Applied zooarchaeology the relevance of faunal analysis to wildlife management

R bee Lagman

Abstract

Zooarchaeology has the opportunity to expand its analytical horizons into the little cxplored rcalrn of modern wildlife management by applying the knowledge it gains from its unique perspcctivc of prehistory Ways in ahich animal populations thscatcned with extinction might be protactcd in perpetuity identifying which Corms or taxa should bc reintroduced to which areas to recreate natural biotas identifying which forms or taxa arc exotic and should bc removcd from an area to create a natural biota and helping to define the boundaries of biological preserves meant to p r e s c r ~ ~ c biota in perpetuity are all subjects to which knowledge gaincd through zooarchaeological research rnight be applied The potential benefits include better informed wildlile management decisions Ccwcr extinctions less loss of biological diversity and increased job opportunities for zooarchaeologists

Keywords

Biological reserves exotic species extinctions nativc spccics reintroductions wildlife managcmcnl

Introduction

Zooarchaeologists have in recent years displayed a remarkably increasing sophisticatiori in analytical techniques and are thus providing an ever-growing set of insights into past human behaviours Much of this sophistication has come from greater attention to taphonomic issues (eg Fisher 1995 Gifford-Gonzalez 1991 Lyman 1994b) and from more detailed ethnoarchaeological studies focused on faunal remains (eg Hudson 1993) Yet the questions we ask of the zooarchaeological record and the problems we seek to solve with faunal data have remained essentially constant virtually since the inception o l zooarchaeological analysis (Robison 1948) The determination of past subsistence systems and paleoecological conditions remains at the forefront of modern zooarchaeology (eg Brewer 1992) although chronological concerns have also benefited from zooarchaeologi- cal research throughout the history of archaeology as a discipline From Edouard Lartets mid-nineteenth-century division of the French Palaeolithic into several mamma ages to

World Arckzaeoogy Vol ZB(1) 110-125 Zooarchaeology (GIRoutledge 1996 0043 -8243

The relevance of faunal anulysis to wildlife management 11 1

modern biostratigraphic analyses such as Cookes (1976) study of suid remains prompting redating of volcanic tuffs in Plio-Pleistocene hominid sites of Africa zooarchaeology has contributed much more to our understanding of the past than just our recreations of the interaction sphere that included people and the fauna with which they were in contact (Olsen and Olsen 1981 193)

In this paper I argue that zooarchaeologists are in a unique position to move beyond tht important anthropological and archaeological issues they seem to prefer and to broaden the scope of their inquiries to include issues that are today and will be in the future important to humanity Specifically I outline here what I perceive to be a little explored avenue of zooarchaeological research --an avenue that could provide not only new sources of funding for zooarchaeologists but new sources of jobs for them as well

Archaeology is generally taught as a part of anthropology in North America In North America anthropological research seeks not only to increase our understanding of bu~ also to better the human condition From the anthropology of the homeless AIDS (acquired immune deficiency syndrome) and domestic violence to the anthropology of criminal behaviour - wherein archaeological forensics or taphonomic research plays a major role - anthropology can and does make a major contribution to making the world what we hope will be a better place Archaeology in general and zooarchaeology in particular with the exception of forensic research may appear to have little to offer in this respect But because so many modern potential ecological problems facing humanity concern our impact on biological phenomena - destruction of the rain forest decreasing harvests of marine fishes and the like - zooarchaeology as well as paleoethnobotany clearly has a major role to play in the resolution of those problems

In the following I describe several instances of the significance of zooarchaeological (and paleontological) data for addressing and helping to resolve particular kinds of modern problems faced by those concerned with wildlife management Many nations have one or more forms of national park andlor game preserve Many of these spatial units are managed in such a manner as to protect i n perpetuity the biota included within their boundaries Sometimes attempts are made to enhance or modify the biotas of these parks and preserves so that they reflect what are variously referred to as pristine ecosystems wilderness or native (pre-industrial) or natural biotas Further many nations have various legal controls on hunting and trapping of animal species and legal controls over which species can be harvested and which are to be unmolested Managing wildlife sometimes involves supplementing resident populations with individual organisms transplanted from elsewhere or attempting to re-establish species in areas where native populations have become extirpated With its unique perspective on the time depth of ecosystems and biotas zooarchaeological research can be applied to ensure that wise wildlife management decisions are made

The kinds of wildlife management problems that zooarchaeological research might help resolve are many They are however often very specific to I particular geographical place andlor species The examples that follow are those with which I am most familiar and thus centre on North America Examples from other continents and regions surely exist None the less I have attempted to select examples that illustrate a broad diversity of wildlife management concerns and how particular kinds of zooarchaeological research might help resolve them

112 R Lee Lymar~

Populations threatened with extirpation

One of the major issues facing wildlife managers working in national parlts kind wiidlifc preserves involves ensuring the wildlife species undcr their carc survivc in perpcinit) Thus wildlife managers attempt lo identify measures thai can be taken ao ensure i81r1t extirpation of species resident on park or preserve lands does not occur ltdne virYal~ic presently being debated in North Amcrican wildlifc management that weighs heavily or1

this issue concerns what is known as SLOSS Simply put should i park or preserve corrsisi of Single Large Or Several Srriall pieces of landscape (see for example Quinn a n d Harrison [I9881and references therein) Thus far the debate his employed daia o n t h e modern distribution of taxa and has typically inferred that tensporal processes havc C I U S C ~

these spatial patterns Zooarchaeological data brought to bear con this dcbate can 111akc 111c temporal inference empirical

Grayson (1991) reviewed the biogeogr-aph~c history CPC smali mamrnals in ihr Gteat Basin physiographic province of western North America and then used zooarchacologicaI and paleontological data to test the implications of a model first proposed by bingiiogra- phei-JFI Brown (197 1 1978) Hm simplest terms Brown rnodel suggests that the i-irotiern distribution of subalpine and alpinc s~nall mammals (c pika [Ocllororrrprinreyial golden mantled ground squirrel [Sper~nophilus llrterulis] ermine [Muslelcl ertvaanriij) oil

Great Basin mountaintops was the result of Pleistocene colonization of those rnountairl followed by Holocene extinction of intermediate lowland populations arid by difcrcrstir extinction of various mountaintop populatioras Qrag~scgtn (1991 372) found that Browsrs model has impressive paleontological support and T note that much of that suppor~ derives from zooarchaeological research

Grayson (1991 373) noted that confirmation of Browns model with thc fossii rccerrci has clear implications for the management of high aleie~ule environr~~erats in the Circlt Basin He was referring to the fact that nnany populations of small mammals iur ahis liaa

are today isolated on mountaintops Basically following thc theory of island biogeography (MacArthur and Wilson 1967) as modified over the years (eg Brown 1986) b3rayrtsrii study underscored -- using zooarchaeological data - the fact that such small gcncticaliy isolated populations of species tend to I-~e more prone inr extinction than larger aa-lciir non-isolated populations (see also Korn 1994) Larger populations provide suificicrii genetic variation Lo ensure survival of a species and larger areas allow migration betwecn population node because corridors bctween thcni are shorter Either isolation of tliose nodes rrom one another or restriction of those node to small geographic areas - s u c l ~as mountaintops or both increases the probability that one or more of thcpse na~dcs wili-

cease to exist Thus the biogeographic histories of alpine small rnanirnals of the Griit

Basin indicate that neithcr alternative of SEOSS is u~orkablc herc We cannot enlarge tile

insular alpine and subalpine habitats found on Great Basin mountaintops bounalarie a i thosc habitat patchcs arc climatically dictateai And these habitat patches are not only small they are also multiple in number Ensuring that thesc patches are not artiliciirlly disrupted in the future would seem to be the only way to guarantee survivii of tl~eir

included small mammals Migration corridors betwecn the habitat patches canraoi hi artificially constructeci or maintained If such corridors could be built where shoa~ldthe

The relevance of faunal analysis to wildlife management 113

be located The fossil record might help us determine where these corridors existed in the past as can be illustrated with another example

The pygmy rabbit (Brachylagi~s idahoensis) is a diminutive leporid with a modern range restricted to two areas of the western United States One area is relatively large and encompasses portions of four states northern Nevada western Utah southern Idaho and southeastern Oregon The other historically documented area where this species occurs is the central portion of eastern Washington state Populations in the two areas are presently isolated from one another The wildlife management coricern here is that the eastern Washington population has over the past forty years shrunk to an alarmingly small size prompting the Washington Wildlife Commission to list this population as threatened and to suggest that i t be listed as endangered (McAllister and Allen 1993) and to develop a management pan with the aim of ensuring survival of the population (McAllister and Allen 1994) The zooarchaeological record of this species provides insight to why this population is threatened with extirpation and how i t might be preserved

The zooarchaeological record for pygmy rabbits indicates this species probably occupied a wider range in central Washington during the Holocene than i t presently does Remains of this species have now been recovered from eleven archaeological and two paleontological sites in the area (updated from Lyman 1991) Some of these are extrali~nitalrecords that is some remains have been recovered from geographic locations where pygmy rabbits have not been historically documented to occur In brief the fossil (botanical and faunal) record suggests that when big sagebrush (Artemisia tridentata) was more widespread in central Washington -- during the middle Holocene climatic interval known as the Altithermal - pygmy rabbits were also more widespread When the Altithermal ended about 4500 years ago the range of sagebrush shrank and so too did the range of pygmy rabbits Then during the late nineteenth and early twentieth centuries land was cleared of sagebrush for agricultural purposes This prompted a second diminution of sagebrush and pygmy rabbit range

Ecological studies of pygmy rabbits indicate that this leporid is dependent on big sagebrush for food and for shelter from predation (Green and Flinders 1980a 1980b) The zooarchaeological record in conjunction with paleobotanical data such as that from pollen records confirms this as both species appear to have responded to climatic change in like manner Washington Department of Wildlife biologists have made use of both modern and zooarchaeological data in their planning efforts (McAllister and Allen 1993 1994) The fossil record suggests that the prehistoric source of Washingtons pygmy rabbits resides in the area of the larger extant population to the south Late Pleistocene fossils of pygmy rabbits recovered from a paleontological site between the two areas (Rensberger and Barnosky 1993) tend to confirm this suggestion While i t would probably be impractical to develop a migration corridor between the two populations today it would also be impossible given the climatic history of the area The probable corridor between the two today is characterized by vegetation habitats that are not conducive to tht survival of pygmy rabbits The zooarchaeological record does indicate however that the extant population might be supplemented by individuals transplanted from southeasterri Oregon and that the maintenance of habitats dominated by big sagebrush is critical to the survival of the species

114 R Lee Lyman

When managing small isolated populations we may not have many options other than to preserve those populations as they presently exist Yn some cases management efforts have failed and populations have been extirpated When this occurs one management alternative is to transplant individuals from other populations to the vacated rangc in an attempt to re-establish a population Zooarchaeological research can also plav an important role in these situations

Re-establishment of extirpated populations

Eighteenth- and nineteenth-century conimercial exploitation of sea otters (Enhydru lirtris) along the western coast of North America ultimately led to the extirpation of this species from much of its range (Kenyon 1969 Ogden 1993 Roest 1973) Attempts to re-establish populations along the middle latitudes of that coast took place in the 1970s and thus far have met with less than perfect success (Jameson et al 1982 1986 MacAskie 1987) The typically expressed reason for the limited success of these efforts is that ecosystems to which sea otters have been transplanted have heen modified too exter~sively by modern human industry to support them Y wonder i f however part of the reason might resicle in a factor that only zooarchaeological research car measure

Sea otter skulls collectetl from much of the once occupied eastern Pacific range of sea otters display various character gradients from the Aleutian Islands to southern California Some of these characters have been used to argue for or against the existence oi multiple subspecies of sea otter (Davis and Lidicker 1975 Roest 1973 1979 Scheffer and Wilke 1950 Wilson et al 1991) Final resolution of this debate will be critical to futtlrc wildlife management activities first because national legislation in the United States concerns species subspecies and populations (OWrien and Mayr 1901) and also beca~ase of conflict between one population (a possibly distinct subspecies) of sea otters and the modern fisheries industry (Wendell et al 1986) lhe significant aspect to all of this Is the transplanting of sea otters ILegally only forms (ecotypes or subspecies) native to ar arcs should be introduced there during efforts to re-establish populations This also m~aites biological sense as i t is likely that those native forms unique to an area are the product of millennia of evolution

The International Union for the Conservation of Nature and Natural Resources PUCN (recently renamed the World Conservation Un~on) has drafted guidelines Cor such -

reintroductions These read in part

An assessment should be made of the taxonomic status of individuals to ba re-introduced They should be of the same taxonomic unit (and ideally closely related genetically) as those which were extirpated An investigation of historical information about the loss and fate of individuals from the re-introduction area as well a mmolecuiar genetic studies should be undertaken in case of doubt Release stock idealiy should be closely-related genetically to the original native stock

(Anonymous 1992 2-31

Clearly a search of not only historic but also prehistoric information nnay be necessary The rapidly developing study of ancient DNA extracted from prehistoric skeletal tissues

The relevance of faunal anuljlsis to wildlife matlugement 115

(eg Richards et al 1993 1995 and references therein) seems to have great potential for contending with IUCN guidelines The technique is applicable to curated as wtll as newly acquired archaeological specimens It has however not yet been applied to Oregon coast sea otter remains

Given the lack of genetic material from eighteenth- and early nineteenth-century sea otters on the Oregon coast the problem becomes one of establishing the age of the character gradients displayed by eastern Pacific rim sea otters (see Wilson et al 1991 for a recent summary) and determining if those gradients have shifted spatially through time IIf those gradients have a prehistoric distribution in time andlor space similar to that documented using historic specimens a potential problem is raised If a particular form -represented by a particular character state -- of sea otter was present prehistorically in one location within the range and another form was present at another location then transplanting individuals from the former to the latter may be doomed from the start In such a case failure to re-establish a population could result on the one hand because the transplanted form is not well adapted to its new environment the extirpated form on the other hand is adapted to the environment of that location hence its difference in form

Several years ago I examined prehistoric sea otter remains recovered from archaeologi- cal sites on the coast of the state of Oregon - the middle latitudes of the historically documented range of sea otters (Lyman 1988b) The size of sea otter teeth recovered from those sites are more or less intermediate to those from Alaska (large) and those from southern California (small) but they also vary somewhat from this simple pattern Prehistoric Oregon sea otters match modern Alaskan sea otters in some characters in others they match modern California sea otters Precise details are unnecessary to the point I wish to make here which is Were the sea otters captured in Alaska and translocated to the Oregon coast in the 1970s of the proper phenotype (or genotype) Zooarchaeological evidence suggests they were not If this is a correct interpretation then perhaps those transplanted sea otters were doomed from the moment they were captured Whether or not my suggestion proves to be true or false and it is unclear which way the cards might fall given that we still have much to learn about this species (VanBlaricom arid Estes 1988) the significance of zooarchaeological data should be clear

Not only can zooarchaeological data inform us as to which form of a species might be the most appropriate one to transplant to an area such data have the potential to indicate the potential success of such transplant efforts Even if the appropriate form to transplant is clear this does not guarantee that the introduced form will survive in its new home This is precisely the argument offered by biologists working with sea otters on the Oregon coast The data substantiaiing their argument can be supplemented with zooarchaeological data An example of this is provided by the California condor (Gymnogyps cnlifornianus) of western North America a species presently near extinction (Emslie 1987) Zooarchaeo- logical and paleontological data suggest this species enjoyed a wider range during the late Pleistocene but that range shrank considerably with the onset of the postglacial Holocene epoch (Emslie 1987 Simons 1983) Those data also suggest that this species was a carrion feeder and that the terminal Pleistocene extinction of many genera of large mammals probably contributed to the decline in frequency and range of the California condor (Emslie 1987) Finally paleoecological data indicate attempts to reintroduce this species to portions of its prehistoric range would probably fail because of major differences

116 R Lee Lytriurz

between late Holocene floral and faunal communities and late Pleistocene commaanities (Emslie 1987)

In the absence of historic data or even with it (see below) zooarchaeological infor- mation may be critical to ensuring that the correct forms of animals are transplanted The wrong forms may be i l l suited to their new habitats or they may be well adapted and pro- liferate in their new homes Of course proliferation of artificially introduced species may on the one hand be beneficial and what is desired On the other hand as H think many of us are aware it may not be desired And the latter brings us to another kind of use that can be made of zooarchaeological data when wildlife management is the issue

Identifying exotic animals

Recalling the TUCNs guidelines for reintroductions if a particular form of organism (ecotype subspecies or species) is exotic to a park or wildlife preserve it map be necessary to remove it from the area in order to preserve the natural status of the ecosystem and the biota of the area Thus we must have a solid definition of exotic ipecieThe United Stttes National Park Service (NPS) defines an exotic species as one that occurs in a given place ts a result of direct or indirect deliberate or accidental actions by humans (not including de- liberate reintroductions) (Hester 1991 127) In contrast native species are those which presently occur or once did occur prior to sonic human influence in a given place area or region as the result of ecological processes that operate and have operatcd without signifi cant direct or indirect deliberate or accidental alteration by humans (NPS documents cited in Lyman 1988a 21 see also Wagner et al 1995) Clearly then if wildlife managcrs are to manage native ecosystems variously referred to as pristine ecosystems or wilder- ness -by exclusion or removal of exotic species and reintroduction of artiticially extirpated native species then they must have that baseline list of native species In the United Stares that list is derived from the earliest historical documents for the area included within a national park (eg Houston and Schreiner 1995 Lgteopold et al 1963see also Hoerr 1993) The ethnocentricity of such a procedure is as might be imagined unavoidable from a przc tical standpoint More importantly historical documents arc sometimes incomplete and are sometimes simply inaccurate Ihose written documents can of course be sup plemented with another kind of document - the zooarchaeological record Here I men tion two examples of such supplementation

Bison (Bison bison) were artificially introduced to a portion of the state of Alaska that was to subsequently become Wrangell-St Elias National Park and Preserve (Peek et al 1987) By definition these introduced bison should be considered exotic and are so corl-sidered by the NPS (Houston and Schreiner 1995) However zooarchaeological evidence indicates that bison were present in the area until at least 500 years ago or slightly later (reviewed in Peek et al 1987) Historic records suggest bison wcre not prcsent when the first white men visited the area in the middle and late nineteenth ccntury but non- fossilized bison skulls were collected from the arca early in the twentieth century (Peek et al 1987) Given the paucity of late Holocene zooarchaeological data in thc area and the unknown nature of the relevant fossilization processes it is unclear if bison were present in Alaska between about 450 years ago and the carly twentieth century (McDonaald 1981)

The relevance of faunal analyl to wrldllfe rnanugerrzent 117

The NPS considers the bison of Wrangell-St Elias National Park to be exotic due to the lack of evidence that bison were present when the tirst white explorers passed through the area (Houston and Schreiner 1995) As well NPS biologists note that the introduced forrn of bison was of an incorrect - non-native -genetic stock (a distinct subspecies or ecotype) given current beliefs about bison taxonomy For this reason alone perhaps the extant bison of Wrangell-St Elias should be removed However the zooarchaeological record is so poorly known that one cannot conclusively demonstrate that native bison were locally extinct by the beginning of the twentieth century This in turn raises the possibility that extant bison are hybrids of native and introduced genetic stocks Whether or not this possibility is in fact reality is irrelevant What is critical here is that only more zooarchaeological and paleontological research will establish first the timing of local extirpation of native Alaskan bison populations and second if all Alaskan populations were extinct when transplanting occurred in the middle of the twentieth century Once the facts are determined we will have strong bases for making a decision regarding the ultimate fate of the bison presently living in Wrangell-St Elias National Park and Preserve

