOPE , RESOLUTION , ANDL IMITATIONS

    Data and techniques

    Historical ecology encompasses all of the data, tech-niques, and perspectives derived from paleoecology,land use history from archival and documentary re-search, and long-term ecological research from moni-toring and experiments extending over decades. Also

    included are time series from instrument-based obser-vations of the environment, such as weather records,stream gauges, and data from satellites. The essentialfeature of historical ecology is a sufciently long timesequence (chronology) of measurements or observa-tions so that meaningful information can be gainedabout changes in populations, ecosystem structures,disturbance frequencies, process rates, trends, perio-dicities, and other dynamical behaviors. To understandecosystem patterns, we must obtain and evaluate datathat match the spatial and temporal scales of the pro-cesses responsible for those patterns (Ricklefs 1987,Levin 1992). Relatively short time frames may be ad-equate for evaluating dynamics of some ecosystemcomponents, but by long time sequences we meanthe longer temporal scales that are usually most rele-vant to understanding and managing landscapes withlong-lived organisms, such as trees. In this context,history typically encompasses time frames from de-cades to millennia.

    The sources, time frames, and temporal/spatial res-olutions of archival information are highly diverse (Fig.1). These archival records are classiable as naturaland documentary. The natural archives are thoserecorded by earth-system processes, such as sedi-

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    F IG . 1. The range of temporal and spatial scales typically encompassed by historical-ecology data (modied from Fosteret al. [1990]).

    mentation (e.g., pollen, charcoal, plant macrofossils,phytoliths, etc., found in lakes, bogs, and soils; Davis1989, Jackson 1984, Jackson et al. 1997), animal de-posits (packrat middens; Betancourt et al. 1990), annualplant and animal growth cycles (e.g., tree rings andcoral layers; Fritts and Swetnam 1989, Dunbar andCole 1993), and other layered records, (e.g., ice cores;(Thompson et al. 1998). Documentary archives are

    written, tabulated, mapped, or photographic records(e.g., chronicles, diaries, cultural histories, land sur-veys, maps, plot measurements, weather observations,etc.; Foster et al. 1990, 1996, Galatowitsch 1990, Rus-sell 1997). (A useful and expanding worldwide databank for many of these types of records is located atthe National Geophysical Data Center, National Oce-anic and Atmospheric Administration, Boulder, Colo-rado, USA. Data, bibliographies, and search tools areaccessible on the World Wide Web.)

    Reconstructions of environmental history are im-proved by complementary and comparative analyses of both natural and documentary records. For example,interpretation of tree-ring reconstructions of climatehistory are most specic and reliable when treegrowthclimate models are statistically calibrated withmeteorological observations and then veried with in-dependent observations (Fritts 1976). Another exampleof combining multiple lines of evidence is the use of both repeat photography and tree demographic data(e.g., based on tree rings) to describe and explain veg-etation change in the late 1920th centuries as a con-sequence of human actions (e.g., Veblen and Lorenz1986, 1991). Blending different methods and data typescan extend information about environmental change

    across a broad range of temporal and spatial scales.Examples include comparative analyses of climate, dis-turbance, vegetation, and land use histories derivedfrom charcoal and pollen in lakes and bog sediments,plant remains in packrat middens, tree rings, and doc-umentary records (Foster et al. 1990, Anderson andCarpenter 1991, Bahre 1991, Betancourt et al. 1991,Fastie 1995, Millspaugh and Whitlock 1995, Baisan

    and Swetnam 1997, Lloyd and Graumlich 1997, Rus-sell 1997, Allen et al. 1998, Allen and Breshears 1998,Swetnam and Betancourt 1998).

    Limitations

    Historical ecology data and methods have a numberof inherent limitations. Natural archives, often referredto as proxy records, are subject to the ltering of past environmental information through physical andbiological processes. The delity of the reconstructedenvironmental variable, therefore, depends on how wellthis ltering process is understood and modeled. Somespecialized elds, such as dendrochronology and pal-ynology, are based upon a solid foundation of extensiveeld and laboratory observations and experimental re-search on mechanisms of the proxy record formation.These elds have also developed rigorous standards of statistical model development and testing that are par-simonious with this physical and biological under-standing (Fritts 1976, Crowley and North 1991). Evenwith careful model calibration and validation, however,interpretations of proxy records of past environmentsare often subject to multiple assumptions. Althoughpaleoecologists and paleoclimatologists seek to mini-mize the effects of confounding environmental factors

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    on the recording unit (i.e., trees, corals, etc.) throughsampling and analytical techniques, it is not possibleto completely control multiple environmental factorsof the past (White and Walker 1997).

    Historical data and interpretations are further chal-lenged by the possibility of changing relations betweenthe physical and biological recording processes and theenvironment that are not captured in modern calibrationperiods. This so-called no analogue problem arisesfrom the lack of modern analogues for past conditions,or vice versa, resulting in violation of some assump-tions of uniformitarianism. The resolution or avail-ability of historical data also tends to be limited by thelack of preservation, degradation, or loss of evidencethrough time. The transient, stochastic, and incompletenature of the paleorecord development and preserva-tion results in missing, patchy, or altered records. Thechief consequences are that (1) most historical time

    series suffer from a fading record problem, wherethe reliability of the time series decreases with increas-ing time before the present, and/or (2) no record isavailable at all for some ecosystem processes or struc-tures that may be of primary management concern.

    Documentary records are also limited in quantity andquality by a kind of cultural ltering that affectstheir availability, completeness, and reliability. Mostareas of the western United States, for example, lack any written, continuous records of weather observa-tions before the late 19th or early 20th century. Evenwhere old weather records, photographs, or other doc-umentary records can be found, they are subject tovarious biases associated with the locations and periodsfor which they are available. For example, weather-recording stations were often located primarily in lowelevations near or within settlements, and they weresometimes moved from one location to another. Over-all, these and other limitations of documentary recordsdo not invalidate their use in environmental history,but they do call for judicious and careful interpreta-tions, with cognizance of their potential biases and dis-tortions.

    Because historical records may not be available fora particular place or time of interest, extrapolation of reference conditions may be the only resort. White andWalker (1997) point out that all ecosystems and their

    histories are unique at some scale of analysis. There-fore, the validity of extrapolating reference conditionsprobably decreases as a function of both increasingspatial and temporal distance from the current ecosys-tem to which those reference conditions might be ap-plied. White and Walker refer to this as the distancedecay problem. Representative sampling is an im-portant concern of landscape ecology (Stohlgren et al.1995), and the added dimension of time further in-creases the need to carefully consider the scales andlocation of available data in judging the validity of

    extrapolations. This problem highlights the need formultiple places and times for reference.

    Historical science is largely inductive, and the in-terpretation of gathered historical facts can be highlysubjective. This challenges ecologists and managers toquantify the historical evidence as thoroughly as pos-sible, and to carefully interpret historical facts withknowledge from other sources (e.g., present-day ob-servations and experiments). A primary strategy forincreasing objectivity and condence in historical in-terpretations is a combination of comparative analysesand testing with multiple, independent data sources andmethods, and the use of converging lines of evidence(White and Walker 1997, Foster and Motzkin 1998).This strategy includes combining historical reconstruc-tions with eld experiments (e.g., Fastie 1995, Lloyd1996, Lloyd and Graumlich 1997), and comparingmodel simulations with historical data (e.g., Anderson

    1995, Miller and Urban 1999). Despite the constraintsand limitations of history, the power and utility of his-torical insight should not be underestimated. Indeed, itis arguable that historical approaches have been as es-sential and effective in advancing basic understandingin ecology as they have been in other natural and phys-ical sciences, such as geology and astronomy (Brown1995).

    E XAMPLES FROM THE S OUTHWESTERNU NITED S TATES

    We focus here on a few examples from our researchin the southwestern USA to illustrate some basic in-sights and uses of historical ecology. These examples

    draw mainly upon the natural archives of packrat mid-dens and tree rings, as well as the documentary archivesof land use history and repeat photography. Our ex-amples begin with millennia-length perspectives of plant migrations and invasions. These perspectives em-phasize the importance of historical knowledge that isspecic to particular landscapes and the danger in ex-trapolating causation of ecosystem changes from onearea or time period to another. We follow with exampleswhere more recent ecological history has been directlyused in justifying and guiding land management de-cisions.

    Long-term perspectives of 20th century invasions

    and changing forest structureManagers of ecosystems sometimes justify their ac-

    tions based on historical assumptions, if not actual ev-idence. In the Southwest, there is no better examplethan blanket acceptance that grazing and re suppres-sion triggered wholesale encroachment of trees andshrubs into rangelands (Archer 1994) and higher stemdensities across woodlands and forests (Covington andMoore 1994). Beginning in the 1950s, assumptionsabout so-called invasions rationalized an aggressivecampaign of pinyon and juniper eradication (through

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    chaining, cabling, burning, and herbicides), presum-ably to increase forage and water yield (Gottfried etal. 1995, Lanner and Van Devender 1998). Having lostpopularity in the Southwest, a similar campaign maynow be contemplated in the Great Basin, where pinyonand juniper are apparently on the move (Miller andWigand 1994).

    The intent in this section is not to question that graz-ing can shift the balance from grasses to trees andshrubs, or that decades of re suppression beget doghair thickets in many forest ecosystems. Rather, weuse the fossil record to point out that modern invasionsand changes in forest and woodland structure can bedriven by something other than human action. Thatinvasions are occurring in the midst of human distur-bance does not invalidate their roots in Holocene mi-gration, demographic responses to climatic variability,or even recovery from past disturbances, including pre-

    historic humans. We provide some important examplesfrom historical ecology in the Southwest.Maps comparing latest Pleistocene and modern veg-

    etation of the Southwest, based on the packrat middenrecord, illustrate the massive redistribution of speciesand reorganization of plant communities that has oc-curred over the past 11 000 yr (Fig. 2; Van Devenderet al. 1987, Betancourt et al. 1990, Thompson et al.1993). These redistributons were regulated not only byintrinsic dispersal rates, but also by climatic variabilityduring the Holocene. Ongoing plant migrations char-acterize the Holocene and surely carry into the modernera. Population dynamics at the leading (northern) edgeof migrations may confound inferences about human

    impacts on present invasions. Notable examples of on-going migrations related to climate change include ex-pansion of creosote bush ( Larrea tridentata ) in the mid-dle Rio Grande Basin and Borderlands of Arizona andNew Mexico (Grover and Musik 1990, Betancourt1996), Colorado pinyon ( Pinus edulis ) on both sidesof the Rockies (Betancourt et al. 1991, Betancourt etal. 1993), single-needle pinyon ( Pinus monophylla ) innorthern Nevada (Miller and Wigand 1994, Nowak etal. 1994), Utah juniper ( Juniperus osteosperma ) in Wy-oming and Montana (Wight and Fisser 1968; J. L. Be-tancourt, S. J. Jackson, and M. Lyford, unpublished data ), western juniper ( Juniperus occidentalis ) in east-ern Oregon (Miller and Rose 1995), and ponderosa pine

    in the northern Rocky Mountain states (Betancourt etal. 1990, Anderson 1989, Weng and Jackson, in press ).