I am much more familiar - and intimately involved -with a virtually identical situation that exists in western Washington state Here mountain goats (Oreamnos urnericanlls) were introduced in the 1920s to the Olympic Mountains A decade later in 1938 these mountains were designated Olympic National Park The present position of the NPS is that the mountain goats are exotic and plans are in mid-1995 under way to eradicate them The plan is controversial (Anunsen and Anunsen 1993 Scheffer 1993) as is the evidence mustered to support it (compare Houston et al 119941 with Lyman 11991a 1995a 1995b1) This is not the place to review that evidence in detail (see references cited) It is relevant however to note the significance of zooarchaeological evidence for final resolution of the controversy

The historical evidence concerning the pre-1920s presence of mountain goats in the Olympic Mountains is ambiguous Some documents indicate goats were absent while others indicate goats were present (Lyman 1994a 1995b) In fact the only way to demonstrate that mountain goats were present or absent from the Olympic Mountains prior to their 1920s introduction requires paleontological andor zooarchaeological data At present there are no paleontological data of any kind for the area and the zooarchaeological record is woefully inadequate and thus cannot serve as a basis for a management decision (Lyman 1995a) Park biologists note that the purpose of national parks in the United States is not to attempt to recreate late-Pleistocene or early Holocene biotas [The NPS] is primarily concerned with historic post-Columbian specres distri- butions (Houston and Schreiner 1995 207) The facts remain however the zooarchaeo- logical record is poorly known and the historic record is internally contradictory

From the perspective of dynamic biogeographic histories provided by zooarchaeology one thing is abundantly clear the definitions of native and exotic species used by the NPS to implement its policies are contradictory Exotic species occur in a given place as a result of actions by humans whereas native species are those which presently occur or once did occ~crprior to some human influence in a place as the result of natural ecological processes The italicized phrase is where the contradiction resides because it indicates that i f for example mountain goats occurred at any tirne in the past in the Olympic Mountains

118 R Lee Lyman

- or bison in Wrangell-St Elias - then by definition the extant individuals represcl~i a native species But the introduced species are also by definition exotic having been transplanted from northwestern Canada and southeastern Alaska Olympic Nationai Parks mountain goats are of a different genetic stock and thus are a form distinct fromi mountain goats that may have been in the Olympic Mountains 200 years ago Rccogr-iizing this contradiction some years ago Y argued that if the prehistoric remains of rnonirrtain goats should be found in the Olympic Mountains then the NPS should rethink policy issues (Lyman I9RWa 22) Some NPS biologists have only recently come to the same conclusion (Houston and Schreiner 1995) And given the discussions above of appropriata forms (ecotypes subspecies species) and the benefit of hindsight I would argue that the policies and definitions of key terms should also be reworded to reflect the reality of wildlife management practice (Anonymous 1992 OBrien and Mayr 1991) Thus given their perspective of time depth zooarchaeologists not only can contribute significant paleobiological information to wildlife management concerns but perhaps they can also help write workable policies (see Wagner et al 1995 for other examples) Paleobiological information provided by zooarchaelogical research may also help wildlife managers plals for the future as the final example demonstrates

Biological preserves and planning for the future

Thus far the examples of zooarchaeology applied to wildlife management concerns have focused on the use of information on prehistoric faunal dynamics to inform management decisions That same kind of information can also be used to make broad predictions about the future effects of certain kinds of decisions For instance as alluded to in the first example presented above the spatial boundaries of parks and wildlife preserves may Piavc significant implications for the long-term survival of populations found withirl those boundaries In an important contribution to the role of planning for the future on tllc trasis of the past Graham (1988) notes that the boundaries of many parks and preserves are defined based on modern climatic and environmental conditions However tkrose conditions as we all know change and the probability of significant change increases as the length of time considered increases Graham underscores the [act that much planning lo preserve terrestrial biological diversity operates under the assumption that biotas or communities of organisms tend to respond to environmental change as intact units The

paleobiological record indicates however that individual species respond to environ- mental changes by migrating in different directions at different rates during different times (Graham 1988 392) Such individualistic responses to environmental change mean that the biological preserves of today are artifacts of the time when they were identified and created Thus Graham (1988 393) argues that constructing future reserves as habitat islands without avenues of migration [corridors] would probably result in [extirpation of bounded populations of managed species]

Graham (1988) thus suggests that as in the example of the Great Basins mountaintop mammals migration corridors are critical to organism survival and that preserves must be located on the landscape in such a rnanner as to be conducive to inter-areal migration I suggest following my earlier discussion that knowing where ihose corridors were located

The relevance of faunal analjgtsis to wildlife management 119

in the past surely will help in locating future preserves And as Graham and Graham (1994) indicate detailed knowledge of the historical and ecological biogeography of a taxon could prove critical to reconstructing appropriate habitats for reintroduced populations of the taxon Such knowledge can often come only from the fossil record

Extinction -of populations of species of lineages - is a natural fact of life (eg Raup 1984) But ignorance resulted in the loss of the passenger pigeon (Ectopistes migratorius) from North America the Great Auk (Pingninns impennis) from the North Atlantic and the dodo (Raphus cuci~llatus) from Mauritius in the Indian Ocean Our often stumbling attempts at wildlife management may due to ignorance ultimately lead to other such losses (eg Graham 1988 Hess 1993 Kay 1994) Ihe international move to maintain biological diversity - not just the taxonomic richness of biotic communities but the particular structure of those communities - seems to be in the correct direction if we are to maintain some quality of life and wilderness Understanding the loss of diversity often demands zooarchaeological and related data (eg Culotta 1995 Steadman 1995) and will help us avoid future inadvertent and unintentional losses Diamonds (1984) Rosetta Stone of using knowledge of modern extinctions to understand prehistoric extinctions could and perhaps should be turned on its head

Discussion

Nearly eighty years ago archaeologist W J Wintemberg (1919) noted how zooarchaeo- logical research could contribute to zoological knowledge and indirectly wildlife management His comments were repeated and elaborated thirty years later by biologist R M Gilmore (1949) Their remarks have largely gone unheeded perhaps because like Legge (1978 129 130) many of us see our job as aiming to contribute to the sum of the archaeological knowledge and that bone data are archaeological data I agree with Legge but also with Wintemberg and Gilmore we should be more than just consumers of wildlife data (cf Will 1982)

The potential topics under the broad umbrella of wildlife management to which zooarchaeological research might contribute are numerous I have touched on only a fevv of the more readily identifiable ones above There are to be sure others of some significance For example the modern fisheries industry of the eastern Pacific Ocean has been disrupted several times in the recent past by major changes in atmospheric circulation patterns causing alteration of ocean currents These events - variously referred to as El Nifio or Southern Oscillation events -appear to have some periodicity (Rasmusson 1985) but the historic record is of limited utility in plotting the precise nature of that periodicity These events clearly influence much of the oceanic biota (eg Arnold and Tissot 1993 Trillmich and Limberger 1985) Given sufficient temporal resolution the zooarchaeologi- cal record thus may eventually provide the time depth necessary to clarify the periodicity and magnitude of El Nifio events and thus allow more accurate predictions of future events

Zooarchaeological research has also shown that humans have long had a hand in modifying ecosystems Based on his extensive research on the prehistoric extinctions of Pacific Island bird populations and species Steadman (1995 1130) concludes that humans

tilay find Yt difficult to manage scarce resources in a sustainable way That humans contributed to these extinctions seems clear (cg Weisler and Cargett 1993) and thus Steadmans conclusion has the proper ominous tone But zooarchaeological research cair also perhaps help us out of such situations The wapiti (Cervns elaphus) of Yellowstone National Park in the United States are an example Herc nearly 60000 wapiti are managed and to a significant degree protected from human predation However historic and zooarchaeological research indicates that this population is a function of modern wildlife management historic records virtually vithout exception fail to mention wapiti i l l

the area and zooarchaeological data indicate this large ungulate was very riirc there during the last several thousand years (Kay 1994) Clearly if the Yellowstone ecosSsten1 is to be managed and maintained as a pristine wilderness in perpetuity (a laudable but in reality unattainable goal see Wager et al 1995) things must change before the ecosystem is irreparably altered A virtually identical situation exists in Rocky Mountain National Park sotile 550 km southwest of Yellowstone (Hcss 1993) where again zooarchaeologicai and other data from prehistoric times might help rcsolve debates

The few examples of how zooarchaeological research can be applied to t i ioder~~ problems I have outlined are rather specific to a place andlor to a taxon I do not perccive that to bc a weakness of an applied zooarchaeology but rather as an intellectual challenge Broadening the scope of zooarchaeology in the manner in which 1 have discussed will not be easy but i t will be beneficial to our Suture not only from the perspective of contributing to ensuring the preservation of biological diversity for future generations but also as 1conclude below from the perspective of our discipline which tilight otherwise become increasingly perceived as the pursuit of esoteric knotvledge of little practical use

One might argue that the discussion here could be expanded by suggesting that the detection of animal residues such as blood and hair (eg Loy 1993) could also providr information on the taxonomic identity of species native to the locale where the tools were collected (eg Hyland et al 1990) I doubt however that such a study would convince wildlife managers or all zooarchaeologists that a species was native given that stone tools were often transported significant distances by their tiiakers (eg Meltzer 1995) 1 beiievc multiple instances of such residues from multiple sites would be necessary to build 21

strong argument A partial skeleton or selected isolated bones and teeth of the species in que5tion recovered from a s~ngle ~ t e uould probably be muclr more conblnclng (Lyman 1994b)

I have attempted here to describe the structure of an applied zooarchaeology This is not a plea to legitimize what some of us do for fun andlor a living Rather i t is an attempt to broaden the scope of zooarchaeological research 1 am all for research for the sake of research if there is a problem in need of solving or a question in need of an answer But B am also concerned with the future of zooarchaeology Too many of our new doctoral str~dcnts are finding that the job market is closed and too marly of us arc finding it increasingly difficult to secure funding for research Perhaps a developed applied zooarchaeology wil

The relevance o f faunal analysis t o wildlife rnanagement 121

provide a new job market and new sources of funding It seems self-evident to me that if a national park for example has a staff archaeologist who deals with the archaeological resources on park lands and has a staff biologist who deals with biological resources on park lands then a marriage of the two would benefit both However given the increasing specialization of both archaeology and biology a viable marriage may not be possible Ihc obvious alternative is a park staff zooarchaeologist who has a deep knowledge ancl understanding of both disciplines and has been trained to integrate the two

New sources of funding may also become available if we can convince the wildlife management community that because its decisions and policies are typically aimed at tht future knowing something of the past can result in wise decisions That knowledge will of course come with a price tag The cost we may encounter is that members of the wildlife management community -while they may well have a solid knowledge of the time depth of evolutionary processes- often tend to view biotas as static ever enduring entities That is biotas have no past or at least their history is insignificant because they are being managed for the future While we certainly cannot predict exactly when the next ice age or glacial period will begin we can I think argue convincingly that climates and environments will change and knowing this we can use the prehistoric record to test our predictions about how certain kinds of changes may affect biotas of the future And confirmed - or even rejected or falsified -predictions would be the strong selling point Federal land-managing agencies whose charge includes wildlife management might well pay a zooarchaeologist to help them make wise management decisions The other selling point is that a less desirable kind of price -ecosystem destruction and loss of biodiversity -might otherwise have to be paid

Acknowledgements

I thank Donald K Grayson Adrian Lister Michael J OBrien and Ken Ihornas for comments on earlier versions of the manuscript

Departtnent o f Anthropology University o f Missouri-Columbia

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The relevance of faunal anulysis to wildlife management 11 1

modern biostratigraphic analyses such as Cookes (1976) study of suid remains prompting redating of volcanic tuffs in Plio-Pleistocene hominid sites of Africa zooarchaeology has contributed much more to our understanding of the past than just our recreations of the interaction sphere that included people and the fauna with which they were in contact (Olsen and Olsen 1981 193)

In this paper I argue that zooarchaeologists are in a unique position to move beyond tht important anthropological and archaeological issues they seem to prefer and to broaden the scope of their inquiries to include issues that are today and will be in the future important to humanity Specifically I outline here what I perceive to be a little explored avenue of zooarchaeological research --an avenue that could provide not only new sources of funding for zooarchaeologists but new sources of jobs for them as well

Archaeology is generally taught as a part of anthropology in North America In North America anthropological research seeks not only to increase our understanding of bu~ also to better the human condition From the anthropology of the homeless AIDS (acquired immune deficiency syndrome) and domestic violence to the anthropology of criminal behaviour - wherein archaeological forensics or taphonomic research plays a major role - anthropology can and does make a major contribution to making the world what we hope will be a better place Archaeology in general and zooarchaeology in particular with the exception of forensic research may appear to have little to offer in this respect But because so many modern potential ecological problems facing humanity concern our impact on biological phenomena - destruction of the rain forest decreasing harvests of marine fishes and the like - zooarchaeology as well as paleoethnobotany clearly has a major role to play in the resolution of those problems

In the following I describe several instances of the significance of zooarchaeological (and paleontological) data for addressing and helping to resolve particular kinds of modern problems faced by those concerned with wildlife management Many nations have one or more forms of national park andlor game preserve Many of these spatial units are managed in such a manner as to protect i n perpetuity the biota included within their boundaries Sometimes attempts are made to enhance or modify the biotas of these parks and preserves so that they reflect what are variously referred to as pristine ecosystems wilderness or native (pre-industrial) or natural biotas Further many nations have various legal controls on hunting and trapping of animal species and legal controls over which species can be harvested and which are to be unmolested Managing wildlife sometimes involves supplementing resident populations with individual organisms transplanted from elsewhere or attempting to re-establish species in areas where native populations have become extirpated With its unique perspective on the time depth of ecosystems and biotas zooarchaeological research can be applied to ensure that wise wildlife management decisions are made

The kinds of wildlife management problems that zooarchaeological research might help resolve are many They are however often very specific to I particular geographical place andlor species The examples that follow are those with which I am most familiar and thus centre on North America Examples from other continents and regions surely exist None the less I have attempted to select examples that illustrate a broad diversity of wildlife management concerns and how particular kinds of zooarchaeological research might help resolve them

112 R Lee Lymar~

Populations threatened with extirpation

One of the major issues facing wildlife managers working in national parlts kind wiidlifc preserves involves ensuring the wildlife species undcr their carc survivc in perpcinit) Thus wildlife managers attempt lo identify measures thai can be taken ao ensure i81r1t extirpation of species resident on park or preserve lands does not occur ltdne virYal~ic presently being debated in North Amcrican wildlifc management that weighs heavily or1

this issue concerns what is known as SLOSS Simply put should i park or preserve corrsisi of Single Large Or Several Srriall pieces of landscape (see for example Quinn a n d Harrison [I9881and references therein) Thus far the debate his employed daia o n t h e modern distribution of taxa and has typically inferred that tensporal processes havc C I U S C ~

these spatial patterns Zooarchaeological data brought to bear con this dcbate can 111akc 111c temporal inference empirical

Grayson (1991) reviewed the biogeogr-aph~c history CPC smali mamrnals in ihr Gteat Basin physiographic province of western North America and then used zooarchacologicaI and paleontological data to test the implications of a model first proposed by bingiiogra- phei-JFI Brown (197 1 1978) Hm simplest terms Brown rnodel suggests that the i-irotiern distribution of subalpine and alpinc s~nall mammals (c pika [Ocllororrrprinreyial golden mantled ground squirrel [Sper~nophilus llrterulis] ermine [Muslelcl ertvaanriij) oil

Great Basin mountaintops was the result of Pleistocene colonization of those rnountairl followed by Holocene extinction of intermediate lowland populations arid by difcrcrstir extinction of various mountaintop populatioras Qrag~scgtn (1991 372) found that Browsrs model has impressive paleontological support and T note that much of that suppor~ derives from zooarchaeological research

Grayson (1991 373) noted that confirmation of Browns model with thc fossii rccerrci has clear implications for the management of high aleie~ule environr~~erats in the Circlt Basin He was referring to the fact that nnany populations of small mammals iur ahis liaa

are today isolated on mountaintops Basically following thc theory of island biogeography (MacArthur and Wilson 1967) as modified over the years (eg Brown 1986) b3rayrtsrii study underscored -- using zooarchaeological data - the fact that such small gcncticaliy isolated populations of species tend to I-~e more prone inr extinction than larger aa-lciir non-isolated populations (see also Korn 1994) Larger populations provide suificicrii genetic variation Lo ensure survival of a species and larger areas allow migration betwecn population node because corridors bctween thcni are shorter Either isolation of tliose nodes rrom one another or restriction of those node to small geographic areas - s u c l ~as mountaintops or both increases the probability that one or more of thcpse na~dcs wili-

cease to exist Thus the biogeographic histories of alpine small rnanirnals of the Griit

Basin indicate that neithcr alternative of SEOSS is u~orkablc herc We cannot enlarge tile

insular alpine and subalpine habitats found on Great Basin mountaintops bounalarie a i thosc habitat patchcs arc climatically dictateai And these habitat patches are not only small they are also multiple in number Ensuring that thesc patches are not artiliciirlly disrupted in the future would seem to be the only way to guarantee survivii of tl~eir

included small mammals Migration corridors betwecn the habitat patches canraoi hi artificially constructeci or maintained If such corridors could be built where shoa~ldthe

The relevance of faunal analysis to wildlife management 113

be located The fossil record might help us determine where these corridors existed in the past as can be illustrated with another example

The pygmy rabbit (Brachylagi~s idahoensis) is a diminutive leporid with a modern range restricted to two areas of the western United States One area is relatively large and encompasses portions of four states northern Nevada western Utah southern Idaho and southeastern Oregon The other historically documented area where this species occurs is the central portion of eastern Washington state Populations in the two areas are presently isolated from one another The wildlife management coricern here is that the eastern Washington population has over the past forty years shrunk to an alarmingly small size prompting the Washington Wildlife Commission to list this population as threatened and to suggest that i t be listed as endangered (McAllister and Allen 1993) and to develop a management pan with the aim of ensuring survival of the population (McAllister and Allen 1994) The zooarchaeological record of this species provides insight to why this population is threatened with extirpation and how i t might be preserved

The zooarchaeological record for pygmy rabbits indicates this species probably occupied a wider range in central Washington during the Holocene than i t presently does Remains of this species have now been recovered from eleven archaeological and two paleontological sites in the area (updated from Lyman 1991) Some of these are extrali~nitalrecords that is some remains have been recovered from geographic locations where pygmy rabbits have not been historically documented to occur In brief the fossil (botanical and faunal) record suggests that when big sagebrush (Artemisia tridentata) was more widespread in central Washington -- during the middle Holocene climatic interval known as the Altithermal - pygmy rabbits were also more widespread When the Altithermal ended about 4500 years ago the range of sagebrush shrank and so too did the range of pygmy rabbits Then during the late nineteenth and early twentieth centuries land was cleared of sagebrush for agricultural purposes This prompted a second diminution of sagebrush and pygmy rabbit range