    Postglacial expansion of pinyon pine offers a well-documented example of broadscale, climate-relatedchange in a species range (Betancourt et al. 1993, Lan-ner and Van Devender 1998). Fig. 3 tracks the migra-tion of Colorado pinyon from the northern ChihuahuanDesert to the Colorado Plateau and along both anksof the Rocky Mountains. Prior to 11 000 14C yr BP,Colorado pinyon was widespread in the northern Chi-huahuan Desert. In New Mexico, USA, its northern

    limits were near Albuquerque and the Sevilleta Long-Term Ecological Research (LTER) Site, coincidingwith the modern northern limits of creosote bush. After11 000 14C yr BP, pinyon was extirpated from desertlowlands and began expanding to higher and morenortherly sites. Many sites in Colorado, USA, werecolonized by pinyon only in the last 2000 yr. The north-ernmost outpost at Owl Canyon, north of Fort Collins,represents a long-distance dispersal event 500 yr ago(Betancourt et al. 1991). Interestingly, this natural in-vasion is taking place in the increasingly urbanizedand disturbed landscapes of the Colorado Front Range(see Mast et al. [1996] for interpretations of tree in-vasion and higher stem densities in the pinegrasslandecotone of the Front Range).

    Owl Canyon offers one paradigm of how recent mi-grations might be imprinted on what we regard as 20thcentury phenomena. This isolated 5-km 2 stand, 200 km

    from the nearest populations to the south, is charac-terized by relatively young trees radiating out from acluster of older pinyons (Wright 1952). The recency of this invasion is also evident in increasing stem density(population growth) during the past 40 yr and second-ary colonizations 230 km from Owl Canyon (Premoliet al. 1994). Secondary outliers are clearly in the earlystages of expansion; the largest population contains 100trees, all 60-yr-old. These expansions probably rep-resent the rising limb of exponential pinyon populationgrowth, which will be followed rapidly by secondarycolonizations and population inlling.

    In the early stages of invasion, an isolated populationlike Owl Canyon may be little more than a curiosity.

    Prior to our paleoecological study, Front Range ecol-ogists speculated on whether it was a relict population,or the result of long-distance dispersal. If we were tofast forward 200300 yr, however, a full-edged pin-yon invasion might be underway in the Front Range.Projecting current rates of urbanization and its effectson ecosystems, would we be able to distinguish naturalfrom cultural causes of invasion in the absence of pa-leoecological perspectives?

    Another source of ambiguity in assigning cause toecological change in the 20th century stems from thelack of regionally scaled, high-resolution demographichistories. Such histories, based on cross dating of birthsand deaths for thousands of trees, are essential to un-

    derstanding demographic responses to decadal-scaleclimatic variability (Swetnam and Betancourt 1998).At the Sevilleta LTER Site, we compared demographictrends in pinyon populations with regional climate re-constructions from tree rings. This comparison sug-gests the following model for the long-term behaviorof these woodlands: Broad-scale mortality during ex-treme, regional drought releases existing seedlings andsaplings from competition for light, water, and nutrientsand opens up niches for recruitment. Recruitment puls-es tend to occur in the rst sustained wet period fol-

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    F IG . 2. As this comparison of modern and Pleistocene vegetation shows, Southwestern landscapes have changed dra-

    matically since the end of the last ice age, 12 000 BP. The Pleistocene map is based on packrat midden vegetation recon-structions (Van Devender et al. 1987, Betancourt et al. 1990, Thompson et al. 1993) and extrapolations using elevations.The modern map is also based on elevations and known distributions of vegetation (adapted from Betancourt et al. [1990]).

    lowing the drought and associated mortality. An ex-ample of this process may be the large pulse in pinyonrecruitment during the last 20 yr. Anomalous seedlingsurvivorship followed the 1950s drought, arguably theworst drought in the last 1000 yr (Grissino-Mayer1996, Swetnam and Betancourt 1998). Pinyon estab-lishment has been sustained by a string of warm, wet

    springs associated with anomalous warming of thetropical Pacic since 1976 (Swetnam and Betancourt1998). This surge in pinyon recruitment will becomeconspicuous on the landscape in another 50 yr andwould likely invoke anthropogenic explanations in theabsence of a comprehensive, demographic history.

    Although re suppression and grazing may be elim-

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    F IG . 3. Pinyonjuniper woodlands presently cover 20 10 6 ha above 1500 m elevation in the western United States,the third largest vegetation type in the contiguous United States. During the late Pleistocene, these woodlands covered whatare now the hot deserts of the southwestern United States, mostly below 1500 m elevation. This diagram shows fossil packratmidden records of pinyon pine along a northsouth transect from Mexico to Colorado, USA. The tick marks on each verticalline represent 350 radiocarbon-dated middens that show the presence or absence of pinyon pines along a 15 latitude( 1600 km) transect from Bermejillo, Mexico (Durango Province) to Fort Collins, Colorado. The diagram depicts the localextinction of pinyon populations growing at desert elevations during the last deglaciation ( 11 000 radiocarbon yr BP) andthe sequential migration to higher elevations and more northerly latitudes during the Holocene (the last 11 000 yr). Note thatpinyons distribution in the state of Colorado may be just a few centuries old and probably is not yet in equilibrium withmodern climate. In Colorado, and probably northern New Mexico, this makes it difcult to distinguish natural migrationduring the late Holocene from historical tree expansion due to re suppression and overgrazing. Tick marks on the y-axesof site graphs represent individual radiocarbon-dated middens, whereas tick marks on the y-axis of the gure indicate

    increments of 1000 yr. Abbreviations: DGO, Durango, Mexico; COA, Coahuila, Mexico; TX, Texas, USA; NM, New Mexico,USA; AZ, Arizona, USA; UT, Utah, USA; CO, Colorado, USA.