Ecological studies of pygmy rabbits indicate that this leporid is dependent on big sagebrush for food and for shelter from predation (Green and Flinders 1980a 1980b) The zooarchaeological record in conjunction with paleobotanical data such as that from pollen records confirms this as both species appear to have responded to climatic change in like manner Washington Department of Wildlife biologists have made use of both modern and zooarchaeological data in their planning efforts (McAllister and Allen 1993 1994) The fossil record suggests that the prehistoric source of Washingtons pygmy rabbits resides in the area of the larger extant population to the south Late Pleistocene fossils of pygmy rabbits recovered from a paleontological site between the two areas (Rensberger and Barnosky 1993) tend to confirm this suggestion While i t would probably be impractical to develop a migration corridor between the two populations today it would also be impossible given the climatic history of the area The probable corridor between the two today is characterized by vegetation habitats that are not conducive to tht survival of pygmy rabbits The zooarchaeological record does indicate however that the extant population might be supplemented by individuals transplanted from southeasterri Oregon and that the maintenance of habitats dominated by big sagebrush is critical to the survival of the species

114 R Lee Lyman

When managing small isolated populations we may not have many options other than to preserve those populations as they presently exist Yn some cases management efforts have failed and populations have been extirpated When this occurs one management alternative is to transplant individuals from other populations to the vacated rangc in an attempt to re-establish a population Zooarchaeological research can also plav an important role in these situations

Re-establishment of extirpated populations

Eighteenth- and nineteenth-century conimercial exploitation of sea otters (Enhydru lirtris) along the western coast of North America ultimately led to the extirpation of this species from much of its range (Kenyon 1969 Ogden 1993 Roest 1973) Attempts to re-establish populations along the middle latitudes of that coast took place in the 1970s and thus far have met with less than perfect success (Jameson et al 1982 1986 MacAskie 1987) The typically expressed reason for the limited success of these efforts is that ecosystems to which sea otters have been transplanted have heen modified too exter~sively by modern human industry to support them Y wonder i f however part of the reason might resicle in a factor that only zooarchaeological research car measure

Sea otter skulls collectetl from much of the once occupied eastern Pacific range of sea otters display various character gradients from the Aleutian Islands to southern California Some of these characters have been used to argue for or against the existence oi multiple subspecies of sea otter (Davis and Lidicker 1975 Roest 1973 1979 Scheffer and Wilke 1950 Wilson et al 1991) Final resolution of this debate will be critical to futtlrc wildlife management activities first because national legislation in the United States concerns species subspecies and populations (OWrien and Mayr 1901) and also beca~ase of conflict between one population (a possibly distinct subspecies) of sea otters and the modern fisheries industry (Wendell et al 1986) lhe significant aspect to all of this Is the transplanting of sea otters ILegally only forms (ecotypes or subspecies) native to ar arcs should be introduced there during efforts to re-establish populations This also m~aites biological sense as i t is likely that those native forms unique to an area are the product of millennia of evolution

The International Union for the Conservation of Nature and Natural Resources PUCN (recently renamed the World Conservation Un~on) has drafted guidelines Cor such -

reintroductions These read in part

An assessment should be made of the taxonomic status of individuals to ba re-introduced They should be of the same taxonomic unit (and ideally closely related genetically) as those which were extirpated An investigation of historical information about the loss and fate of individuals from the re-introduction area as well a mmolecuiar genetic studies should be undertaken in case of doubt Release stock idealiy should be closely-related genetically to the original native stock

(Anonymous 1992 2-31

Clearly a search of not only historic but also prehistoric information nnay be necessary The rapidly developing study of ancient DNA extracted from prehistoric skeletal tissues

The relevance of faunal anuljlsis to wildlife matlugement 115

(eg Richards et al 1993 1995 and references therein) seems to have great potential for contending with IUCN guidelines The technique is applicable to curated as wtll as newly acquired archaeological specimens It has however not yet been applied to Oregon coast sea otter remains

Given the lack of genetic material from eighteenth- and early nineteenth-century sea otters on the Oregon coast the problem becomes one of establishing the age of the character gradients displayed by eastern Pacific rim sea otters (see Wilson et al 1991 for a recent summary) and determining if those gradients have shifted spatially through time IIf those gradients have a prehistoric distribution in time andlor space similar to that documented using historic specimens a potential problem is raised If a particular form -represented by a particular character state -- of sea otter was present prehistorically in one location within the range and another form was present at another location then transplanting individuals from the former to the latter may be doomed from the start In such a case failure to re-establish a population could result on the one hand because the transplanted form is not well adapted to its new environment the extirpated form on the other hand is adapted to the environment of that location hence its difference in form

Several years ago I examined prehistoric sea otter remains recovered from archaeologi- cal sites on the coast of the state of Oregon - the middle latitudes of the historically documented range of sea otters (Lyman 1988b) The size of sea otter teeth recovered from those sites are more or less intermediate to those from Alaska (large) and those from southern California (small) but they also vary somewhat from this simple pattern Prehistoric Oregon sea otters match modern Alaskan sea otters in some characters in others they match modern California sea otters Precise details are unnecessary to the point I wish to make here which is Were the sea otters captured in Alaska and translocated to the Oregon coast in the 1970s of the proper phenotype (or genotype) Zooarchaeological evidence suggests they were not If this is a correct interpretation then perhaps those transplanted sea otters were doomed from the moment they were captured Whether or not my suggestion proves to be true or false and it is unclear which way the cards might fall given that we still have much to learn about this species (VanBlaricom arid Estes 1988) the significance of zooarchaeological data should be clear

Not only can zooarchaeological data inform us as to which form of a species might be the most appropriate one to transplant to an area such data have the potential to indicate the potential success of such transplant efforts Even if the appropriate form to transplant is clear this does not guarantee that the introduced form will survive in its new home This is precisely the argument offered by biologists working with sea otters on the Oregon coast The data substantiaiing their argument can be supplemented with zooarchaeological data An example of this is provided by the California condor (Gymnogyps cnlifornianus) of western North America a species presently near extinction (Emslie 1987) Zooarchaeo- logical and paleontological data suggest this species enjoyed a wider range during the late Pleistocene but that range shrank considerably with the onset of the postglacial Holocene epoch (Emslie 1987 Simons 1983) Those data also suggest that this species was a carrion feeder and that the terminal Pleistocene extinction of many genera of large mammals probably contributed to the decline in frequency and range of the California condor (Emslie 1987) Finally paleoecological data indicate attempts to reintroduce this species to portions of its prehistoric range would probably fail because of major differences

116 R Lee Lytriurz

between late Holocene floral and faunal communities and late Pleistocene commaanities (Emslie 1987)

In the absence of historic data or even with it (see below) zooarchaeological infor- mation may be critical to ensuring that the correct forms of animals are transplanted The wrong forms may be i l l suited to their new habitats or they may be well adapted and pro- liferate in their new homes Of course proliferation of artificially introduced species may on the one hand be beneficial and what is desired On the other hand as H think many of us are aware it may not be desired And the latter brings us to another kind of use that can be made of zooarchaeological data when wildlife management is the issue

Identifying exotic animals

Recalling the TUCNs guidelines for reintroductions if a particular form of organism (ecotype subspecies or species) is exotic to a park or wildlife preserve it map be necessary to remove it from the area in order to preserve the natural status of the ecosystem and the biota of the area Thus we must have a solid definition of exotic ipecieThe United Stttes National Park Service (NPS) defines an exotic species as one that occurs in a given place ts a result of direct or indirect deliberate or accidental actions by humans (not including de- liberate reintroductions) (Hester 1991 127) In contrast native species are those which presently occur or once did occur prior to sonic human influence in a given place area or region as the result of ecological processes that operate and have operatcd without signifi cant direct or indirect deliberate or accidental alteration by humans (NPS documents cited in Lyman 1988a 21 see also Wagner et al 1995) Clearly then if wildlife managcrs are to manage native ecosystems variously referred to as pristine ecosystems or wilder- ness -by exclusion or removal of exotic species and reintroduction of artiticially extirpated native species then they must have that baseline list of native species In the United Stares that list is derived from the earliest historical documents for the area included within a national park (eg Houston and Schreiner 1995 Lgteopold et al 1963see also Hoerr 1993) The ethnocentricity of such a procedure is as might be imagined unavoidable from a przc tical standpoint More importantly historical documents arc sometimes incomplete and are sometimes simply inaccurate Ihose written documents can of course be sup plemented with another kind of document - the zooarchaeological record Here I men tion two examples of such supplementation

Bison (Bison bison) were artificially introduced to a portion of the state of Alaska that was to subsequently become Wrangell-St Elias National Park and Preserve (Peek et al 1987) By definition these introduced bison should be considered exotic and are so corl-sidered by the NPS (Houston and Schreiner 1995) However zooarchaeological evidence indicates that bison were present in the area until at least 500 years ago or slightly later (reviewed in Peek et al 1987) Historic records suggest bison wcre not prcsent when the first white men visited the area in the middle and late nineteenth ccntury but non- fossilized bison skulls were collected from the arca early in the twentieth century (Peek et al 1987) Given the paucity of late Holocene zooarchaeological data in thc area and the unknown nature of the relevant fossilization processes it is unclear if bison were present in Alaska between about 450 years ago and the carly twentieth century (McDonaald 1981)

The relevance of faunal analyl to wrldllfe rnanugerrzent 117

The NPS considers the bison of Wrangell-St Elias National Park to be exotic due to the lack of evidence that bison were present when the tirst white explorers passed through the area (Houston and Schreiner 1995) As well NPS biologists note that the introduced forrn of bison was of an incorrect - non-native -genetic stock (a distinct subspecies or ecotype) given current beliefs about bison taxonomy For this reason alone perhaps the extant bison of Wrangell-St Elias should be removed However the zooarchaeological record is so poorly known that one cannot conclusively demonstrate that native bison were locally extinct by the beginning of the twentieth century This in turn raises the possibility that extant bison are hybrids of native and introduced genetic stocks Whether or not this possibility is in fact reality is irrelevant What is critical here is that only more zooarchaeological and paleontological research will establish first the timing of local extirpation of native Alaskan bison populations and second if all Alaskan populations were extinct when transplanting occurred in the middle of the twentieth century Once the facts are determined we will have strong bases for making a decision regarding the ultimate fate of the bison presently living in Wrangell-St Elias National Park and Preserve

I am much more familiar - and intimately involved -with a virtually identical situation that exists in western Washington state Here mountain goats (Oreamnos urnericanlls) were introduced in the 1920s to the Olympic Mountains A decade later in 1938 these mountains were designated Olympic National Park The present position of the NPS is that the mountain goats are exotic and plans are in mid-1995 under way to eradicate them The plan is controversial (Anunsen and Anunsen 1993 Scheffer 1993) as is the evidence mustered to support it (compare Houston et al 119941 with Lyman 11991a 1995a 1995b1) This is not the place to review that evidence in detail (see references cited) It is relevant however to note the significance of zooarchaeological evidence for final resolution of the controversy

The historical evidence concerning the pre-1920s presence of mountain goats in the Olympic Mountains is ambiguous Some documents indicate goats were absent while others indicate goats were present (Lyman 1994a 1995b) In fact the only way to demonstrate that mountain goats were present or absent from the Olympic Mountains prior to their 1920s introduction requires paleontological andor zooarchaeological data At present there are no paleontological data of any kind for the area and the zooarchaeological record is woefully inadequate and thus cannot serve as a basis for a management decision (Lyman 1995a) Park biologists note that the purpose of national parks in the United States is not to attempt to recreate late-Pleistocene or early Holocene biotas [The NPS] is primarily concerned with historic post-Columbian specres distri- butions (Houston and Schreiner 1995 207) The facts remain however the zooarchaeo- logical record is poorly known and the historic record is internally contradictory

From the perspective of dynamic biogeographic histories provided by zooarchaeology one thing is abundantly clear the definitions of native and exotic species used by the NPS to implement its policies are contradictory Exotic species occur in a given place as a result of actions by humans whereas native species are those which presently occur or once did occ~crprior to some human influence in a place as the result of natural ecological processes The italicized phrase is where the contradiction resides because it indicates that i f for example mountain goats occurred at any tirne in the past in the Olympic Mountains

118 R Lee Lyman

- or bison in Wrangell-St Elias - then by definition the extant individuals represcl~i a native species But the introduced species are also by definition exotic having been transplanted from northwestern Canada and southeastern Alaska Olympic Nationai Parks mountain goats are of a different genetic stock and thus are a form distinct fromi mountain goats that may have been in the Olympic Mountains 200 years ago Rccogr-iizing this contradiction some years ago Y argued that if the prehistoric remains of rnonirrtain goats should be found in the Olympic Mountains then the NPS should rethink policy issues (Lyman I9RWa 22) Some NPS biologists have only recently come to the same conclusion (Houston and Schreiner 1995) And given the discussions above of appropriata forms (ecotypes subspecies species) and the benefit of hindsight I would argue that the policies and definitions of key terms should also be reworded to reflect the reality of wildlife management practice (Anonymous 1992 OBrien and Mayr 1991) Thus given their perspective of time depth zooarchaeologists not only can contribute significant paleobiological information to wildlife management concerns but perhaps they can also help write workable policies (see Wagner et al 1995 for other examples) Paleobiological information provided by zooarchaelogical research may also help wildlife managers plals for the future as the final example demonstrates

Biological preserves and planning for the future

Thus far the examples of zooarchaeology applied to wildlife management concerns have focused on the use of information on prehistoric faunal dynamics to inform management decisions That same kind of information can also be used to make broad predictions about the future effects of certain kinds of decisions For instance as alluded to in the first example presented above the spatial boundaries of parks and wildlife preserves may Piavc significant implications for the long-term survival of populations found withirl those boundaries In an important contribution to the role of planning for the future on tllc trasis of the past Graham (1988) notes that the boundaries of many parks and preserves are defined based on modern climatic and environmental conditions However tkrose conditions as we all know change and the probability of significant change increases as the length of time considered increases Graham underscores the [act that much planning lo preserve terrestrial biological diversity operates under the assumption that biotas or communities of organisms tend to respond to environmental change as intact units The

paleobiological record indicates however that individual species respond to environ- mental changes by migrating in different directions at different rates during different times (Graham 1988 392) Such individualistic responses to environmental change mean that the biological preserves of today are artifacts of the time when they were identified and created Thus Graham (1988 393) argues that constructing future reserves as habitat islands without avenues of migration [corridors] would probably result in [extirpation of bounded populations of managed species]

Graham (1988) thus suggests that as in the example of the Great Basins mountaintop mammals migration corridors are critical to organism survival and that preserves must be located on the landscape in such a rnanner as to be conducive to inter-areal migration I suggest following my earlier discussion that knowing where ihose corridors were located

The relevance of faunal analjgtsis to wildlife management 119

in the past surely will help in locating future preserves And as Graham and Graham (1994) indicate detailed knowledge of the historical and ecological biogeography of a taxon could prove critical to reconstructing appropriate habitats for reintroduced populations of the taxon Such knowledge can often come only from the fossil record

Extinction -of populations of species of lineages - is a natural fact of life (eg Raup 1984) But ignorance resulted in the loss of the passenger pigeon (Ectopistes migratorius) from North America the Great Auk (Pingninns impennis) from the North Atlantic and the dodo (Raphus cuci~llatus) from Mauritius in the Indian Ocean Our often stumbling attempts at wildlife management may due to ignorance ultimately lead to other such losses (eg Graham 1988 Hess 1993 Kay 1994) Ihe international move to maintain biological diversity - not just the taxonomic richness of biotic communities but the particular structure of those communities - seems to be in the correct direction if we are to maintain some quality of life and wilderness Understanding the loss of diversity often demands zooarchaeological and related data (eg Culotta 1995 Steadman 1995) and will help us avoid future inadvertent and unintentional losses Diamonds (1984) Rosetta Stone of using knowledge of modern extinctions to understand prehistoric extinctions could and perhaps should be turned on its head

Discussion

Nearly eighty years ago archaeologist W J Wintemberg (1919) noted how zooarchaeo- logical research could contribute to zoological knowledge and indirectly wildlife management His comments were repeated and elaborated thirty years later by biologist R M Gilmore (1949) Their remarks have largely gone unheeded perhaps because like Legge (1978 129 130) many of us see our job as aiming to contribute to the sum of the archaeological knowledge and that bone data are archaeological data I agree with Legge but also with Wintemberg and Gilmore we should be more than just consumers of wildlife data (cf Will 1982)

The potential topics under the broad umbrella of wildlife management to which zooarchaeological research might contribute are numerous I have touched on only a fevv of the more readily identifiable ones above There are to be sure others of some significance For example the modern fisheries industry of the eastern Pacific Ocean has been disrupted several times in the recent past by major changes in atmospheric circulation patterns causing alteration of ocean currents These events - variously referred to as El Nifio or Southern Oscillation events -appear to have some periodicity (Rasmusson 1985) but the historic record is of limited utility in plotting the precise nature of that periodicity These events clearly influence much of the oceanic biota (eg Arnold and Tissot 1993 Trillmich and Limberger 1985) Given sufficient temporal resolution the zooarchaeologi- cal record thus may eventually provide the time depth necessary to clarify the periodicity and magnitude of El Nifio events and thus allow more accurate predictions of future events

Zooarchaeological research has also shown that humans have long had a hand in modifying ecosystems Based on his extensive research on the prehistoric extinctions of Pacific Island bird populations and species Steadman (1995 1130) concludes that humans

tilay find Yt difficult to manage scarce resources in a sustainable way That humans contributed to these extinctions seems clear (cg Weisler and Cargett 1993) and thus Steadmans conclusion has the proper ominous tone But zooarchaeological research cair also perhaps help us out of such situations The wapiti (Cervns elaphus) of Yellowstone National Park in the United States are an example Herc nearly 60000 wapiti are managed and to a significant degree protected from human predation However historic and zooarchaeological research indicates that this population is a function of modern wildlife management historic records virtually vithout exception fail to mention wapiti i l l

the area and zooarchaeological data indicate this large ungulate was very riirc there during the last several thousand years (Kay 1994) Clearly if the Yellowstone ecosSsten1 is to be managed and maintained as a pristine wilderness in perpetuity (a laudable but in reality unattainable goal see Wager et al 1995) things must change before the ecosystem is irreparably altered A virtually identical situation exists in Rocky Mountain National Park sotile 550 km southwest of Yellowstone (Hcss 1993) where again zooarchaeologicai and other data from prehistoric times might help rcsolve debates

The few examples of how zooarchaeological research can be applied to t i ioder~~ problems I have outlined are rather specific to a place andlor to a taxon I do not perccive that to bc a weakness of an applied zooarchaeology but rather as an intellectual challenge Broadening the scope of zooarchaeology in the manner in which 1 have discussed will not be easy but i t will be beneficial to our Suture not only from the perspective of contributing to ensuring the preservation of biological diversity for future generations but also as 1conclude below from the perspective of our discipline which tilight otherwise become increasingly perceived as the pursuit of esoteric knotvledge of little practical use

One might argue that the discussion here could be expanded by suggesting that the detection of animal residues such as blood and hair (eg Loy 1993) could also providr information on the taxonomic identity of species native to the locale where the tools were collected (eg Hyland et al 1990) I doubt however that such a study would convince wildlife managers or all zooarchaeologists that a species was native given that stone tools were often transported significant distances by their tiiakers (eg Meltzer 1995) 1 beiievc multiple instances of such residues from multiple sites would be necessary to build 21

strong argument A partial skeleton or selected isolated bones and teeth of the species in que5tion recovered from a s~ngle ~ t e uould probably be muclr more conblnclng (Lyman 1994b)

I have attempted here to describe the structure of an applied zooarchaeology This is not a plea to legitimize what some of us do for fun andlor a living Rather i t is an attempt to broaden the scope of zooarchaeological research 1 am all for research for the sake of research if there is a problem in need of solving or a question in need of an answer But B am also concerned with the future of zooarchaeology Too many of our new doctoral str~dcnts are finding that the job market is closed and too marly of us arc finding it increasingly difficult to secure funding for research Perhaps a developed applied zooarchaeology wil

The relevance o f faunal analysis t o wildlife rnanagement 121

provide a new job market and new sources of funding It seems self-evident to me that if a national park for example has a staff archaeologist who deals with the archaeological resources on park lands and has a staff biologist who deals with biological resources on park lands then a marriage of the two would benefit both However given the increasing specialization of both archaeology and biology a viable marriage may not be possible Ihc obvious alternative is a park staff zooarchaeologist who has a deep knowledge ancl understanding of both disciplines and has been trained to integrate the two

New sources of funding may also become available if we can convince the wildlife management community that because its decisions and policies are typically aimed at tht future knowing something of the past can result in wise decisions That knowledge will of course come with a price tag The cost we may encounter is that members of the wildlife management community -while they may well have a solid knowledge of the time depth of evolutionary processes- often tend to view biotas as static ever enduring entities That is biotas have no past or at least their history is insignificant because they are being managed for the future While we certainly cannot predict exactly when the next ice age or glacial period will begin we can I think argue convincingly that climates and environments will change and knowing this we can use the prehistoric record to test our predictions about how certain kinds of changes may affect biotas of the future And confirmed - or even rejected or falsified -predictions would be the strong selling point Federal land-managing agencies whose charge includes wildlife management might well pay a zooarchaeologist to help them make wise management decisions The other selling point is that a less desirable kind of price -ecosystem destruction and loss of biodiversity -might otherwise have to be paid

Acknowledgements

I thank Donald K Grayson Adrian Lister Michael J OBrien and Ken Ihornas for comments on earlier versions of the manuscript

Departtnent o f Anthropology University o f Missouri-Columbia

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112 R Lee Lymar~

Populations threatened with extirpation

One of the major issues facing wildlife managers working in national parlts kind wiidlifc preserves involves ensuring the wildlife species undcr their carc survivc in perpcinit) Thus wildlife managers attempt lo identify measures thai can be taken ao ensure i81r1t extirpation of species resident on park or preserve lands does not occur ltdne virYal~ic presently being debated in North Amcrican wildlifc management that weighs heavily or1

this issue concerns what is known as SLOSS Simply put should i park or preserve corrsisi of Single Large Or Several Srriall pieces of landscape (see for example Quinn a n d Harrison [I9881and references therein) Thus far the debate his employed daia o n t h e modern distribution of taxa and has typically inferred that tensporal processes havc C I U S C ~

these spatial patterns Zooarchaeological data brought to bear con this dcbate can 111akc 111c temporal inference empirical

Grayson (1991) reviewed the biogeogr-aph~c history CPC smali mamrnals in ihr Gteat Basin physiographic province of western North America and then used zooarchacologicaI and paleontological data to test the implications of a model first proposed by bingiiogra- phei-JFI Brown (197 1 1978) Hm simplest terms Brown rnodel suggests that the i-irotiern distribution of subalpine and alpinc s~nall mammals (c pika [Ocllororrrprinreyial golden mantled ground squirrel [Sper~nophilus llrterulis] ermine [Muslelcl ertvaanriij) oil

Great Basin mountaintops was the result of Pleistocene colonization of those rnountairl followed by Holocene extinction of intermediate lowland populations arid by difcrcrstir extinction of various mountaintop populatioras Qrag~scgtn (1991 372) found that Browsrs model has impressive paleontological support and T note that much of that suppor~ derives from zooarchaeological research

Grayson (1991 373) noted that confirmation of Browns model with thc fossii rccerrci has clear implications for the management of high aleie~ule environr~~erats in the Circlt Basin He was referring to the fact that nnany populations of small mammals iur ahis liaa

are today isolated on mountaintops Basically following thc theory of island biogeography (MacArthur and Wilson 1967) as modified over the years (eg Brown 1986) b3rayrtsrii study underscored -- using zooarchaeological data - the fact that such small gcncticaliy isolated populations of species tend to I-~e more prone inr extinction than larger aa-lciir non-isolated populations (see also Korn 1994) Larger populations provide suificicrii genetic variation Lo ensure survival of a species and larger areas allow migration betwecn population node because corridors bctween thcni are shorter Either isolation of tliose nodes rrom one another or restriction of those node to small geographic areas - s u c l ~as mountaintops or both increases the probability that one or more of thcpse na~dcs wili-

cease to exist Thus the biogeographic histories of alpine small rnanirnals of the Griit

Basin indicate that neithcr alternative of SEOSS is u~orkablc herc We cannot enlarge tile

insular alpine and subalpine habitats found on Great Basin mountaintops bounalarie a i thosc habitat patchcs arc climatically dictateai And these habitat patches are not only small they are also multiple in number Ensuring that thesc patches are not artiliciirlly disrupted in the future would seem to be the only way to guarantee survivii of tl~eir

included small mammals Migration corridors betwecn the habitat patches canraoi hi artificially constructeci or maintained If such corridors could be built where shoa~ldthe

The relevance of faunal analysis to wildlife management 113

be located The fossil record might help us determine where these corridors existed in the past as can be illustrated with another example

The pygmy rabbit (Brachylagi~s idahoensis) is a diminutive leporid with a modern range restricted to two areas of the western United States One area is relatively large and encompasses portions of four states northern Nevada western Utah southern Idaho and southeastern Oregon The other historically documented area where this species occurs is the central portion of eastern Washington state Populations in the two areas are presently isolated from one another The wildlife management coricern here is that the eastern Washington population has over the past forty years shrunk to an alarmingly small size prompting the Washington Wildlife Commission to list this population as threatened and to suggest that i t be listed as endangered (McAllister and Allen 1993) and to develop a management pan with the aim of ensuring survival of the population (McAllister and Allen 1994) The zooarchaeological record of this species provides insight to why this population is threatened with extirpation and how i t might be preserved

The zooarchaeological record for pygmy rabbits indicates this species probably occupied a wider range in central Washington during the Holocene than i t presently does Remains of this species have now been recovered from eleven archaeological and two paleontological sites in the area (updated from Lyman 1991) Some of these are extrali~nitalrecords that is some remains have been recovered from geographic locations where pygmy rabbits have not been historically documented to occur In brief the fossil (botanical and faunal) record suggests that when big sagebrush (Artemisia tridentata) was more widespread in central Washington -- during the middle Holocene climatic interval known as the Altithermal - pygmy rabbits were also more widespread When the Altithermal ended about 4500 years ago the range of sagebrush shrank and so too did the range of pygmy rabbits Then during the late nineteenth and early twentieth centuries land was cleared of sagebrush for agricultural purposes This prompted a second diminution of sagebrush and pygmy rabbit range

Ecological studies of pygmy rabbits indicate that this leporid is dependent on big sagebrush for food and for shelter from predation (Green and Flinders 1980a 1980b) The zooarchaeological record in conjunction with paleobotanical data such as that from pollen records confirms this as both species appear to have responded to climatic change in like manner Washington Department of Wildlife biologists have made use of both modern and zooarchaeological data in their planning efforts (McAllister and Allen 1993 1994) The fossil record suggests that the prehistoric source of Washingtons pygmy rabbits resides in the area of the larger extant population to the south Late Pleistocene fossils of pygmy rabbits recovered from a paleontological site between the two areas (Rensberger and Barnosky 1993) tend to confirm this suggestion While i t would probably be impractical to develop a migration corridor between the two populations today it would also be impossible given the climatic history of the area The probable corridor between the two today is characterized by vegetation habitats that are not conducive to tht survival of pygmy rabbits The zooarchaeological record does indicate however that the extant population might be supplemented by individuals transplanted from southeasterri Oregon and that the maintenance of habitats dominated by big sagebrush is critical to the survival of the species

114 R Lee Lyman

When managing small isolated populations we may not have many options other than to preserve those populations as they presently exist Yn some cases management efforts have failed and populations have been extirpated When this occurs one management alternative is to transplant individuals from other populations to the vacated rangc in an attempt to re-establish a population Zooarchaeological research can also plav an important role in these situations

Re-establishment of extirpated populations

Eighteenth- and nineteenth-century conimercial exploitation of sea otters (Enhydru lirtris) along the western coast of North America ultimately led to the extirpation of this species from much of its range (Kenyon 1969 Ogden 1993 Roest 1973) Attempts to re-establish populations along the middle latitudes of that coast took place in the 1970s and thus far have met with less than perfect success (Jameson et al 1982 1986 MacAskie 1987) The typically expressed reason for the limited success of these efforts is that ecosystems to which sea otters have been transplanted have heen modified too exter~sively by modern human industry to support them Y wonder i f however part of the reason might resicle in a factor that only zooarchaeological research car measure

Sea otter skulls collectetl from much of the once occupied eastern Pacific range of sea otters display various character gradients from the Aleutian Islands to southern California Some of these characters have been used to argue for or against the existence oi multiple subspecies of sea otter (Davis and Lidicker 1975 Roest 1973 1979 Scheffer and Wilke 1950 Wilson et al 1991) Final resolution of this debate will be critical to futtlrc wildlife management activities first because national legislation in the United States concerns species subspecies and populations (OWrien and Mayr 1901) and also beca~ase of conflict between one population (a possibly distinct subspecies) of sea otters and the modern fisheries industry (Wendell et al 1986) lhe significant aspect to all of this Is the transplanting of sea otters ILegally only forms (ecotypes or subspecies) native to ar arcs should be introduced there during efforts to re-establish populations This also m~aites biological sense as i t is likely that those native forms unique to an area are the product of millennia of evolution

The International Union for the Conservation of Nature and Natural Resources PUCN (recently renamed the World Conservation Un~on) has drafted guidelines Cor such -

reintroductions These read in part

An assessment should be made of the taxonomic status of individuals to ba re-introduced They should be of the same taxonomic unit (and ideally closely related genetically) as those which were extirpated An investigation of historical information about the loss and fate of individuals from the re-introduction area as well a mmolecuiar genetic studies should be undertaken in case of doubt Release stock idealiy should be closely-related genetically to the original native stock

(Anonymous 1992 2-31

Clearly a search of not only historic but also prehistoric information nnay be necessary The rapidly developing study of ancient DNA extracted from prehistoric skeletal tissues

The relevance of faunal anuljlsis to wildlife matlugement 115

(eg Richards et al 1993 1995 and references therein) seems to have great potential for contending with IUCN guidelines The technique is applicable to curated as wtll as newly acquired archaeological specimens It has however not yet been applied to Oregon coast sea otter remains

Given the lack of genetic material from eighteenth- and early nineteenth-century sea otters on the Oregon coast the problem becomes one of establishing the age of the character gradients displayed by eastern Pacific rim sea otters (see Wilson et al 1991 for a recent summary) and determining if those gradients have shifted spatially through time IIf those gradients have a prehistoric distribution in time andlor space similar to that documented using historic specimens a potential problem is raised If a particular form -represented by a particular character state -- of sea otter was present prehistorically in one location within the range and another form was present at another location then transplanting individuals from the former to the latter may be doomed from the start In such a case failure to re-establish a population could result on the one hand because the transplanted form is not well adapted to its new environment the extirpated form on the other hand is adapted to the environment of that location hence its difference in form

Several years ago I examined prehistoric sea otter remains recovered from archaeologi- cal sites on the coast of the state of Oregon - the middle latitudes of the historically documented range of sea otters (Lyman 1988b) The size of sea otter teeth recovered from those sites are more or less intermediate to those from Alaska (large) and those from southern California (small) but they also vary somewhat from this simple pattern Prehistoric Oregon sea otters match modern Alaskan sea otters in some characters in others they match modern California sea otters Precise details are unnecessary to the point I wish to make here which is Were the sea otters captured in Alaska and translocated to the Oregon coast in the 1970s of the proper phenotype (or genotype) Zooarchaeological evidence suggests they were not If this is a correct interpretation then perhaps those transplanted sea otters were doomed from the moment they were captured Whether or not my suggestion proves to be true or false and it is unclear which way the cards might fall given that we still have much to learn about this species (VanBlaricom arid Estes 1988) the significance of zooarchaeological data should be clear

Not only can zooarchaeological data inform us as to which form of a species might be the most appropriate one to transplant to an area such data have the potential to indicate the potential success of such transplant efforts Even if the appropriate form to transplant is clear this does not guarantee that the introduced form will survive in its new home This is precisely the argument offered by biologists working with sea otters on the Oregon coast The data substantiaiing their argument can be supplemented with zooarchaeological data An example of this is provided by the California condor (Gymnogyps cnlifornianus) of western North America a species presently near extinction (Emslie 1987) Zooarchaeo- logical and paleontological data suggest this species enjoyed a wider range during the late Pleistocene but that range shrank considerably with the onset of the postglacial Holocene epoch (Emslie 1987 Simons 1983) Those data also suggest that this species was a carrion feeder and that the terminal Pleistocene extinction of many genera of large mammals probably contributed to the decline in frequency and range of the California condor (Emslie 1987) Finally paleoecological data indicate attempts to reintroduce this species to portions of its prehistoric range would probably fail because of major differences

116 R Lee Lytriurz

between late Holocene floral and faunal communities and late Pleistocene commaanities (Emslie 1987)

In the absence of historic data or even with it (see below) zooarchaeological infor- mation may be critical to ensuring that the correct forms of animals are transplanted The wrong forms may be i l l suited to their new habitats or they may be well adapted and pro- liferate in their new homes Of course proliferation of artificially introduced species may on the one hand be beneficial and what is desired On the other hand as H think many of us are aware it may not be desired And the latter brings us to another kind of use that can be made of zooarchaeological data when wildlife management is the issue

Identifying exotic animals

Recalling the TUCNs guidelines for reintroductions if a particular form of organism (ecotype subspecies or species) is exotic to a park or wildlife preserve it map be necessary to remove it from the area in order to preserve the natural status of the ecosystem and the biota of the area Thus we must have a solid definition of exotic ipecieThe United Stttes National Park Service (NPS) defines an exotic species as one that occurs in a given place ts a result of direct or indirect deliberate or accidental actions by humans (not including de- liberate reintroductions) (Hester 1991 127) In contrast native species are those which presently occur or once did occur prior to sonic human influence in a given place area or region as the result of ecological processes that operate and have operatcd without signifi cant direct or indirect deliberate or accidental alteration by humans (NPS documents cited in Lyman 1988a 21 see also Wagner et al 1995) Clearly then if wildlife managcrs are to manage native ecosystems variously referred to as pristine ecosystems or wilder- ness -by exclusion or removal of exotic species and reintroduction of artiticially extirpated native species then they must have that baseline list of native species In the United Stares that list is derived from the earliest historical documents for the area included within a national park (eg Houston and Schreiner 1995 Lgteopold et al 1963see also Hoerr 1993) The ethnocentricity of such a procedure is as might be imagined unavoidable from a przc tical standpoint More importantly historical documents arc sometimes incomplete and are sometimes simply inaccurate Ihose written documents can of course be sup plemented with another kind of document - the zooarchaeological record Here I men tion two examples of such supplementation

Bison (Bison bison) were artificially introduced to a portion of the state of Alaska that was to subsequently become Wrangell-St Elias National Park and Preserve (Peek et al 1987) By definition these introduced bison should be considered exotic and are so corl-sidered by the NPS (Houston and Schreiner 1995) However zooarchaeological evidence indicates that bison were present in the area until at least 500 years ago or slightly later (reviewed in Peek et al 1987) Historic records suggest bison wcre not prcsent when the first white men visited the area in the middle and late nineteenth ccntury but non- fossilized bison skulls were collected from the arca early in the twentieth century (Peek et al 1987) Given the paucity of late Holocene zooarchaeological data in thc area and the unknown nature of the relevant fossilization processes it is unclear if bison were present in Alaska between about 450 years ago and the carly twentieth century (McDonaald 1981)

The relevance of faunal analyl to wrldllfe rnanugerrzent 117

The NPS considers the bison of Wrangell-St Elias National Park to be exotic due to the lack of evidence that bison were present when the tirst white explorers passed through the area (Houston and Schreiner 1995) As well NPS biologists note that the introduced forrn of bison was of an incorrect - non-native -genetic stock (a distinct subspecies or ecotype) given current beliefs about bison taxonomy For this reason alone perhaps the extant bison of Wrangell-St Elias should be removed However the zooarchaeological record is so poorly known that one cannot conclusively demonstrate that native bison were locally extinct by the beginning of the twentieth century This in turn raises the possibility that extant bison are hybrids of native and introduced genetic stocks Whether or not this possibility is in fact reality is irrelevant What is critical here is that only more zooarchaeological and paleontological research will establish first the timing of local extirpation of native Alaskan bison populations and second if all Alaskan populations were extinct when transplanting occurred in the middle of the twentieth century Once the facts are determined we will have strong bases for making a decision regarding the ultimate fate of the bison presently living in Wrangell-St Elias National Park and Preserve

I am much more familiar - and intimately involved -with a virtually identical situation that exists in western Washington state Here mountain goats (Oreamnos urnericanlls) were introduced in the 1920s to the Olympic Mountains A decade later in 1938 these mountains were designated Olympic National Park The present position of the NPS is that the mountain goats are exotic and plans are in mid-1995 under way to eradicate them The plan is controversial (Anunsen and Anunsen 1993 Scheffer 1993) as is the evidence mustered to support it (compare Houston et al 119941 with Lyman 11991a 1995a 1995b1) This is not the place to review that evidence in detail (see references cited) It is relevant however to note the significance of zooarchaeological evidence for final resolution of the controversy

The historical evidence concerning the pre-1920s presence of mountain goats in the Olympic Mountains is ambiguous Some documents indicate goats were absent while others indicate goats were present (Lyman 1994a 1995b) In fact the only way to demonstrate that mountain goats were present or absent from the Olympic Mountains prior to their 1920s introduction requires paleontological andor zooarchaeological data At present there are no paleontological data of any kind for the area and the zooarchaeological record is woefully inadequate and thus cannot serve as a basis for a management decision (Lyman 1995a) Park biologists note that the purpose of national parks in the United States is not to attempt to recreate late-Pleistocene or early Holocene biotas [The NPS] is primarily concerned with historic post-Columbian specres distri- butions (Houston and Schreiner 1995 207) The facts remain however the zooarchaeo- logical record is poorly known and the historic record is internally contradictory

From the perspective of dynamic biogeographic histories provided by zooarchaeology one thing is abundantly clear the definitions of native and exotic species used by the NPS to implement its policies are contradictory Exotic species occur in a given place as a result of actions by humans whereas native species are those which presently occur or once did occ~crprior to some human influence in a place as the result of natural ecological processes The italicized phrase is where the contradiction resides because it indicates that i f for example mountain goats occurred at any tirne in the past in the Olympic Mountains