    inated as reasons for at least some of the ecologicalshifts in the 20th century, global change may furthermuddle the distinction between natural and cultural ef-fects. Hypothetically, atmospheric CO 2 enrichmentshould affect plant performance in the southwest; someauthors have been quick to suggest that it is alreadyplaying a role in driving encroachment of C 3 trees andshrubs into C 4 grasslands of the southwest (Idso 1992,

    Johnson et al. 1993; see objections by Archer et al.[1995]). The fossil record of the last deglaciation, whenCO 2 increased from 180 to 280 ppmv ( L CO 2 /L), of-fers some perspective. Based on reduced stomatal den-sities on limber pine needles from packrat middens,Van de Water et al. (1994) inferred an increase in wateruse efciency (WUE) in response to the 30% CO 2 en-richment (from 180 to 280 ppmv [ L CO 2 /L]) during

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    the last deglaciation. However, the increase in WUEapparently was not enough to offset increasing Holo-cene aridity, and desert scrub replaced conifer wood-lands throughout much of the southwest. Clearly, cli-mate variability strongly modulates the importance of CO 2 variations in these water-limited ecosystems; e.g.,the frequency and intensity of summer drought willlargely determine the pace of shrub invasion and de-sertication (Betancourt 1996, Neilson 1986). A blan-ket acceptance of large CO 2 effects sends an erroneousmessage to land managers and ranchers that shrub in-vasion of grasslands is inevitable, no matter what theclimate.

    Southwestern paleoecology also instructs us thatEuropeans do not have a monopoly on human impacts.At Chaco Canyon, New Mexico, the Anasazi appar-ently overharvested pinyon populations for fuelwood8001000 yr ago (Betancourt and Van Devender 1981,

    Samuels and Betancourt 1982). In some places, thiswoodland has yet to recover; in others, relatively youngpinyon populations suggest that recovery in these mar-ginal woodlands is sluggish. Given the density and per-sistence of prehistoric humans across the Southwest, itis not farfetched to imagine that, at least in some places,what may seem like invasion is actually recovery. Asimilar case has been made for the apparent invasionof one-needle pinyon ( P. monophylla ) in the Great Ba-sin, which in some areas may actually represent re-covery from 19th century clear-cutting for charcoalmanufacture (Lanner and Van Devender 1998).

    Although highly informative, the long-term histori-cal record has several shortcomings. For example,

    packrat middens are limited to rocky terrain and, andas such, fail to census the open terrain and deeper soilsthat are the typical staging grounds for modern en-croachments (Connin et al. 1998). Historical demo-graphic data are similarly patchy in time and space,and there are important limitations in reconstructingdynamics of forests and woodlands from extant, livingpopulations or dead tees (Gore et al. 1985, Johnson etal. 1994). Moreover, whether the Anasazi or droughtwiped out the Chaco woodland is still subject to in-terpretation. Nevertheless, our brief examples of nat-ural migration, demographic responses to climate, andprehistoric human impact testify that management de-cisions to halt or reverse woody plant encroachments

    or changes in forest structure can easily be misguided.

    Repeat photography: 20th century landscape change

    Historical photographs of landscapes, from foreststo wetlands, are available for practically any area of the western United States (Rogers et al. 1984). As arst approximation, past environmental change can bemeasured by nding the site of a historical photograph,reoccupying the original camera position, and makinga new photograph of the same scene. Differences be-tween then and now provide a basis for identifying and

    quantifying changes, while the new photograph estab-lishes a benchmark for future evaluation (Malde 1973).Repeat photography is a simple, inexpensive, and el-egant tool for reconstructing past environmentalchanges and monitoring future ones; it is particularlywell suited for the relatively open landscapes of thewestern U.S. Rephotography in the southwest, for ex-ample, has focused on key ecological concerns that arerelevant to management of public lands, includingshrub and tree encroachment upon grasslands, climaticeffects on demographic trends in woodlands, postdis-turbance histories, and geomorphic, hydrologic, andvegetation changes in riparian areas (Hastings and Tur-ner 1965, Dutton and Bunting 1981, Rogers et al. 1984,Humphrey 1987, Bahre 1991, Webb 1996, Allen et al.1998). Matched photographs and their graphic evi-dence of change have the unique potential of galva-nizing public dialogue about the timing, magnitude,

    and causes of environmental change.Repeat photography has a variety of limitations thatmust be considered to accurately interpret apparentlandscape changes (Rogers et al. 1984). Early equip-ment had constraints, such as the fading of distant ob- jects on blue-sensi tive plates. Bellows lenses could dis-tort the appearance of an image on the photographicplate. Artistic motives of early photographers some-times resulted in misleading images, and old photo-graphs could have been altered deliberately. Individualphotographs can be unrepresentative of overall land-scape conditions, while retakes may also be selectivelybiased if only unrepresentative scenes are chosen to berelocated. Interpretation errors may occur from an in-

    complete understanding of the natural and cultural pro-cesses responsible for landscape changes documentedin the images. Photographs are also simply unavailablefor many time periods and locations.

    Repeated photographs from the air or space can pro-vide broadscale perspectives of landscape to regional-scale ecological change, thereby overcoming some of the biases of selecting specic, smaller localities. Re-peat aerial photography and other remote sensing data(e.g., satellite imagery) can be ver y useful for assessinglandscape changes in recent decades, particularly whenused in concert with geographic information systems(Mast et al. 1996, Allen 1994, Allen and Breshears1998). For example, repeat aerial photography and geo-

    graphic information systems were extensively used toassess twentieth century landscape changes in the in-terior Columbia River Basin in one of the largest re-gional ecosystem studies ever conducted (Quigley etal. 1996, Hessburg et al. 1999).