118 R Lee Lyman

- or bison in Wrangell-St Elias - then by definition the extant individuals represcl~i a native species But the introduced species are also by definition exotic having been transplanted from northwestern Canada and southeastern Alaska Olympic Nationai Parks mountain goats are of a different genetic stock and thus are a form distinct fromi mountain goats that may have been in the Olympic Mountains 200 years ago Rccogr-iizing this contradiction some years ago Y argued that if the prehistoric remains of rnonirrtain goats should be found in the Olympic Mountains then the NPS should rethink policy issues (Lyman I9RWa 22) Some NPS biologists have only recently come to the same conclusion (Houston and Schreiner 1995) And given the discussions above of appropriata forms (ecotypes subspecies species) and the benefit of hindsight I would argue that the policies and definitions of key terms should also be reworded to reflect the reality of wildlife management practice (Anonymous 1992 OBrien and Mayr 1991) Thus given their perspective of time depth zooarchaeologists not only can contribute significant paleobiological information to wildlife management concerns but perhaps they can also help write workable policies (see Wagner et al 1995 for other examples) Paleobiological information provided by zooarchaelogical research may also help wildlife managers plals for the future as the final example demonstrates

Biological preserves and planning for the future

Thus far the examples of zooarchaeology applied to wildlife management concerns have focused on the use of information on prehistoric faunal dynamics to inform management decisions That same kind of information can also be used to make broad predictions about the future effects of certain kinds of decisions For instance as alluded to in the first example presented above the spatial boundaries of parks and wildlife preserves may Piavc significant implications for the long-term survival of populations found withirl those boundaries In an important contribution to the role of planning for the future on tllc trasis of the past Graham (1988) notes that the boundaries of many parks and preserves are defined based on modern climatic and environmental conditions However tkrose conditions as we all know change and the probability of significant change increases as the length of time considered increases Graham underscores the [act that much planning lo preserve terrestrial biological diversity operates under the assumption that biotas or communities of organisms tend to respond to environmental change as intact units The

paleobiological record indicates however that individual species respond to environ- mental changes by migrating in different directions at different rates during different times (Graham 1988 392) Such individualistic responses to environmental change mean that the biological preserves of today are artifacts of the time when they were identified and created Thus Graham (1988 393) argues that constructing future reserves as habitat islands without avenues of migration [corridors] would probably result in [extirpation of bounded populations of managed species]

Graham (1988) thus suggests that as in the example of the Great Basins mountaintop mammals migration corridors are critical to organism survival and that preserves must be located on the landscape in such a rnanner as to be conducive to inter-areal migration I suggest following my earlier discussion that knowing where ihose corridors were located

The relevance of faunal analjgtsis to wildlife management 119

in the past surely will help in locating future preserves And as Graham and Graham (1994) indicate detailed knowledge of the historical and ecological biogeography of a taxon could prove critical to reconstructing appropriate habitats for reintroduced populations of the taxon Such knowledge can often come only from the fossil record

Extinction -of populations of species of lineages - is a natural fact of life (eg Raup 1984) But ignorance resulted in the loss of the passenger pigeon (Ectopistes migratorius) from North America the Great Auk (Pingninns impennis) from the North Atlantic and the dodo (Raphus cuci~llatus) from Mauritius in the Indian Ocean Our often stumbling attempts at wildlife management may due to ignorance ultimately lead to other such losses (eg Graham 1988 Hess 1993 Kay 1994) Ihe international move to maintain biological diversity - not just the taxonomic richness of biotic communities but the particular structure of those communities - seems to be in the correct direction if we are to maintain some quality of life and wilderness Understanding the loss of diversity often demands zooarchaeological and related data (eg Culotta 1995 Steadman 1995) and will help us avoid future inadvertent and unintentional losses Diamonds (1984) Rosetta Stone of using knowledge of modern extinctions to understand prehistoric extinctions could and perhaps should be turned on its head

Discussion

Nearly eighty years ago archaeologist W J Wintemberg (1919) noted how zooarchaeo- logical research could contribute to zoological knowledge and indirectly wildlife management His comments were repeated and elaborated thirty years later by biologist R M Gilmore (1949) Their remarks have largely gone unheeded perhaps because like Legge (1978 129 130) many of us see our job as aiming to contribute to the sum of the archaeological knowledge and that bone data are archaeological data I agree with Legge but also with Wintemberg and Gilmore we should be more than just consumers of wildlife data (cf Will 1982)

The potential topics under the broad umbrella of wildlife management to which zooarchaeological research might contribute are numerous I have touched on only a fevv of the more readily identifiable ones above There are to be sure others of some significance For example the modern fisheries industry of the eastern Pacific Ocean has been disrupted several times in the recent past by major changes in atmospheric circulation patterns causing alteration of ocean currents These events - variously referred to as El Nifio or Southern Oscillation events -appear to have some periodicity (Rasmusson 1985) but the historic record is of limited utility in plotting the precise nature of that periodicity These events clearly influence much of the oceanic biota (eg Arnold and Tissot 1993 Trillmich and Limberger 1985) Given sufficient temporal resolution the zooarchaeologi- cal record thus may eventually provide the time depth necessary to clarify the periodicity and magnitude of El Nifio events and thus allow more accurate predictions of future events

Zooarchaeological research has also shown that humans have long had a hand in modifying ecosystems Based on his extensive research on the prehistoric extinctions of Pacific Island bird populations and species Steadman (1995 1130) concludes that humans

tilay find Yt difficult to manage scarce resources in a sustainable way That humans contributed to these extinctions seems clear (cg Weisler and Cargett 1993) and thus Steadmans conclusion has the proper ominous tone But zooarchaeological research cair also perhaps help us out of such situations The wapiti (Cervns elaphus) of Yellowstone National Park in the United States are an example Herc nearly 60000 wapiti are managed and to a significant degree protected from human predation However historic and zooarchaeological research indicates that this population is a function of modern wildlife management historic records virtually vithout exception fail to mention wapiti i l l

the area and zooarchaeological data indicate this large ungulate was very riirc there during the last several thousand years (Kay 1994) Clearly if the Yellowstone ecosSsten1 is to be managed and maintained as a pristine wilderness in perpetuity (a laudable but in reality unattainable goal see Wager et al 1995) things must change before the ecosystem is irreparably altered A virtually identical situation exists in Rocky Mountain National Park sotile 550 km southwest of Yellowstone (Hcss 1993) where again zooarchaeologicai and other data from prehistoric times might help rcsolve debates

The few examples of how zooarchaeological research can be applied to t i ioder~~ problems I have outlined are rather specific to a place andlor to a taxon I do not perccive that to bc a weakness of an applied zooarchaeology but rather as an intellectual challenge Broadening the scope of zooarchaeology in the manner in which 1 have discussed will not be easy but i t will be beneficial to our Suture not only from the perspective of contributing to ensuring the preservation of biological diversity for future generations but also as 1conclude below from the perspective of our discipline which tilight otherwise become increasingly perceived as the pursuit of esoteric knotvledge of little practical use

One might argue that the discussion here could be expanded by suggesting that the detection of animal residues such as blood and hair (eg Loy 1993) could also providr information on the taxonomic identity of species native to the locale where the tools were collected (eg Hyland et al 1990) I doubt however that such a study would convince wildlife managers or all zooarchaeologists that a species was native given that stone tools were often transported significant distances by their tiiakers (eg Meltzer 1995) 1 beiievc multiple instances of such residues from multiple sites would be necessary to build 21

strong argument A partial skeleton or selected isolated bones and teeth of the species in que5tion recovered from a s~ngle ~ t e uould probably be muclr more conblnclng (Lyman 1994b)

I have attempted here to describe the structure of an applied zooarchaeology This is not a plea to legitimize what some of us do for fun andlor a living Rather i t is an attempt to broaden the scope of zooarchaeological research 1 am all for research for the sake of research if there is a problem in need of solving or a question in need of an answer But B am also concerned with the future of zooarchaeology Too many of our new doctoral str~dcnts are finding that the job market is closed and too marly of us arc finding it increasingly difficult to secure funding for research Perhaps a developed applied zooarchaeology wil

The relevance o f faunal analysis t o wildlife rnanagement 121

provide a new job market and new sources of funding It seems self-evident to me that if a national park for example has a staff archaeologist who deals with the archaeological resources on park lands and has a staff biologist who deals with biological resources on park lands then a marriage of the two would benefit both However given the increasing specialization of both archaeology and biology a viable marriage may not be possible Ihc obvious alternative is a park staff zooarchaeologist who has a deep knowledge ancl understanding of both disciplines and has been trained to integrate the two

New sources of funding may also become available if we can convince the wildlife management community that because its decisions and policies are typically aimed at tht future knowing something of the past can result in wise decisions That knowledge will of course come with a price tag The cost we may encounter is that members of the wildlife management community -while they may well have a solid knowledge of the time depth of evolutionary processes- often tend to view biotas as static ever enduring entities That is biotas have no past or at least their history is insignificant because they are being managed for the future While we certainly cannot predict exactly when the next ice age or glacial period will begin we can I think argue convincingly that climates and environments will change and knowing this we can use the prehistoric record to test our predictions about how certain kinds of changes may affect biotas of the future And confirmed - or even rejected or falsified -predictions would be the strong selling point Federal land-managing agencies whose charge includes wildlife management might well pay a zooarchaeologist to help them make wise management decisions The other selling point is that a less desirable kind of price -ecosystem destruction and loss of biodiversity -might otherwise have to be paid

Acknowledgements

I thank Donald K Grayson Adrian Lister Michael J OBrien and Ken Ihornas for comments on earlier versions of the manuscript

Departtnent o f Anthropology University o f Missouri-Columbia

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Mcltzcr D J 1995 Clocking the first Americans Annual Review of Anthropology 24 71-45

124 R Lee E y n ~ u n

OBricn S J and Mays E 1991 Burcauc1atic mischief rccogni~ing enciangcrcd spccics and subspecies Scicncc 251 1 187--8

Ogden A 1933 Russian sea ottcr anal seal hunting on the Califorilia coast Caij(~rrrirrNisroricci1 Qmcrrrerly 12 29-5 1

Olscn S I and Olscn I W 1981 A conin-icnt on nomcnclaturc in faunal studies iui~cric~un Antiquity 46 192-4

Peck J M Miclucllc 14 C and Wright M 61987 Arc bison cuotic in the Wrangcll-St Elia National Park and Pre~cr~c Envivnnnlcrzrcrl iWrlrlcrgcineilr I I l(195 3

Quinn J Iand ifarrison S P 1988 Effects of habitat fragmentation anhi isolation on spci-is richness evidence from biogeographic patterns Oecologiii 75 132-40

Rasmusson E M 1985 El Nino and asiations in climate Atzcricr~z Scic3nritt 73 168-77

Raup D M 1984 Death of spccics I n Errirrcrions (etl M 14 Nitccllti) dlhicago University of Chicago Press pp 1-19

IcnsbcrgcrJ M and Barnosky A I) 1993 Short-term fluctuations in slniill mammals ot the lato Pleistocene from eastern Washington Wn iWorphologicrrl Chr~rigc irr C)iir~t~rricrr~ Mrrinnzcrls (J ~ i i~ t l~ Alzcrica (eds R A Martin and 4 Igt 131rnosky) Qambridgc Ciimbsidgc University Prcss pp 299-342

Richards M Smallcy KSykcs R and Iledges BP 1993 iir-ch21eology and genetics anaiysirig DNA from skeletal remains Worlrl~lrchtrcolog~5 18-28

Richards M B Sykcs B C and Hedges MF M 1995 Authenticating D N A extriicted Irorri ancient skeletal remains Iourr7crl of Arcl~aeologicril Scicncc 22 2W-9

Faohison ND 1978 Zooarchaeology its hiirto~y and dcvclopnirn~7r7rrcssrc Anthropologic~ul A~orirition Miscellancotrs Pripcr 2 1-2

Rocst A 1 1973 Subspecies of the sea ottcs Er7llytira Iiitris Io5 ii7gcles Coirnty Nnturcrl tiisrory Mltscirrn (ontributior~s in Scicncr 252

Rocst A 1 1979 A re-evaluation oi sea o t t c ~ taxonomy Paper prcscntcci at the Sea Ottcr Workshop Santa Barbara Cilifornia

Scheffer V B 1993 Tlie Olympic Mo~rntain goat conrrovcrs) a pcr5pectivc Coriscrvtrtior Biology 7 916-19

Sclicffcr V B and Wilkc F 1950 Validity of the subspccics E~llzydrii 1iltris r7crcic the southcrn s ix ottcr Joltvnill qf the Wcr~I7ir7gton Accrtlcuny 01 S~icr~cr402672

Simons I) R 1083 Interactions lsctween Calilornin corrdors and humans in prehistoric far wcstern North America In Vcrltcrrr Biology und ~tlunuget~ierlt (cds S KWilbur and J A Jackson) Berkeley University or Ciliforni~i Press pp 470-91

Steadman D xi 1995 Prehistoric cstinctions of Pacific isiarld birds biodiversity nlccts zooarchacology Scieilce 267 1 123-3 1

Trillrnich F and Limbcrger D 1985 Drastic effects of 871 Nirio on Ga1apigos piniiipcd Oecologiu 67 19-22

VanBlaricorn 6Rand Estcs 9 A (cds) 1988 l h p Bot~irnlrniij bcologj oj Seu O t r e r ~ (Berlin

Springer-Verliig

Wagner F 11 Forest RC i i l l R R Mcdullough 14 R Pcltoil M MPorTcr W F and Salwasscr 13 1995 WildlifePolicies in the U S Nirrionul Purks Washington DC Island Prcsi

Weisler M I and Gargett R H 1993 Pacific island avian cstinctions the taphonorny or human predation Arclzucoloyj in Occuniu 28 85-03

Wendell F E flardy R A An-rc5 1 land Burgi K T 1986 Tcrnporalanci spatial patterns in

The relevance of faunal analysis to wildlife management 113

be located The fossil record might help us determine where these corridors existed in the past as can be illustrated with another example

The pygmy rabbit (Brachylagi~s idahoensis) is a diminutive leporid with a modern range restricted to two areas of the western United States One area is relatively large and encompasses portions of four states northern Nevada western Utah southern Idaho and southeastern Oregon The other historically documented area where this species occurs is the central portion of eastern Washington state Populations in the two areas are presently isolated from one another The wildlife management coricern here is that the eastern Washington population has over the past forty years shrunk to an alarmingly small size prompting the Washington Wildlife Commission to list this population as threatened and to suggest that i t be listed as endangered (McAllister and Allen 1993) and to develop a management pan with the aim of ensuring survival of the population (McAllister and Allen 1994) The zooarchaeological record of this species provides insight to why this population is threatened with extirpation and how i t might be preserved

The zooarchaeological record for pygmy rabbits indicates this species probably occupied a wider range in central Washington during the Holocene than i t presently does Remains of this species have now been recovered from eleven archaeological and two paleontological sites in the area (updated from Lyman 1991) Some of these are extrali~nitalrecords that is some remains have been recovered from geographic locations where pygmy rabbits have not been historically documented to occur In brief the fossil (botanical and faunal) record suggests that when big sagebrush (Artemisia tridentata) was more widespread in central Washington -- during the middle Holocene climatic interval known as the Altithermal - pygmy rabbits were also more widespread When the Altithermal ended about 4500 years ago the range of sagebrush shrank and so too did the range of pygmy rabbits Then during the late nineteenth and early twentieth centuries land was cleared of sagebrush for agricultural purposes This prompted a second diminution of sagebrush and pygmy rabbit range

Ecological studies of pygmy rabbits indicate that this leporid is dependent on big sagebrush for food and for shelter from predation (Green and Flinders 1980a 1980b) The zooarchaeological record in conjunction with paleobotanical data such as that from pollen records confirms this as both species appear to have responded to climatic change in like manner Washington Department of Wildlife biologists have made use of both modern and zooarchaeological data in their planning efforts (McAllister and Allen 1993 1994) The fossil record suggests that the prehistoric source of Washingtons pygmy rabbits resides in the area of the larger extant population to the south Late Pleistocene fossils of pygmy rabbits recovered from a paleontological site between the two areas (Rensberger and Barnosky 1993) tend to confirm this suggestion While i t would probably be impractical to develop a migration corridor between the two populations today it would also be impossible given the climatic history of the area The probable corridor between the two today is characterized by vegetation habitats that are not conducive to tht survival of pygmy rabbits The zooarchaeological record does indicate however that the extant population might be supplemented by individuals transplanted from southeasterri Oregon and that the maintenance of habitats dominated by big sagebrush is critical to the survival of the species

114 R Lee Lyman

When managing small isolated populations we may not have many options other than to preserve those populations as they presently exist Yn some cases management efforts have failed and populations have been extirpated When this occurs one management alternative is to transplant individuals from other populations to the vacated rangc in an attempt to re-establish a population Zooarchaeological research can also plav an important role in these situations

Re-establishment of extirpated populations

Eighteenth- and nineteenth-century conimercial exploitation of sea otters (Enhydru lirtris) along the western coast of North America ultimately led to the extirpation of this species from much of its range (Kenyon 1969 Ogden 1993 Roest 1973) Attempts to re-establish populations along the middle latitudes of that coast took place in the 1970s and thus far have met with less than perfect success (Jameson et al 1982 1986 MacAskie 1987) The typically expressed reason for the limited success of these efforts is that ecosystems to which sea otters have been transplanted have heen modified too exter~sively by modern human industry to support them Y wonder i f however part of the reason might resicle in a factor that only zooarchaeological research car measure

Sea otter skulls collectetl from much of the once occupied eastern Pacific range of sea otters display various character gradients from the Aleutian Islands to southern California Some of these characters have been used to argue for or against the existence oi multiple subspecies of sea otter (Davis and Lidicker 1975 Roest 1973 1979 Scheffer and Wilke 1950 Wilson et al 1991) Final resolution of this debate will be critical to futtlrc wildlife management activities first because national legislation in the United States concerns species subspecies and populations (OWrien and Mayr 1901) and also beca~ase of conflict between one population (a possibly distinct subspecies) of sea otters and the modern fisheries industry (Wendell et al 1986) lhe significant aspect to all of this Is the transplanting of sea otters ILegally only forms (ecotypes or subspecies) native to ar arcs should be introduced there during efforts to re-establish populations This also m~aites biological sense as i t is likely that those native forms unique to an area are the product of millennia of evolution

The International Union for the Conservation of Nature and Natural Resources PUCN (recently renamed the World Conservation Un~on) has drafted guidelines Cor such -

reintroductions These read in part

An assessment should be made of the taxonomic status of individuals to ba re-introduced They should be of the same taxonomic unit (and ideally closely related genetically) as those which were extirpated An investigation of historical information about the loss and fate of individuals from the re-introduction area as well a mmolecuiar genetic studies should be undertaken in case of doubt Release stock idealiy should be closely-related genetically to the original native stock

(Anonymous 1992 2-31

Clearly a search of not only historic but also prehistoric information nnay be necessary The rapidly developing study of ancient DNA extracted from prehistoric skeletal tissues

The relevance of faunal anuljlsis to wildlife matlugement 115

(eg Richards et al 1993 1995 and references therein) seems to have great potential for contending with IUCN guidelines The technique is applicable to curated as wtll as newly acquired archaeological specimens It has however not yet been applied to Oregon coast sea otter remains