    Our example from the Southwest is a repeat aerialphotography study of landscape change in support of a montane grassland restoration project in the JemezMountains of northern New Mexico (Allen 1989, 1994)(Fig. 4). These grasslands have a restricted distributionin northern New Mexico where soil patterns show they

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    F IG . 4. Aerial photographs of the Cerro Grande montane grassland show vegetation changes between 1935 (left) and1979 (right). Recent invasion by ponderosa pine and Douglas-r trees has overwhelmed most of this ancient grassland. Theeld of view is 2 km across each photo.

    have been persistent landscape features for millennia(Allen 1989). Tree-ring evidence indicated that duringthis century conifer trees had greatly encroached uponthe ancient grasslands. Aerial photographs conrmed

    these observations and revealed the extent of the treeinvasion, which reduced the area of open montanegrasslands by 55% across a 100 000-ha region during19351981 (Fig. 4). Comparative study of tree de-mography, soils, re, climate, and land use history atmultiple sites across the study area showed that treeinvasion was a modern process caused primarily byhumans. Specically, tree invasion was due to changesin livestock grazing and re exclusion, and not climaticchanges (Allen 1989).

    Before this historical assessment of tree invasion andits causes, neither ecologists nor local land managersrecognized that major vegetation changes had recentlyoccurred in these grasslands. Initial assessments by ex-perienced ecologists focused only on the current con-guration of forests and grasslands, noting that theseareas had the highest grass density and productivity ina forest understory that they had ever seen in the South-west (Potter and Foxx, unpublished manuscript ). Thedense understory coverage appears less unusual onceit is recognized as a formerly open, montane grasslandthat has been invaded by trees. Lacking a historical-ecology perspective, the National Park Service failedto discern the anomalous, human-induced changes inthe ancient vegetation patterns, and so incorrectly be-

    lieved that the best management option was to leavethe area in its current state.

    Now, a restoration program has been initiated basedon (1) historical knowledge that the ancient grasslands

    were declining in extent because of 20th century humanactions, and (2) the National Park Services mandateto preserve natural biotic communities. Historical ev-idence of grassland extent early in the twentieth centuryand tree recruitment patterns in the area have servedas objective guides to the restoration procedures in-volving tree cutting and prescribed burning. Managerson the adjoining Santa Fe National Forest have recentlyconducted a similar grassland restoration project in anearby area and are considering other sites for treat-ment. The multiple lines of historical evidence mar-shaled in this case were scientically compelling, butthe visual presentation of the extensive tree invasionclearly evident in the air photo time series (Fig. 4)probably had the largest effect in mustering supportfor grassland restoration among resource managers andthe public.

    Fire scar histories: local and regional patterns

    Prior to the 19th century westward expansion andEuro-American settlement in North America, frequent,low-intensity surface res were a dominant ecologicalprocess in most pine and mixed-conifer forests (Agee1993, Swetnam 1993, Swetnam and Baisan 1996). De-tailed and long chronologies of surface res can be

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    assembled by sampling re scars preserved within theboles of surviving trees (Swetnam and Baisan 1996,Arno et al. 1997, Fule et al. 1997, Brown et al. 1999).Most trees producing annual growth rings have alsodeveloped a physiological capacity to grow new ringsover injuries caused by res. This capacity, plus theenhanced ammability of exposed wood and seepingresin within previously formed wounds, often leads tomultiple, overlapping re scars on individual trees. Thecalendar years, and usually the approximate seasons,of past res can be determined by cross dating theannual rings and by carefully observing the position of re scars within the dated rings (Dieterich and Swet-nam 1984). Compiled re scar records from numerousspecimens collected throughout forest stands often pro-vide nearly complete inventories of re events overperiods of centuries (Baisan and Swetnam 1990, 1997,Swetnam 1993, Fule et al. 1997, Brown et al. 1999).

    Fire scar based re histories are limited in a numberof important ways. First, as with most historical re-constructions, they depend upon the development andpreservation of the record; in this case, re scarred treesthat can be dated by dendrochronological techniques.Hence, re history reconstructions are generally pre-cluded across large areas by the absence of annual ringproducing trees or shrubs. Past harvesting and recent,intense res can also remove the accumulated re scarrecord from some landscapes where it might have pre-viously been preserved in the resinous lower boles of living and dead trees (i.e., snags, stumps, and logs).The re scar record generally decreases in complete-ness with increasing time before the present because

    of burning and decay processes. Another limitation of tree-ring-based re history reconstructions is relativelylow spatial resolution of past re perimeters. Firescarred trees are essentially point recorders of reevents, and they probably do not record every re thatburns near their base. In low to moderate severity sur-face re regimes, there may be no other recoverableevidence of past res than a fragmentary and incom-plete re scar record.

    Although these problems constrain our ability to re-construct or use re history in some locations and forsome purposes, the tree-ring record has an advantageof high temporal resolution. Seasonal or annual datingresolution of re scars enables re historians to eval-

    uate patterns of re event synchrony across multiplespatial scales. From evaluation of these patterns, it ispossible to make reasonable inferences about past reextent, and human vs. natural causes of change (Allenet al. 1998). Networks of systematically collected re-scarred trees, for example, have been used to infer reextent within forest stands (Arno et al. 1997, Fule etal. 1997) and across large landscapes (Allen 1989,Brown et al. 1999).

    Our example from the Southwest is a composite of re event chronologies from very widely dispersed

    mountain ranges over the entire region. The regionalre years evident in the southwestern composite (Fig.5), were almost certainly related to regional climaticpatterns, since no other known regional-scale factor islikely to have caused such a high degree of repeatedre synchrony at this spatial scale over such a longperiod of time. Comparison of these multicentury chro-nologies with independent tree-ring-based drought re-constructions conrms that extreme climatic oscilla-tions of wet and dry years were strongly related toregional re years (Swetnam and Betancourt 1990,1998).