Given the lack of genetic material from eighteenth- and early nineteenth-century sea otters on the Oregon coast the problem becomes one of establishing the age of the character gradients displayed by eastern Pacific rim sea otters (see Wilson et al 1991 for a recent summary) and determining if those gradients have shifted spatially through time IIf those gradients have a prehistoric distribution in time andlor space similar to that documented using historic specimens a potential problem is raised If a particular form -represented by a particular character state -- of sea otter was present prehistorically in one location within the range and another form was present at another location then transplanting individuals from the former to the latter may be doomed from the start In such a case failure to re-establish a population could result on the one hand because the transplanted form is not well adapted to its new environment the extirpated form on the other hand is adapted to the environment of that location hence its difference in form

Several years ago I examined prehistoric sea otter remains recovered from archaeologi- cal sites on the coast of the state of Oregon - the middle latitudes of the historically documented range of sea otters (Lyman 1988b) The size of sea otter teeth recovered from those sites are more or less intermediate to those from Alaska (large) and those from southern California (small) but they also vary somewhat from this simple pattern Prehistoric Oregon sea otters match modern Alaskan sea otters in some characters in others they match modern California sea otters Precise details are unnecessary to the point I wish to make here which is Were the sea otters captured in Alaska and translocated to the Oregon coast in the 1970s of the proper phenotype (or genotype) Zooarchaeological evidence suggests they were not If this is a correct interpretation then perhaps those transplanted sea otters were doomed from the moment they were captured Whether or not my suggestion proves to be true or false and it is unclear which way the cards might fall given that we still have much to learn about this species (VanBlaricom arid Estes 1988) the significance of zooarchaeological data should be clear

Not only can zooarchaeological data inform us as to which form of a species might be the most appropriate one to transplant to an area such data have the potential to indicate the potential success of such transplant efforts Even if the appropriate form to transplant is clear this does not guarantee that the introduced form will survive in its new home This is precisely the argument offered by biologists working with sea otters on the Oregon coast The data substantiaiing their argument can be supplemented with zooarchaeological data An example of this is provided by the California condor (Gymnogyps cnlifornianus) of western North America a species presently near extinction (Emslie 1987) Zooarchaeo- logical and paleontological data suggest this species enjoyed a wider range during the late Pleistocene but that range shrank considerably with the onset of the postglacial Holocene epoch (Emslie 1987 Simons 1983) Those data also suggest that this species was a carrion feeder and that the terminal Pleistocene extinction of many genera of large mammals probably contributed to the decline in frequency and range of the California condor (Emslie 1987) Finally paleoecological data indicate attempts to reintroduce this species to portions of its prehistoric range would probably fail because of major differences

116 R Lee Lytriurz

between late Holocene floral and faunal communities and late Pleistocene commaanities (Emslie 1987)

In the absence of historic data or even with it (see below) zooarchaeological infor- mation may be critical to ensuring that the correct forms of animals are transplanted The wrong forms may be i l l suited to their new habitats or they may be well adapted and pro- liferate in their new homes Of course proliferation of artificially introduced species may on the one hand be beneficial and what is desired On the other hand as H think many of us are aware it may not be desired And the latter brings us to another kind of use that can be made of zooarchaeological data when wildlife management is the issue

Identifying exotic animals

Recalling the TUCNs guidelines for reintroductions if a particular form of organism (ecotype subspecies or species) is exotic to a park or wildlife preserve it map be necessary to remove it from the area in order to preserve the natural status of the ecosystem and the biota of the area Thus we must have a solid definition of exotic ipecieThe United Stttes National Park Service (NPS) defines an exotic species as one that occurs in a given place ts a result of direct or indirect deliberate or accidental actions by humans (not including de- liberate reintroductions) (Hester 1991 127) In contrast native species are those which presently occur or once did occur prior to sonic human influence in a given place area or region as the result of ecological processes that operate and have operatcd without signifi cant direct or indirect deliberate or accidental alteration by humans (NPS documents cited in Lyman 1988a 21 see also Wagner et al 1995) Clearly then if wildlife managcrs are to manage native ecosystems variously referred to as pristine ecosystems or wilder- ness -by exclusion or removal of exotic species and reintroduction of artiticially extirpated native species then they must have that baseline list of native species In the United Stares that list is derived from the earliest historical documents for the area included within a national park (eg Houston and Schreiner 1995 Lgteopold et al 1963see also Hoerr 1993) The ethnocentricity of such a procedure is as might be imagined unavoidable from a przc tical standpoint More importantly historical documents arc sometimes incomplete and are sometimes simply inaccurate Ihose written documents can of course be sup plemented with another kind of document - the zooarchaeological record Here I men tion two examples of such supplementation

Bison (Bison bison) were artificially introduced to a portion of the state of Alaska that was to subsequently become Wrangell-St Elias National Park and Preserve (Peek et al 1987) By definition these introduced bison should be considered exotic and are so corl-sidered by the NPS (Houston and Schreiner 1995) However zooarchaeological evidence indicates that bison were present in the area until at least 500 years ago or slightly later (reviewed in Peek et al 1987) Historic records suggest bison wcre not prcsent when the first white men visited the area in the middle and late nineteenth ccntury but non- fossilized bison skulls were collected from the arca early in the twentieth century (Peek et al 1987) Given the paucity of late Holocene zooarchaeological data in thc area and the unknown nature of the relevant fossilization processes it is unclear if bison were present in Alaska between about 450 years ago and the carly twentieth century (McDonaald 1981)

The relevance of faunal analyl to wrldllfe rnanugerrzent 117

The NPS considers the bison of Wrangell-St Elias National Park to be exotic due to the lack of evidence that bison were present when the tirst white explorers passed through the area (Houston and Schreiner 1995) As well NPS biologists note that the introduced forrn of bison was of an incorrect - non-native -genetic stock (a distinct subspecies or ecotype) given current beliefs about bison taxonomy For this reason alone perhaps the extant bison of Wrangell-St Elias should be removed However the zooarchaeological record is so poorly known that one cannot conclusively demonstrate that native bison were locally extinct by the beginning of the twentieth century This in turn raises the possibility that extant bison are hybrids of native and introduced genetic stocks Whether or not this possibility is in fact reality is irrelevant What is critical here is that only more zooarchaeological and paleontological research will establish first the timing of local extirpation of native Alaskan bison populations and second if all Alaskan populations were extinct when transplanting occurred in the middle of the twentieth century Once the facts are determined we will have strong bases for making a decision regarding the ultimate fate of the bison presently living in Wrangell-St Elias National Park and Preserve

I am much more familiar - and intimately involved -with a virtually identical situation that exists in western Washington state Here mountain goats (Oreamnos urnericanlls) were introduced in the 1920s to the Olympic Mountains A decade later in 1938 these mountains were designated Olympic National Park The present position of the NPS is that the mountain goats are exotic and plans are in mid-1995 under way to eradicate them The plan is controversial (Anunsen and Anunsen 1993 Scheffer 1993) as is the evidence mustered to support it (compare Houston et al 119941 with Lyman 11991a 1995a 1995b1) This is not the place to review that evidence in detail (see references cited) It is relevant however to note the significance of zooarchaeological evidence for final resolution of the controversy

The historical evidence concerning the pre-1920s presence of mountain goats in the Olympic Mountains is ambiguous Some documents indicate goats were absent while others indicate goats were present (Lyman 1994a 1995b) In fact the only way to demonstrate that mountain goats were present or absent from the Olympic Mountains prior to their 1920s introduction requires paleontological andor zooarchaeological data At present there are no paleontological data of any kind for the area and the zooarchaeological record is woefully inadequate and thus cannot serve as a basis for a management decision (Lyman 1995a) Park biologists note that the purpose of national parks in the United States is not to attempt to recreate late-Pleistocene or early Holocene biotas [The NPS] is primarily concerned with historic post-Columbian specres distri- butions (Houston and Schreiner 1995 207) The facts remain however the zooarchaeo- logical record is poorly known and the historic record is internally contradictory

From the perspective of dynamic biogeographic histories provided by zooarchaeology one thing is abundantly clear the definitions of native and exotic species used by the NPS to implement its policies are contradictory Exotic species occur in a given place as a result of actions by humans whereas native species are those which presently occur or once did occ~crprior to some human influence in a place as the result of natural ecological processes The italicized phrase is where the contradiction resides because it indicates that i f for example mountain goats occurred at any tirne in the past in the Olympic Mountains

118 R Lee Lyman

- or bison in Wrangell-St Elias - then by definition the extant individuals represcl~i a native species But the introduced species are also by definition exotic having been transplanted from northwestern Canada and southeastern Alaska Olympic Nationai Parks mountain goats are of a different genetic stock and thus are a form distinct fromi mountain goats that may have been in the Olympic Mountains 200 years ago Rccogr-iizing this contradiction some years ago Y argued that if the prehistoric remains of rnonirrtain goats should be found in the Olympic Mountains then the NPS should rethink policy issues (Lyman I9RWa 22) Some NPS biologists have only recently come to the same conclusion (Houston and Schreiner 1995) And given the discussions above of appropriata forms (ecotypes subspecies species) and the benefit of hindsight I would argue that the policies and definitions of key terms should also be reworded to reflect the reality of wildlife management practice (Anonymous 1992 OBrien and Mayr 1991) Thus given their perspective of time depth zooarchaeologists not only can contribute significant paleobiological information to wildlife management concerns but perhaps they can also help write workable policies (see Wagner et al 1995 for other examples) Paleobiological information provided by zooarchaelogical research may also help wildlife managers plals for the future as the final example demonstrates

Biological preserves and planning for the future

Thus far the examples of zooarchaeology applied to wildlife management concerns have focused on the use of information on prehistoric faunal dynamics to inform management decisions That same kind of information can also be used to make broad predictions about the future effects of certain kinds of decisions For instance as alluded to in the first example presented above the spatial boundaries of parks and wildlife preserves may Piavc significant implications for the long-term survival of populations found withirl those boundaries In an important contribution to the role of planning for the future on tllc trasis of the past Graham (1988) notes that the boundaries of many parks and preserves are defined based on modern climatic and environmental conditions However tkrose conditions as we all know change and the probability of significant change increases as the length of time considered increases Graham underscores the [act that much planning lo preserve terrestrial biological diversity operates under the assumption that biotas or communities of organisms tend to respond to environmental change as intact units The

paleobiological record indicates however that individual species respond to environ- mental changes by migrating in different directions at different rates during different times (Graham 1988 392) Such individualistic responses to environmental change mean that the biological preserves of today are artifacts of the time when they were identified and created Thus Graham (1988 393) argues that constructing future reserves as habitat islands without avenues of migration [corridors] would probably result in [extirpation of bounded populations of managed species]

Graham (1988) thus suggests that as in the example of the Great Basins mountaintop mammals migration corridors are critical to organism survival and that preserves must be located on the landscape in such a rnanner as to be conducive to inter-areal migration I suggest following my earlier discussion that knowing where ihose corridors were located

The relevance of faunal analjgtsis to wildlife management 119

in the past surely will help in locating future preserves And as Graham and Graham (1994) indicate detailed knowledge of the historical and ecological biogeography of a taxon could prove critical to reconstructing appropriate habitats for reintroduced populations of the taxon Such knowledge can often come only from the fossil record

Extinction -of populations of species of lineages - is a natural fact of life (eg Raup 1984) But ignorance resulted in the loss of the passenger pigeon (Ectopistes migratorius) from North America the Great Auk (Pingninns impennis) from the North Atlantic and the dodo (Raphus cuci~llatus) from Mauritius in the Indian Ocean Our often stumbling attempts at wildlife management may due to ignorance ultimately lead to other such losses (eg Graham 1988 Hess 1993 Kay 1994) Ihe international move to maintain biological diversity - not just the taxonomic richness of biotic communities but the particular structure of those communities - seems to be in the correct direction if we are to maintain some quality of life and wilderness Understanding the loss of diversity often demands zooarchaeological and related data (eg Culotta 1995 Steadman 1995) and will help us avoid future inadvertent and unintentional losses Diamonds (1984) Rosetta Stone of using knowledge of modern extinctions to understand prehistoric extinctions could and perhaps should be turned on its head

Discussion

Nearly eighty years ago archaeologist W J Wintemberg (1919) noted how zooarchaeo- logical research could contribute to zoological knowledge and indirectly wildlife management His comments were repeated and elaborated thirty years later by biologist R M Gilmore (1949) Their remarks have largely gone unheeded perhaps because like Legge (1978 129 130) many of us see our job as aiming to contribute to the sum of the archaeological knowledge and that bone data are archaeological data I agree with Legge but also with Wintemberg and Gilmore we should be more than just consumers of wildlife data (cf Will 1982)

The potential topics under the broad umbrella of wildlife management to which zooarchaeological research might contribute are numerous I have touched on only a fevv of the more readily identifiable ones above There are to be sure others of some significance For example the modern fisheries industry of the eastern Pacific Ocean has been disrupted several times in the recent past by major changes in atmospheric circulation patterns causing alteration of ocean currents These events - variously referred to as El Nifio or Southern Oscillation events -appear to have some periodicity (Rasmusson 1985) but the historic record is of limited utility in plotting the precise nature of that periodicity These events clearly influence much of the oceanic biota (eg Arnold and Tissot 1993 Trillmich and Limberger 1985) Given sufficient temporal resolution the zooarchaeologi- cal record thus may eventually provide the time depth necessary to clarify the periodicity and magnitude of El Nifio events and thus allow more accurate predictions of future events

Zooarchaeological research has also shown that humans have long had a hand in modifying ecosystems Based on his extensive research on the prehistoric extinctions of Pacific Island bird populations and species Steadman (1995 1130) concludes that humans

tilay find Yt difficult to manage scarce resources in a sustainable way That humans contributed to these extinctions seems clear (cg Weisler and Cargett 1993) and thus Steadmans conclusion has the proper ominous tone But zooarchaeological research cair also perhaps help us out of such situations The wapiti (Cervns elaphus) of Yellowstone National Park in the United States are an example Herc nearly 60000 wapiti are managed and to a significant degree protected from human predation However historic and zooarchaeological research indicates that this population is a function of modern wildlife management historic records virtually vithout exception fail to mention wapiti i l l

the area and zooarchaeological data indicate this large ungulate was very riirc there during the last several thousand years (Kay 1994) Clearly if the Yellowstone ecosSsten1 is to be managed and maintained as a pristine wilderness in perpetuity (a laudable but in reality unattainable goal see Wager et al 1995) things must change before the ecosystem is irreparably altered A virtually identical situation exists in Rocky Mountain National Park sotile 550 km southwest of Yellowstone (Hcss 1993) where again zooarchaeologicai and other data from prehistoric times might help rcsolve debates

The few examples of how zooarchaeological research can be applied to t i ioder~~ problems I have outlined are rather specific to a place andlor to a taxon I do not perccive that to bc a weakness of an applied zooarchaeology but rather as an intellectual challenge Broadening the scope of zooarchaeology in the manner in which 1 have discussed will not be easy but i t will be beneficial to our Suture not only from the perspective of contributing to ensuring the preservation of biological diversity for future generations but also as 1conclude below from the perspective of our discipline which tilight otherwise become increasingly perceived as the pursuit of esoteric knotvledge of little practical use

One might argue that the discussion here could be expanded by suggesting that the detection of animal residues such as blood and hair (eg Loy 1993) could also providr information on the taxonomic identity of species native to the locale where the tools were collected (eg Hyland et al 1990) I doubt however that such a study would convince wildlife managers or all zooarchaeologists that a species was native given that stone tools were often transported significant distances by their tiiakers (eg Meltzer 1995) 1 beiievc multiple instances of such residues from multiple sites would be necessary to build 21

strong argument A partial skeleton or selected isolated bones and teeth of the species in que5tion recovered from a s~ngle ~ t e uould probably be muclr more conblnclng (Lyman 1994b)

I have attempted here to describe the structure of an applied zooarchaeology This is not a plea to legitimize what some of us do for fun andlor a living Rather i t is an attempt to broaden the scope of zooarchaeological research 1 am all for research for the sake of research if there is a problem in need of solving or a question in need of an answer But B am also concerned with the future of zooarchaeology Too many of our new doctoral str~dcnts are finding that the job market is closed and too marly of us arc finding it increasingly difficult to secure funding for research Perhaps a developed applied zooarchaeology wil

The relevance o f faunal analysis t o wildlife rnanagement 121

provide a new job market and new sources of funding It seems self-evident to me that if a national park for example has a staff archaeologist who deals with the archaeological resources on park lands and has a staff biologist who deals with biological resources on park lands then a marriage of the two would benefit both However given the increasing specialization of both archaeology and biology a viable marriage may not be possible Ihc obvious alternative is a park staff zooarchaeologist who has a deep knowledge ancl understanding of both disciplines and has been trained to integrate the two

New sources of funding may also become available if we can convince the wildlife management community that because its decisions and policies are typically aimed at tht future knowing something of the past can result in wise decisions That knowledge will of course come with a price tag The cost we may encounter is that members of the wildlife management community -while they may well have a solid knowledge of the time depth of evolutionary processes- often tend to view biotas as static ever enduring entities That is biotas have no past or at least their history is insignificant because they are being managed for the future While we certainly cannot predict exactly when the next ice age or glacial period will begin we can I think argue convincingly that climates and environments will change and knowing this we can use the prehistoric record to test our predictions about how certain kinds of changes may affect biotas of the future And confirmed - or even rejected or falsified -predictions would be the strong selling point Federal land-managing agencies whose charge includes wildlife management might well pay a zooarchaeologist to help them make wise management decisions The other selling point is that a less desirable kind of price -ecosystem destruction and loss of biodiversity -might otherwise have to be paid

Acknowledgements

I thank Donald K Grayson Adrian Lister Michael J OBrien and Ken Ihornas for comments on earlier versions of the manuscript

Departtnent o f Anthropology University o f Missouri-Columbia

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Graham RW and Graham M A 1994 Late Quaternary distribution of Marter in North America In Mcrrten~ Sublcs urtd Fislrcrs Biology and Conservation (eds S W Ruskirk 14 S Harestad M 6Raphael and RA Powell) Itllaca NY Corncll University Prcss pp 2 6 5 8

Grayson D M 1991 Tllc biogeographic history of small mammals in the Great Basin observations on thc last 20000 years Journal of Mammalogy 68 359-75

Green J S and Flinders J T 1980a Brachylrig~s idahoerisis Manznzalian Specics 125 1--4

Green J S and Flindcrs J T 1980b Habitat and dietary relationships of the pygmy rabbit dournnl Kringe Marlrigernent 33 13amp42

I3css K J r 1993 Rocky Times in Rocky Mourltairl hcrtiorzril Park Niwot Colo University Prcss of Colorado

Flestcr F E 1991 The U S National Park Scrvicc experience with exotic species Nriturril Areri ~ Jourr~ril1 1 127-8

Hoerr W 1993 The concept of naturalness in environmental tliscoursc Natural Areas Journal 1329-32

Ilouston D B and Schrciner E 6 1995 Alien specie in national parks drawing lines in space and time Conservation Biology 9 204-9

IHouston D B Schrciner E 6and Moorhcad R R 1994 Mourltairl Goats in Olympic Nutionul Park Biologjl and Marlugernerlt of an Introduced Species U S National Park Scrvicc Scientific Monograph NPSINROLYMINRSM-9325

I-Iudson J (cd) 1993 From Bones to Bc~lzavior Etlznotrrchaeologicul and E~plcrimcntill Coiltri- butiorls ro rhe Intcrprc~tation of Fblrr~al liernairls Carbondalc Southern Illinois University Center for Archaeological Investigations Occasional Papcr No 21

T h e relevance of faunal analysis t o wildlife management 123

Hyland D C Tersak J M Adovasio J M and Sicgal M I 1990 Identification of thc speciesof origin of residual blood on lithic material Americntl Antiquity 55 104-12

Jameson R J Kenyon K W Johnson A M and Wight H M 1982 History and status of translocated sea otter populations in North America Wildlife Society Bulletin 10 100-7

Jameson R J Kenyon K W Jeffries S and VanBlaricom G R 1986 Status of a I ranslocatcd sea otter population and its habitat in Washington The Murrelet 67 84-7

Kay C E 1994 Aboriginal overkill the role of Native Americans in structur~ng western ecosystems Hurnnt~ Nrrrure 5 359-98

Kenyon K W 1969 The sea otter In the eastern Pacific Ocean Worth Amer~crrtz Firuna 68

Korn H 1994 Genetic demographic spatial environmental and catastrophic effects on the survival probability of small populations of mammals In Mitlimum Animrrl Popu1rrrior1s (ed H Remmert) Berlin Springer-Verlag pp 3)-49

Lcggc A J 1978 Archacozoology - or zooarchacology In Research Prohlems in Zoourchueology (eds D R Brothwell K D Thomas and J Clutton-Brock) Univcrsity of London Institute of Archaeology Occasional Publications No 3 pp 129-32

Leopold A S Cain S A Cottam C IM Gabrielson I N and Kimhall T L 1963 Wildlife management in the national parks Trunsuctions of the Twenty-Eighth North American Wildlife und Nutural Resoirrce~ Cd70nferencepp 2945

Loy T H 1993 The artifact as site an example of the biomolccular analysis of organic residues on prehistoric tools World Archaeology 25 44-63

Lyman R L 1988a Significance for wildlife management of the late Quaternary biogeography of mountain goats (Oreurnnos urnericunus) in the Pacific Northwest USA Arctic and Alpine Reseurch 20 13-23

Lyman R L 1988b Zoogeography of Oregon coast marine mammals the last 3000 years Murine Murnrnul Science 4 247-64

Lyman R L 1991 Late Quaternary biogeography of the pygmy rabbit (Brachylagirs iduhoensis) in eastern Washington Jo~lrnalof Mammalogy 72 11amp17

Lyman R L 1994a The Olympic Mountain goat controversy a different perspective Conservation Biology 8 898-901

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Lyman R L 199 Inaccurate data and the Olympic National Park mountain goat controversy North wect Science 69 234-8

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MacAskie I 1987 Updated status of the sea otter Enhydru lutris in Canada Cunudiun Field- Nuturcrlist 101 279-83

McDonald J N 1981 North American Bison Their Clascicution and Evolution Berkeley University of California Press

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124 R Lee E y n ~ u n

OBricn S J and Mays E 1991 Burcauc1atic mischief rccogni~ing enciangcrcd spccics and subspecies Scicncc 251 1 187--8

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Peck J M Miclucllc 14 C and Wright M 61987 Arc bison cuotic in the Wrangcll-St Elia National Park and Pre~cr~c Envivnnnlcrzrcrl iWrlrlcrgcineilr I I l(195 3

Quinn J Iand ifarrison S P 1988 Effects of habitat fragmentation anhi isolation on spci-is richness evidence from biogeographic patterns Oecologiii 75 132-40

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IcnsbcrgcrJ M and Barnosky A I) 1993 Short-term fluctuations in slniill mammals ot the lato Pleistocene from eastern Washington Wn iWorphologicrrl Chr~rigc irr C)iir~t~rricrr~ Mrrinnzcrls (J ~ i i~ t l~ Alzcrica (eds R A Martin and 4 Igt 131rnosky) Qambridgc Ciimbsidgc University Prcss pp 299-342

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Richards M B Sykcs B C and Hedges MF M 1995 Authenticating D N A extriicted Irorri ancient skeletal remains Iourr7crl of Arcl~aeologicril Scicncc 22 2W-9

Faohison ND 1978 Zooarchaeology its hiirto~y and dcvclopnirn~7r7rrcssrc Anthropologic~ul A~orirition Miscellancotrs Pripcr 2 1-2

Rocst A 1 1973 Subspecies of the sea ottcs Er7llytira Iiitris Io5 ii7gcles Coirnty Nnturcrl tiisrory Mltscirrn (ontributior~s in Scicncr 252

Rocst A 1 1979 A re-evaluation oi sea o t t c ~ taxonomy Paper prcscntcci at the Sea Ottcr Workshop Santa Barbara Cilifornia

Scheffer V B 1993 Tlie Olympic Mo~rntain goat conrrovcrs) a pcr5pectivc Coriscrvtrtior Biology 7 916-19

Sclicffcr V B and Wilkc F 1950 Validity of the subspccics E~llzydrii 1iltris r7crcic the southcrn s ix ottcr Joltvnill qf the Wcr~I7ir7gton Accrtlcuny 01 S~icr~cr402672

Simons I) R 1083 Interactions lsctween Calilornin corrdors and humans in prehistoric far wcstern North America In Vcrltcrrr Biology und ~tlunuget~ierlt (cds S KWilbur and J A Jackson) Berkeley University or Ciliforni~i Press pp 470-91

Steadman D xi 1995 Prehistoric cstinctions of Pacific isiarld birds biodiversity nlccts zooarchacology Scieilce 267 1 123-3 1

Trillrnich F and Limbcrger D 1985 Drastic effects of 871 Nirio on Ga1apigos piniiipcd Oecologiu 67 19-22

VanBlaricorn 6Rand Estcs 9 A (cds) 1988 l h p Bot~irnlrniij bcologj oj Seu O t r e r ~ (Berlin

Springer-Verliig

Wagner F 11 Forest RC i i l l R R Mcdullough 14 R Pcltoil M MPorTcr W F and Salwasscr 13 1995 WildlifePolicies in the U S Nirrionul Purks Washington DC Island Prcsi

Weisler M I and Gargett R H 1993 Pacific island avian cstinctions the taphonorny or human predation Arclzucoloyj in Occuniu 28 85-03

Wendell F E flardy R A An-rc5 1 land Burgi K T 1986 Tcrnporalanci spatial patterns in

114 R Lee Lyman

When managing small isolated populations we may not have many options other than to preserve those populations as they presently exist Yn some cases management efforts have failed and populations have been extirpated When this occurs one management alternative is to transplant individuals from other populations to the vacated rangc in an attempt to re-establish a population Zooarchaeological research can also plav an important role in these situations

Re-establishment of extirpated populations

Eighteenth- and nineteenth-century conimercial exploitation of sea otters (Enhydru lirtris) along the western coast of North America ultimately led to the extirpation of this species from much of its range (Kenyon 1969 Ogden 1993 Roest 1973) Attempts to re-establish populations along the middle latitudes of that coast took place in the 1970s and thus far have met with less than perfect success (Jameson et al 1982 1986 MacAskie 1987) The typically expressed reason for the limited success of these efforts is that ecosystems to which sea otters have been transplanted have heen modified too exter~sively by modern human industry to support them Y wonder i f however part of the reason might resicle in a factor that only zooarchaeological research car measure

Sea otter skulls collectetl from much of the once occupied eastern Pacific range of sea otters display various character gradients from the Aleutian Islands to southern California Some of these characters have been used to argue for or against the existence oi multiple subspecies of sea otter (Davis and Lidicker 1975 Roest 1973 1979 Scheffer and Wilke 1950 Wilson et al 1991) Final resolution of this debate will be critical to futtlrc wildlife management activities first because national legislation in the United States concerns species subspecies and populations (OWrien and Mayr 1901) and also beca~ase of conflict between one population (a possibly distinct subspecies) of sea otters and the modern fisheries industry (Wendell et al 1986) lhe significant aspect to all of this Is the transplanting of sea otters ILegally only forms (ecotypes or subspecies) native to ar arcs should be introduced there during efforts to re-establish populations This also m~aites biological sense as i t is likely that those native forms unique to an area are the product of millennia of evolution

The International Union for the Conservation of Nature and Natural Resources PUCN (recently renamed the World Conservation Un~on) has drafted guidelines Cor such -

reintroductions These read in part

An assessment should be made of the taxonomic status of individuals to ba re-introduced They should be of the same taxonomic unit (and ideally closely related genetically) as those which were extirpated An investigation of historical information about the loss and fate of individuals from the re-introduction area as well a mmolecuiar genetic studies should be undertaken in case of doubt Release stock idealiy should be closely-related genetically to the original native stock

(Anonymous 1992 2-31

Clearly a search of not only historic but also prehistoric information nnay be necessary The rapidly developing study of ancient DNA extracted from prehistoric skeletal tissues

The relevance of faunal anuljlsis to wildlife matlugement 115

(eg Richards et al 1993 1995 and references therein) seems to have great potential for contending with IUCN guidelines The technique is applicable to curated as wtll as newly acquired archaeological specimens It has however not yet been applied to Oregon coast sea otter remains

Given the lack of genetic material from eighteenth- and early nineteenth-century sea otters on the Oregon coast the problem becomes one of establishing the age of the character gradients displayed by eastern Pacific rim sea otters (see Wilson et al 1991 for a recent summary) and determining if those gradients have shifted spatially through time IIf those gradients have a prehistoric distribution in time andlor space similar to that documented using historic specimens a potential problem is raised If a particular form -represented by a particular character state -- of sea otter was present prehistorically in one location within the range and another form was present at another location then transplanting individuals from the former to the latter may be doomed from the start In such a case failure to re-establish a population could result on the one hand because the transplanted form is not well adapted to its new environment the extirpated form on the other hand is adapted to the environment of that location hence its difference in form

Several years ago I examined prehistoric sea otter remains recovered from archaeologi- cal sites on the coast of the state of Oregon - the middle latitudes of the historically documented range of sea otters (Lyman 1988b) The size of sea otter teeth recovered from those sites are more or less intermediate to those from Alaska (large) and those from southern California (small) but they also vary somewhat from this simple pattern Prehistoric Oregon sea otters match modern Alaskan sea otters in some characters in others they match modern California sea otters Precise details are unnecessary to the point I wish to make here which is Were the sea otters captured in Alaska and translocated to the Oregon coast in the 1970s of the proper phenotype (or genotype) Zooarchaeological evidence suggests they were not If this is a correct interpretation then perhaps those transplanted sea otters were doomed from the moment they were captured Whether or not my suggestion proves to be true or false and it is unclear which way the cards might fall given that we still have much to learn about this species (VanBlaricom arid Estes 1988) the significance of zooarchaeological data should be clear

Not only can zooarchaeological data inform us as to which form of a species might be the most appropriate one to transplant to an area such data have the potential to indicate the potential success of such transplant efforts Even if the appropriate form to transplant is clear this does not guarantee that the introduced form will survive in its new home This is precisely the argument offered by biologists working with sea otters on the Oregon coast The data substantiaiing their argument can be supplemented with zooarchaeological data An example of this is provided by the California condor (Gymnogyps cnlifornianus) of western North America a species presently near extinction (Emslie 1987) Zooarchaeo- logical and paleontological data suggest this species enjoyed a wider range during the late Pleistocene but that range shrank considerably with the onset of the postglacial Holocene epoch (Emslie 1987 Simons 1983) Those data also suggest that this species was a carrion feeder and that the terminal Pleistocene extinction of many genera of large mammals probably contributed to the decline in frequency and range of the California condor (Emslie 1987) Finally paleoecological data indicate attempts to reintroduce this species to portions of its prehistoric range would probably fail because of major differences

116 R Lee Lytriurz

between late Holocene floral and faunal communities and late Pleistocene commaanities (Emslie 1987)

In the absence of historic data or even with it (see below) zooarchaeological infor- mation may be critical to ensuring that the correct forms of animals are transplanted The wrong forms may be i l l suited to their new habitats or they may be well adapted and pro- liferate in their new homes Of course proliferation of artificially introduced species may on the one hand be beneficial and what is desired On the other hand as H think many of us are aware it may not be desired And the latter brings us to another kind of use that can be made of zooarchaeological data when wildlife management is the issue

Identifying exotic animals

Recalling the TUCNs guidelines for reintroductions if a particular form of organism (ecotype subspecies or species) is exotic to a park or wildlife preserve it map be necessary to remove it from the area in order to preserve the natural status of the ecosystem and the biota of the area Thus we must have a solid definition of exotic ipecieThe United Stttes National Park Service (NPS) defines an exotic species as one that occurs in a given place ts a result of direct or indirect deliberate or accidental actions by humans (not including de- liberate reintroductions) (Hester 1991 127) In contrast native species are those which presently occur or once did occur prior to sonic human influence in a given place area or region as the result of ecological processes that operate and have operatcd without signifi cant direct or indirect deliberate or accidental alteration by humans (NPS documents cited in Lyman 1988a 21 see also Wagner et al 1995) Clearly then if wildlife managcrs are to manage native ecosystems variously referred to as pristine ecosystems or wilder- ness -by exclusion or removal of exotic species and reintroduction of artiticially extirpated native species then they must have that baseline list of native species In the United Stares that list is derived from the earliest historical documents for the area included within a national park (eg Houston and Schreiner 1995 Lgteopold et al 1963see also Hoerr 1993) The ethnocentricity of such a procedure is as might be imagined unavoidable from a przc tical standpoint More importantly historical documents arc sometimes incomplete and are sometimes simply inaccurate Ihose written documents can of course be sup plemented with another kind of document - the zooarchaeological record Here I men tion two examples of such supplementation

Bison (Bison bison) were artificially introduced to a portion of the state of Alaska that was to subsequently become Wrangell-St Elias National Park and Preserve (Peek et al 1987) By definition these introduced bison should be considered exotic and are so corl-sidered by the NPS (Houston and Schreiner 1995) However zooarchaeological evidence indicates that bison were present in the area until at least 500 years ago or slightly later (reviewed in Peek et al 1987) Historic records suggest bison wcre not prcsent when the first white men visited the area in the middle and late nineteenth ccntury but non- fossilized bison skulls were collected from the arca early in the twentieth century (Peek et al 1987) Given the paucity of late Holocene zooarchaeological data in thc area and the unknown nature of the relevant fossilization processes it is unclear if bison were present in Alaska between about 450 years ago and the carly twentieth century (McDonaald 1981)

The relevance of faunal analyl to wrldllfe rnanugerrzent 117

The NPS considers the bison of Wrangell-St Elias National Park to be exotic due to the lack of evidence that bison were present when the tirst white explorers passed through the area (Houston and Schreiner 1995) As well NPS biologists note that the introduced forrn of bison was of an incorrect - non-native -genetic stock (a distinct subspecies or ecotype) given current beliefs about bison taxonomy For this reason alone perhaps the extant bison of Wrangell-St Elias should be removed However the zooarchaeological record is so poorly known that one cannot conclusively demonstrate that native bison were locally extinct by the beginning of the twentieth century This in turn raises the possibility that extant bison are hybrids of native and introduced genetic stocks Whether or not this possibility is in fact reality is irrelevant What is critical here is that only more zooarchaeological and paleontological research will establish first the timing of local extirpation of native Alaskan bison populations and second if all Alaskan populations were extinct when transplanting occurred in the middle of the twentieth century Once the facts are determined we will have strong bases for making a decision regarding the ultimate fate of the bison presently living in Wrangell-St Elias National Park and Preserve

I am much more familiar - and intimately involved -with a virtually identical situation that exists in western Washington state Here mountain goats (Oreamnos urnericanlls) were introduced in the 1920s to the Olympic Mountains A decade later in 1938 these mountains were designated Olympic National Park The present position of the NPS is that the mountain goats are exotic and plans are in mid-1995 under way to eradicate them The plan is controversial (Anunsen and Anunsen 1993 Scheffer 1993) as is the evidence mustered to support it (compare Houston et al 119941 with Lyman 11991a 1995a 1995b1) This is not the place to review that evidence in detail (see references cited) It is relevant however to note the significance of zooarchaeological evidence for final resolution of the controversy

The historical evidence concerning the pre-1920s presence of mountain goats in the Olympic Mountains is ambiguous Some documents indicate goats were absent while others indicate goats were present (Lyman 1994a 1995b) In fact the only way to demonstrate that mountain goats were present or absent from the Olympic Mountains prior to their 1920s introduction requires paleontological andor zooarchaeological data At present there are no paleontological data of any kind for the area and the zooarchaeological record is woefully inadequate and thus cannot serve as a basis for a management decision (Lyman 1995a) Park biologists note that the purpose of national parks in the United States is not to attempt to recreate late-Pleistocene or early Holocene biotas [The NPS] is primarily concerned with historic post-Columbian specres distri- butions (Houston and Schreiner 1995 207) The facts remain however the zooarchaeo- logical record is poorly known and the historic record is internally contradictory

From the perspective of dynamic biogeographic histories provided by zooarchaeology one thing is abundantly clear the definitions of native and exotic species used by the NPS to implement its policies are contradictory Exotic species occur in a given place as a result of actions by humans whereas native species are those which presently occur or once did occ~crprior to some human influence in a place as the result of natural ecological processes The italicized phrase is where the contradiction resides because it indicates that i f for example mountain goats occurred at any tirne in the past in the Olympic Mountains

118 R Lee Lyman

- or bison in Wrangell-St Elias - then by definition the extant individuals represcl~i a native species But the introduced species are also by definition exotic having been transplanted from northwestern Canada and southeastern Alaska Olympic Nationai Parks mountain goats are of a different genetic stock and thus are a form distinct fromi mountain goats that may have been in the Olympic Mountains 200 years ago Rccogr-iizing this contradiction some years ago Y argued that if the prehistoric remains of rnonirrtain goats should be found in the Olympic Mountains then the NPS should rethink policy issues (Lyman I9RWa 22) Some NPS biologists have only recently come to the same conclusion (Houston and Schreiner 1995) And given the discussions above of appropriata forms (ecotypes subspecies species) and the benefit of hindsight I would argue that the policies and definitions of key terms should also be reworded to reflect the reality of wildlife management practice (Anonymous 1992 OBrien and Mayr 1991) Thus given their perspective of time depth zooarchaeologists not only can contribute significant paleobiological information to wildlife management concerns but perhaps they can also help write workable policies (see Wagner et al 1995 for other examples) Paleobiological information provided by zooarchaelogical research may also help wildlife managers plals for the future as the final example demonstrates

Biological preserves and planning for the future

Thus far the examples of zooarchaeology applied to wildlife management concerns have focused on the use of information on prehistoric faunal dynamics to inform management decisions That same kind of information can also be used to make broad predictions about the future effects of certain kinds of decisions For instance as alluded to in the first example presented above the spatial boundaries of parks and wildlife preserves may Piavc significant implications for the long-term survival of populations found withirl those boundaries In an important contribution to the role of planning for the future on tllc trasis of the past Graham (1988) notes that the boundaries of many parks and preserves are defined based on modern climatic and environmental conditions However tkrose conditions as we all know change and the probability of significant change increases as the length of time considered increases Graham underscores the [act that much planning lo preserve terrestrial biological diversity operates under the assumption that biotas or communities of organisms tend to respond to environmental change as intact units The

paleobiological record indicates however that individual species respond to environ- mental changes by migrating in different directions at different rates during different times (Graham 1988 392) Such individualistic responses to environmental change mean that the biological preserves of today are artifacts of the time when they were identified and created Thus Graham (1988 393) argues that constructing future reserves as habitat islands without avenues of migration [corridors] would probably result in [extirpation of bounded populations of managed species]