    Numerous re and land use histories in the South-west indicate that intense livestock grazing was theinitial cause of decline in natural re regimes at re-gional and local scales in the late 1800s (Leopold 1924,Swetnam and Baisan 1996, Swetnam et al. in press ).The general pattern documented in these histories was

    a decrease in widespread, frequent res that coincidedclosely with the beginning of intense grazing by largenumbers of sheep, goats, cattle, or horses in that par-ticular area (Leopold 1924, Dahm and Geils 1997,Swetnam and Baisan 1996, Swetnam et al. in press ).While these events occurred during the late 1800s inmost locations, a few exceptions offer further supportto the interpretation that livestock grazing was the ini-tial cause of re regime disruptions. A few sites innorthern New Mexico and Arizona, for example, showre frequencies declining in the early 19th century (orearlier) and corresponding to within a few years of thedocumented timing of pastoral activities by Spanishcolonists and Navajos in these areas (Allen et al. 1995,

    Touchan et al. 1995, Baisan and Swetnam 1997). (Notethe decreased re occurrence in the early 1800s in theuppermost chronology labeled (a) in Fig. 5 from tra-ditional Navajo pasturelands in the Chuska Mountains[Savage and Swetnam 1990].) In contrast, remote foreststands surrounded by lava ows, with no evidence of intensive grazing, sustained some surface res into themiddle of the 20th century, when aerial re-ghtingresources began to be most effective in suppressingres (chronologies labeled (b) in Fig. 5, from El Mal-pais National Monument [Grissino-Mayer and Swet-nam 1997]). Finally, a remote mountain in northernSonora, Mexico (lowermost re scar chronology la-beled (c) in Fig. 5), where neither intensive livestock

    grazing nor effective re suppression occurred, showsepisodic surface res burning throughout the 20th cen-tury in the highest elevations of this range (Swetnamet al. in press ). A similar re history from remote areasnear Durango, Mexico shows continued occurrence of surface res until the middle of the 20th centur y (Fuleand Covington 1997).

    One of the ways that re history research has beenuseful to managers in the southwestern United Statesis in providing the primary evidence of frequent surfaceres in pine and mixed conifer forests before circa

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    F IG . 5. Composite re scar chronologies from 55 forest and woodland sites in Arizona, New Mexico, and northern Mexico(AD 1600present) reveal regional synchrony. In the upper chart, each horizontal line represents the composite re chronologyfrom a different site, and the tick marks are the re dates recorded by 10% re-scarred trees (and 2 trees) within thatsite. The long tick marks are re dates recorded by 10 sites in the southwestern network. Note that when adjacentchronologies

    have long tick marks for the same year, i.e., a regional re event, this forms a continuous vertical line between the chronologies.Most of the sites were in ponderosa pine or mixed conifer forests, and the sampled areas typically ranged from 10 to 100ha, although a few sites exceeded 1000 ha. The average number of trees sampled per site was 20, with a maximum of 56trees. See the text ( Examples from the southwestern United States: Fire scar histories ) for explanations of the specic rechronologies that are labeled (a, b, c) on the right side. The line graph on the bottom shows the total number of sites recordingre dates each year. The labeled years with arrows were regional re dates, i.e., res occurred in 10 sites across the network.

    1900. These histories also indicate that large, high in-tensity, stand replacement res in southwestern pon-derosa pine forests were rare. Other historical data sup-porting this interpretation include forest age structureand stand density reconstructions from dendrochro-

    nology, repeat photographs, historical writings, andearly land survey records (Covington and Moore 1994).These data also show that the end of frequent surfacere regimes was followed in subsequent decades bymassive tree irruptions, leading to highly anoma-

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    lous, dense stands of dog hair thickets (Covington andMoore 1994, Fule et al. 1997). These thickets are nowa key factor contributing to catastrophic, stand-replac-ing res in some areas (Covington and Moore 1994).Twentieth century re statistics from the Southwestshow clear trends of increasing number, size, and in-tensity of such catastrophic res (Swetnam 1990, Dahmand Geils 1997, Swetnam and Betancourt 1998). Thecombination of all of these lines of historical evidenceis the strongest argument managers have for forest res-toration initiatives involving tree thinning and reintro-duction of natural re patterns (Moore et al. 1999).Without such compelling historical evidence, publicopposition would probably preclude most ecologicalrestoration work.

    Thus, at the most basic level, re history researchhas been a major impetus and justication for reintro-duction of surface re in pine and mixed-conifer for-

    ests. This is especially the case in landscapes wherenatural-process maintenance is legally mandated, suchas in designated wilderness areas and in some park lands (Christensen 1989, Parsons et al. 1986, Allen etal. 1995). Even on landscapes where restoration orpreservation of naturalness per se is not a goal, rehistories still provide basic information about the mag-nitude and extent of changes that have occurred in the20th century, relative to earlier centuries. This infor-mation is often crucial for determining if current con-ditions are practically sustainable, or if they are soanomalous in both a historical and ecological sense,that ecosystem sustainability is degraded.

    Although re histories have so far had the greatest

    impact in simply justifying re restoration programs,ultimately, the specic characteristics of past re in-terval distributions may also serve as a guide to rerestoration programs involving repeat, interval burn-ing. Measures of central tendency (e.g., means, me-dians, modes, etc.), range, and higher moments (e.g.,variance, skewness, etc.) of re interval distributionsdetermined from re chronologies, for example, mayprovide a basis for modeling, planning, and imple-menting re restoration programs designed to emulatenatural processes. An ecological reason for concerningourselves with the distinctiveness of historical re in-terval distributions on particular landscapes is that thecharacter of these distributions is intimately linked with

    the presence or absence of certain species (e.g., seeClark 1996), as well as the age structure, composition,and spatial arrangement of extant communities (Arnoet al. 1997, Fule et al. 1997, Brown et al. 1999).

    A potential application of re history studies is toidentify regional-scale, seasonal to decadal climatererelations that might be used in re hazard forecastingmodels. Using historical data, for example, several re-searchers have found that years of increased or de-creased regional burning were closely related to theextreme phases of the El NinoSouthern Oscillation

    (ENSO) (Simard et al. 1985, Swetnam and Betancourt1990, 1998, Brenner 1991). In the southwestern U.S.,extreme El Nino events typically result in increasedcool season moisture with reduced re activity in thefollowing spring and summer. These wet conditionsalso lead to growth and accumulation of ne fuels im-portant to re ignition and spread in subsequent dryseasons. A key situation for regional re years is thecontingency of a series of wet years, reduced re ac-tivity, and fuel accumulation, followed by a La Ninaevent when precipitation is reduced and extreme burn-ing conditions arise (Swetnam and Betancourt 1998).The inverse of this reclimate pattern may exist inthe Northern Rockies and Pacic Northwest, becauseENSO-related precipitation amounts tend to be oppo-site between this region and the Southwest (Dettingeret al. 1998). Useful interseasonal re hazard forecastingsystems might be developed using a combination of

    lagged climatefuel patterns and lagged ENSOpre-cipitation relations.

    A DVANTAGES AND C ONSTRAINTS OF A PPLIEDH ISTORICAL E COLOGY

    Historical ecology can be very useful to manage-ment, but there are number of constraints in its appli-cation to management problems. Specic limitations of data and methods were briey pointed out in our ex-amples from the Southwest. Now, we will further dis-cuss several general advantages and constraints in us-ing history.

    Predictability and historical context

    Although direct or simple extrapolation of historicalpatterns or trends into the future is usually erroneous,history can be useful for guiding the development andtesting of predictive models. The re and climate his-tory ndings in the Southwest are encouraging in thisregard, because they indicate that there is some poten-tial for using historical knowledge to help developlong-range, re hazard forecasting models. Part of thispotential derives from recent understanding that theglobal climate system includes oscillatory dynamics(e.g., ENSO, North American Oscillation, and Quasi-Biennial Oscillation), driven by long temporal lags inthe transfer of energy (heat) through and between theoceans and the atmosphere. Improved physical knowl-edge of these seasonal to decadal-scale patterns andtheir impacts on regional climates have resulted in ma- jor improvements in long-range climate forecastingmodels (Barnett et al. 1994, U.S. Global Change Re-search Program 1999).

    Historical climatology has played an important rolein this scientic advance by discovering the existenceof some of the long-term, regional- to global-scale pat-terns, and by providing important clues to the under-lying mechanisms. Knowledge of past patterns oftengenerates hypotheses or interpretations that can be test-

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    ed and evaluated with modern observations, experi-ments, and models. Historical ecology offers similaropportunities for evaluating our understanding of eco-systems by comparing model simulations with histor-ical reconstructions of processes and structures (e.g.,Anderson 1995, Miller and Urban 1999). Although pre-diction (or forecasting) of many ecological processes,particularly chaotic ones, probably will never be pos-sible, it is premature to assume that this is true for allsituations. Some forest insect population outbreaks, forexample, are quasiperiodic and inuenced by long-termclimatic uctuations (Swetnam and Lynch 1993).Therefore, they might hold some promise for predict-ability with statistical or mechanistic models.

    Rather than a direct use of history for prediction,most historical ecologists emphasize the importance of knowing history, because it informs us about what ispossible within the context of certain locations and

    times, and it places current conditions into this context.This knowledge often informs us about the potentialcauses of change and the historical pathways thatbrought ecosystems to their current condition (Kauf-mann et al. 1994, Morgan et al. 1994, Swanson et al.1994). Because history provides multiple points of tem-poral reference, it can also tell us if current conditionsare highly anomalous and, therefore, in some casesdeserving of greater priority in management decisionsand actions. Providing a historical context for evalu-ating present conditions may be one of the most im-portant uses of historical knowledge as we face in-creasing concerns about human-caused environmentalchanges in the 20th century (e.g., global warming). For

    example, the unusual rate and magnitude of 20th cen-tury atmospheric CO 2 enrichment is placed in historicalcontext by the records from monitoring at Mauna Loa,Hawaii, USA, and from gas bubbles trapped in Green-land and Antarctic ice. We also know from historicalstudies that the 1990s included the warmest years of the 20th century on a global scale (U.S. Global ChangeResearch Program 1999) and that it was probably thewarmest decade in the Northern Hemisphere over thepast 600 yr (Mann et al. 1998). In light of the enormousconsequences of these environmental changes, it is cru-cial that we evaluate them within a long-term historicalcontext.

    Fragmentary nature of historyIn our view, the most important limitation of his-

    torical ecology is that the record of the past is oftenbrief, fragmentary, or simply unobtainable for the pro-cess or structure of interest. Almost all historical re-constructions also suffer from a fading record prob-lem; i.e., the completeness and reliability of the recorddecreases with increasing time before the present. Thisis usually due to progressive loss of older evidence.Most historical-ecology studies partially deal with thislimitation by truncating the period of analyses to the

    best replicated and most reliable period. When histor-ical reconstructions are possible, they are often limitedto only a few ecological variables or processes. In thebest circumstances, these are reconstructions of essen-tial processes, such as climate and re for many ter-restrial ecosystems.

    Although lack of historical evidence can not be rem-edied for many localities and specic ecosystem pro-cesses, it is usually possible to assemble time series of at least some relevant variables at broader spatial scales(e.g., regions) using spatial networks, as in our packratmidden and re history examples. Historical processesand patterns for specic locations and ecosystems maynot be recoverable or justiably extrapolated frombroader scale histories. When lacking local historicalevidence, ecologists and managers should be particu-larly cautious about making assumptions based onknowledge of historical pathways of vegetation change

    from other regions or ecosystem types.Because of the fragmentary nature of the historicalrecord, and uncertainties about causation and the po-tential role of unmeasured variables, the highest degreeof condence is achieved by combining multiple, in-dependent data sources and historical methods (e.g.,our example of tree invasion in montane grasslands).Converging lines of historical evidence from many lo-cations are most convincing, particularly when coupledwith sound knowledge of mechanisms established bymodern observations and experiments.

    Nonequilibrium paradigm

    Other challenges to the usefulness or relevance of

    history derive from the nonequilibrium paradigm. If ecosystems are necessarily dynamic, then it may bemisguided and fruitless to choose a single xed pointor period of time in the past for establishing a static,desired future condition (Sprugel 1991). The Holocenehistory of southwestern plant communities is a graphicexample of the remarkable changeability of ecosystems(Fig. 2). The use of historical information for identi-fying desired future conditions, however, does not im-ply management for static conditions. The importanceof temporal change is partly reected in the phraserange of natural variability (or historical range of variability), i.e., the focus is not on a single condition,but on a range of conditions and the variability underwhich ecosystems were sustained in the past. Recog-nition of the importance of maintaining or reestablish-ing the changing nature of ecosystems is reected in apolicy adopted by some natural re programs inparks and wilderness areas. In the Sierra Nevada Na-tional Parks, for example, restoration of the historicalor natural re process is preferred over restoration of a particular ecosystem structure that existed at somepoint of time in the past (Parsons et al. 1986, Chris-tensen 1988, 1989, Parsons 1990). Natural re regimes,however, cannot be safely or effectively reintroduced

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    in many ecosystems without rst restoring related for-est structures (see Stephenson 1999, Moore et al. 1999).

    Ultimately, decisions about desired conditions,and what is natural are inherently subjective andvalue laden. History can inform this decision process,but it does not provide unequivocal policy or manage-ment guidelines (Foster and Motzkin 1998, Schmidt etal. 1998). People must decide if past mechanisms of change and the dynamical processes involved are de-sirable for todays landscapes. There are some land-scapes where maintenance or restoration of natural orhistorical processes and structures are probably desiredby most people and are legally mandated (e.g., des-ignated wilderness and some parks). There are manyother places where it is very difcult to determine whathistorical processes and structures are compatible withcurrent human desires, as well as being necessary tosustain ecosystems. We suspect, however, that the high

    costs and consequences of excluding necessary eco-logical processes (e.g., re) will soon shape humandesires and decisions more than they have in the past.

    Humans and nature

    A problem with using history to identify naturalconditions is the fact that humans were an importantcause of past variability in most ecosystems (Shrader-Frechette and McCoy 1995, Hunter 1996). Our south-western research, for example, indicates that NativeAmericans were probably very inuential, at least atlocal scales, in the pre-Columbian era (e.g., at ChacoCanyon). Extreme re regime changes in some forestswere also caused by human activities in the 1920th

    centuries (e.g., livestock grazing and active re sup-pression). Pre-Columbian human impacts on NorthAmerican landscapes have often been underestimated(Denevan 1992), but there seems to be a tendency inrecent years to overgeneralize these impacts (Vale1998). Population densities, resource demands, andconsequent ecological effects of humans were highlyvariable through space and time (Russell 1997). Somelandscapes were managed and manipulated for millen-nia, while it is likely that some large regions and eco-systems were not substantially altered by humans untilthe 19th or 20th centuries (Vale 1998).

    It is obvious that many ecosystems are greatly mod-ied today by humans, and for many areas restorationof historical processes or structures is impossible orundesirable. In these areas, we often trade off ecolog-ical integrity for short-term economic gains, as manyecosystem components (e.g., biodiversity) and pro-cesses (e.g., ecosystem services, like water purica-tion) are lost in humanized landscapes. Moreover, hu-man needs are increasing, and humans will largely de-termine the future of all ecosystems, perhaps regardlessof history. Despite the prevalence of high temporalvariability in many ecosystems, whether caused by hu-mans or climate, this does not mean that we can prac-

    tically manage and sustain ecosystems under any con-dition that we choose, regardless of the inherent limitsand tolerances of species and communities. As pointedout by Pickett et al. (1992) and a recent EcologicalSociety of America report:

    . . . Human-generated changes must be constrained because nature has functional, historical, and evo-lutionary limits. Nature has a range of ways to be,but there is a limit to those ways, and therefore,human changes must be within those limits.

    Christensen et al. (1996)

    A primary objective of historical ecology should beto help dene those limits. In the face of imperfectunderstanding of ecosystem functional and evolution-ary limits, knowledge of historical patterns and pro-cesses can be of considerable value. Applying histor-ical knowledge to guiding and constraining manage-

    ment actions is then an informed and conservative ap-proach to managing ecosystems.

    A CKNOWLEDGMENTS

    For discussions and ideas we thank all participants of a1996 workshop held near Missoula, Montana, USA, spon-sored by the U.S. Forest Service, Rocky Mountain ResearchStation, and Aldo Leopold Wilderness Research Institute. Wealso thank Chris Baisan, Wally Covington, Amy Hessl, EmilyHeyerdahl, Steve Jackson, Kurt Kipfmueller, MargaretMoore, Dave Parsons, Matt Rollins, and two anonymous re-viewers for comments on review drafts. Our research wassupported by the U.S. Geological Survey, U.S. Forest Ser-vice, National Park Service, The Nature Conservancy, TheUniversity of Arizona, and the National Science Foundation(Geography, Atmospheric Dynamics, and Earth System His-tory programs).

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