Graham (1988) thus suggests that as in the example of the Great Basins mountaintop mammals migration corridors are critical to organism survival and that preserves must be located on the landscape in such a rnanner as to be conducive to inter-areal migration I suggest following my earlier discussion that knowing where ihose corridors were located

The relevance of faunal analjgtsis to wildlife management 119

in the past surely will help in locating future preserves And as Graham and Graham (1994) indicate detailed knowledge of the historical and ecological biogeography of a taxon could prove critical to reconstructing appropriate habitats for reintroduced populations of the taxon Such knowledge can often come only from the fossil record

Extinction -of populations of species of lineages - is a natural fact of life (eg Raup 1984) But ignorance resulted in the loss of the passenger pigeon (Ectopistes migratorius) from North America the Great Auk (Pingninns impennis) from the North Atlantic and the dodo (Raphus cuci~llatus) from Mauritius in the Indian Ocean Our often stumbling attempts at wildlife management may due to ignorance ultimately lead to other such losses (eg Graham 1988 Hess 1993 Kay 1994) Ihe international move to maintain biological diversity - not just the taxonomic richness of biotic communities but the particular structure of those communities - seems to be in the correct direction if we are to maintain some quality of life and wilderness Understanding the loss of diversity often demands zooarchaeological and related data (eg Culotta 1995 Steadman 1995) and will help us avoid future inadvertent and unintentional losses Diamonds (1984) Rosetta Stone of using knowledge of modern extinctions to understand prehistoric extinctions could and perhaps should be turned on its head

Discussion

Nearly eighty years ago archaeologist W J Wintemberg (1919) noted how zooarchaeo- logical research could contribute to zoological knowledge and indirectly wildlife management His comments were repeated and elaborated thirty years later by biologist R M Gilmore (1949) Their remarks have largely gone unheeded perhaps because like Legge (1978 129 130) many of us see our job as aiming to contribute to the sum of the archaeological knowledge and that bone data are archaeological data I agree with Legge but also with Wintemberg and Gilmore we should be more than just consumers of wildlife data (cf Will 1982)

The potential topics under the broad umbrella of wildlife management to which zooarchaeological research might contribute are numerous I have touched on only a fevv of the more readily identifiable ones above There are to be sure others of some significance For example the modern fisheries industry of the eastern Pacific Ocean has been disrupted several times in the recent past by major changes in atmospheric circulation patterns causing alteration of ocean currents These events - variously referred to as El Nifio or Southern Oscillation events -appear to have some periodicity (Rasmusson 1985) but the historic record is of limited utility in plotting the precise nature of that periodicity These events clearly influence much of the oceanic biota (eg Arnold and Tissot 1993 Trillmich and Limberger 1985) Given sufficient temporal resolution the zooarchaeologi- cal record thus may eventually provide the time depth necessary to clarify the periodicity and magnitude of El Nifio events and thus allow more accurate predictions of future events

Zooarchaeological research has also shown that humans have long had a hand in modifying ecosystems Based on his extensive research on the prehistoric extinctions of Pacific Island bird populations and species Steadman (1995 1130) concludes that humans

tilay find Yt difficult to manage scarce resources in a sustainable way That humans contributed to these extinctions seems clear (cg Weisler and Cargett 1993) and thus Steadmans conclusion has the proper ominous tone But zooarchaeological research cair also perhaps help us out of such situations The wapiti (Cervns elaphus) of Yellowstone National Park in the United States are an example Herc nearly 60000 wapiti are managed and to a significant degree protected from human predation However historic and zooarchaeological research indicates that this population is a function of modern wildlife management historic records virtually vithout exception fail to mention wapiti i l l

the area and zooarchaeological data indicate this large ungulate was very riirc there during the last several thousand years (Kay 1994) Clearly if the Yellowstone ecosSsten1 is to be managed and maintained as a pristine wilderness in perpetuity (a laudable but in reality unattainable goal see Wager et al 1995) things must change before the ecosystem is irreparably altered A virtually identical situation exists in Rocky Mountain National Park sotile 550 km southwest of Yellowstone (Hcss 1993) where again zooarchaeologicai and other data from prehistoric times might help rcsolve debates

The few examples of how zooarchaeological research can be applied to t i ioder~~ problems I have outlined are rather specific to a place andlor to a taxon I do not perccive that to bc a weakness of an applied zooarchaeology but rather as an intellectual challenge Broadening the scope of zooarchaeology in the manner in which 1 have discussed will not be easy but i t will be beneficial to our Suture not only from the perspective of contributing to ensuring the preservation of biological diversity for future generations but also as 1conclude below from the perspective of our discipline which tilight otherwise become increasingly perceived as the pursuit of esoteric knotvledge of little practical use

One might argue that the discussion here could be expanded by suggesting that the detection of animal residues such as blood and hair (eg Loy 1993) could also providr information on the taxonomic identity of species native to the locale where the tools were collected (eg Hyland et al 1990) I doubt however that such a study would convince wildlife managers or all zooarchaeologists that a species was native given that stone tools were often transported significant distances by their tiiakers (eg Meltzer 1995) 1 beiievc multiple instances of such residues from multiple sites would be necessary to build 21

strong argument A partial skeleton or selected isolated bones and teeth of the species in que5tion recovered from a s~ngle ~ t e uould probably be muclr more conblnclng (Lyman 1994b)

I have attempted here to describe the structure of an applied zooarchaeology This is not a plea to legitimize what some of us do for fun andlor a living Rather i t is an attempt to broaden the scope of zooarchaeological research 1 am all for research for the sake of research if there is a problem in need of solving or a question in need of an answer But B am also concerned with the future of zooarchaeology Too many of our new doctoral str~dcnts are finding that the job market is closed and too marly of us arc finding it increasingly difficult to secure funding for research Perhaps a developed applied zooarchaeology wil

The relevance o f faunal analysis t o wildlife rnanagement 121

provide a new job market and new sources of funding It seems self-evident to me that if a national park for example has a staff archaeologist who deals with the archaeological resources on park lands and has a staff biologist who deals with biological resources on park lands then a marriage of the two would benefit both However given the increasing specialization of both archaeology and biology a viable marriage may not be possible Ihc obvious alternative is a park staff zooarchaeologist who has a deep knowledge ancl understanding of both disciplines and has been trained to integrate the two

New sources of funding may also become available if we can convince the wildlife management community that because its decisions and policies are typically aimed at tht future knowing something of the past can result in wise decisions That knowledge will of course come with a price tag The cost we may encounter is that members of the wildlife management community -while they may well have a solid knowledge of the time depth of evolutionary processes- often tend to view biotas as static ever enduring entities That is biotas have no past or at least their history is insignificant because they are being managed for the future While we certainly cannot predict exactly when the next ice age or glacial period will begin we can I think argue convincingly that climates and environments will change and knowing this we can use the prehistoric record to test our predictions about how certain kinds of changes may affect biotas of the future And confirmed - or even rejected or falsified -predictions would be the strong selling point Federal land-managing agencies whose charge includes wildlife management might well pay a zooarchaeologist to help them make wise management decisions The other selling point is that a less desirable kind of price -ecosystem destruction and loss of biodiversity -might otherwise have to be paid

Acknowledgements

I thank Donald K Grayson Adrian Lister Michael J OBrien and Ken Ihornas for comments on earlier versions of the manuscript

Departtnent o f Anthropology University o f Missouri-Columbia

References

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Anunscn C S and Anunsen R 1993 Rcsponsc to Sclleffer Conservation Biology 7 054-7

Arnold J E andTissot B N 1993 Measurcmcnt of significant rnarine palcotcmperature variation using black abalone shells from prehistoric middens Quaternurj~ Iesearch 39 3901

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Culotta E 1995 Many suspects to blame in Madagascar cxtinctions Science 268 1568-9

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Emslic S D 1987 Age and diet of fossil California condors in Grand Canyon Arizona Science 237 768-70

Fisller J W Jr 1995 Bone surface modifications in 7ooarchaeology Journal ofArchacologica1 Method rind Theory 2 7-68

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T h e relevance of faunal analysis t o wildlife management 123

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McDonald J N 1981 North American Bison Their Clascicution and Evolution Berkeley University of California Press

Mcltzcr D J 1995 Clocking the first Americans Annual Review of Anthropology 24 71-45

124 R Lee E y n ~ u n

OBricn S J and Mays E 1991 Burcauc1atic mischief rccogni~ing enciangcrcd spccics and subspecies Scicncc 251 1 187--8

Ogden A 1933 Russian sea ottcr anal seal hunting on the Califorilia coast Caij(~rrrirrNisroricci1 Qmcrrrerly 12 29-5 1

Olscn S I and Olscn I W 1981 A conin-icnt on nomcnclaturc in faunal studies iui~cric~un Antiquity 46 192-4

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Quinn J Iand ifarrison S P 1988 Effects of habitat fragmentation anhi isolation on spci-is richness evidence from biogeographic patterns Oecologiii 75 132-40

Rasmusson E M 1985 El Nino and asiations in climate Atzcricr~z Scic3nritt 73 168-77

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IcnsbcrgcrJ M and Barnosky A I) 1993 Short-term fluctuations in slniill mammals ot the lato Pleistocene from eastern Washington Wn iWorphologicrrl Chr~rigc irr C)iir~t~rricrr~ Mrrinnzcrls (J ~ i i~ t l~ Alzcrica (eds R A Martin and 4 Igt 131rnosky) Qambridgc Ciimbsidgc University Prcss pp 299-342

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Faohison ND 1978 Zooarchaeology its hiirto~y and dcvclopnirn~7r7rrcssrc Anthropologic~ul A~orirition Miscellancotrs Pripcr 2 1-2

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Sclicffcr V B and Wilkc F 1950 Validity of the subspccics E~llzydrii 1iltris r7crcic the southcrn s ix ottcr Joltvnill qf the Wcr~I7ir7gton Accrtlcuny 01 S~icr~cr402672

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Steadman D xi 1995 Prehistoric cstinctions of Pacific isiarld birds biodiversity nlccts zooarchacology Scieilce 267 1 123-3 1

Trillrnich F and Limbcrger D 1985 Drastic effects of 871 Nirio on Ga1apigos piniiipcd Oecologiu 67 19-22

VanBlaricorn 6Rand Estcs 9 A (cds) 1988 l h p Bot~irnlrniij bcologj oj Seu O t r e r ~ (Berlin

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Wagner F 11 Forest RC i i l l R R Mcdullough 14 R Pcltoil M MPorTcr W F and Salwasscr 13 1995 WildlifePolicies in the U S Nirrionul Purks Washington DC Island Prcsi

Weisler M I and Gargett R H 1993 Pacific island avian cstinctions the taphonorny or human predation Arclzucoloyj in Occuniu 28 85-03

Wendell F E flardy R A An-rc5 1 land Burgi K T 1986 Tcrnporalanci spatial patterns in

The relevance of faunal anuljlsis to wildlife matlugement 115

(eg Richards et al 1993 1995 and references therein) seems to have great potential for contending with IUCN guidelines The technique is applicable to curated as wtll as newly acquired archaeological specimens It has however not yet been applied to Oregon coast sea otter remains

Given the lack of genetic material from eighteenth- and early nineteenth-century sea otters on the Oregon coast the problem becomes one of establishing the age of the character gradients displayed by eastern Pacific rim sea otters (see Wilson et al 1991 for a recent summary) and determining if those gradients have shifted spatially through time IIf those gradients have a prehistoric distribution in time andlor space similar to that documented using historic specimens a potential problem is raised If a particular form -represented by a particular character state -- of sea otter was present prehistorically in one location within the range and another form was present at another location then transplanting individuals from the former to the latter may be doomed from the start In such a case failure to re-establish a population could result on the one hand because the transplanted form is not well adapted to its new environment the extirpated form on the other hand is adapted to the environment of that location hence its difference in form

Several years ago I examined prehistoric sea otter remains recovered from archaeologi- cal sites on the coast of the state of Oregon - the middle latitudes of the historically documented range of sea otters (Lyman 1988b) The size of sea otter teeth recovered from those sites are more or less intermediate to those from Alaska (large) and those from southern California (small) but they also vary somewhat from this simple pattern Prehistoric Oregon sea otters match modern Alaskan sea otters in some characters in others they match modern California sea otters Precise details are unnecessary to the point I wish to make here which is Were the sea otters captured in Alaska and translocated to the Oregon coast in the 1970s of the proper phenotype (or genotype) Zooarchaeological evidence suggests they were not If this is a correct interpretation then perhaps those transplanted sea otters were doomed from the moment they were captured Whether or not my suggestion proves to be true or false and it is unclear which way the cards might fall given that we still have much to learn about this species (VanBlaricom arid Estes 1988) the significance of zooarchaeological data should be clear

Not only can zooarchaeological data inform us as to which form of a species might be the most appropriate one to transplant to an area such data have the potential to indicate the potential success of such transplant efforts Even if the appropriate form to transplant is clear this does not guarantee that the introduced form will survive in its new home This is precisely the argument offered by biologists working with sea otters on the Oregon coast The data substantiaiing their argument can be supplemented with zooarchaeological data An example of this is provided by the California condor (Gymnogyps cnlifornianus) of western North America a species presently near extinction (Emslie 1987) Zooarchaeo- logical and paleontological data suggest this species enjoyed a wider range during the late Pleistocene but that range shrank considerably with the onset of the postglacial Holocene epoch (Emslie 1987 Simons 1983) Those data also suggest that this species was a carrion feeder and that the terminal Pleistocene extinction of many genera of large mammals probably contributed to the decline in frequency and range of the California condor (Emslie 1987) Finally paleoecological data indicate attempts to reintroduce this species to portions of its prehistoric range would probably fail because of major differences

116 R Lee Lytriurz

between late Holocene floral and faunal communities and late Pleistocene commaanities (Emslie 1987)

In the absence of historic data or even with it (see below) zooarchaeological infor- mation may be critical to ensuring that the correct forms of animals are transplanted The wrong forms may be i l l suited to their new habitats or they may be well adapted and pro- liferate in their new homes Of course proliferation of artificially introduced species may on the one hand be beneficial and what is desired On the other hand as H think many of us are aware it may not be desired And the latter brings us to another kind of use that can be made of zooarchaeological data when wildlife management is the issue

Identifying exotic animals

Recalling the TUCNs guidelines for reintroductions if a particular form of organism (ecotype subspecies or species) is exotic to a park or wildlife preserve it map be necessary to remove it from the area in order to preserve the natural status of the ecosystem and the biota of the area Thus we must have a solid definition of exotic ipecieThe United Stttes National Park Service (NPS) defines an exotic species as one that occurs in a given place ts a result of direct or indirect deliberate or accidental actions by humans (not including de- liberate reintroductions) (Hester 1991 127) In contrast native species are those which presently occur or once did occur prior to sonic human influence in a given place area or region as the result of ecological processes that operate and have operatcd without signifi cant direct or indirect deliberate or accidental alteration by humans (NPS documents cited in Lyman 1988a 21 see also Wagner et al 1995) Clearly then if wildlife managcrs are to manage native ecosystems variously referred to as pristine ecosystems or wilder- ness -by exclusion or removal of exotic species and reintroduction of artiticially extirpated native species then they must have that baseline list of native species In the United Stares that list is derived from the earliest historical documents for the area included within a national park (eg Houston and Schreiner 1995 Lgteopold et al 1963see also Hoerr 1993) The ethnocentricity of such a procedure is as might be imagined unavoidable from a przc tical standpoint More importantly historical documents arc sometimes incomplete and are sometimes simply inaccurate Ihose written documents can of course be sup plemented with another kind of document - the zooarchaeological record Here I men tion two examples of such supplementation

Bison (Bison bison) were artificially introduced to a portion of the state of Alaska that was to subsequently become Wrangell-St Elias National Park and Preserve (Peek et al 1987) By definition these introduced bison should be considered exotic and are so corl-sidered by the NPS (Houston and Schreiner 1995) However zooarchaeological evidence indicates that bison were present in the area until at least 500 years ago or slightly later (reviewed in Peek et al 1987) Historic records suggest bison wcre not prcsent when the first white men visited the area in the middle and late nineteenth ccntury but non- fossilized bison skulls were collected from the arca early in the twentieth century (Peek et al 1987) Given the paucity of late Holocene zooarchaeological data in thc area and the unknown nature of the relevant fossilization processes it is unclear if bison were present in Alaska between about 450 years ago and the carly twentieth century (McDonaald 1981)

The relevance of faunal analyl to wrldllfe rnanugerrzent 117

The NPS considers the bison of Wrangell-St Elias National Park to be exotic due to the lack of evidence that bison were present when the tirst white explorers passed through the area (Houston and Schreiner 1995) As well NPS biologists note that the introduced forrn of bison was of an incorrect - non-native -genetic stock (a distinct subspecies or ecotype) given current beliefs about bison taxonomy For this reason alone perhaps the extant bison of Wrangell-St Elias should be removed However the zooarchaeological record is so poorly known that one cannot conclusively demonstrate that native bison were locally extinct by the beginning of the twentieth century This in turn raises the possibility that extant bison are hybrids of native and introduced genetic stocks Whether or not this possibility is in fact reality is irrelevant What is critical here is that only more zooarchaeological and paleontological research will establish first the timing of local extirpation of native Alaskan bison populations and second if all Alaskan populations were extinct when transplanting occurred in the middle of the twentieth century Once the facts are determined we will have strong bases for making a decision regarding the ultimate fate of the bison presently living in Wrangell-St Elias National Park and Preserve

I am much more familiar - and intimately involved -with a virtually identical situation that exists in western Washington state Here mountain goats (Oreamnos urnericanlls) were introduced in the 1920s to the Olympic Mountains A decade later in 1938 these mountains were designated Olympic National Park The present position of the NPS is that the mountain goats are exotic and plans are in mid-1995 under way to eradicate them The plan is controversial (Anunsen and Anunsen 1993 Scheffer 1993) as is the evidence mustered to support it (compare Houston et al 119941 with Lyman 11991a 1995a 1995b1) This is not the place to review that evidence in detail (see references cited) It is relevant however to note the significance of zooarchaeological evidence for final resolution of the controversy

The historical evidence concerning the pre-1920s presence of mountain goats in the Olympic Mountains is ambiguous Some documents indicate goats were absent while others indicate goats were present (Lyman 1994a 1995b) In fact the only way to demonstrate that mountain goats were present or absent from the Olympic Mountains prior to their 1920s introduction requires paleontological andor zooarchaeological data At present there are no paleontological data of any kind for the area and the zooarchaeological record is woefully inadequate and thus cannot serve as a basis for a management decision (Lyman 1995a) Park biologists note that the purpose of national parks in the United States is not to attempt to recreate late-Pleistocene or early Holocene biotas [The NPS] is primarily concerned with historic post-Columbian specres distri- butions (Houston and Schreiner 1995 207) The facts remain however the zooarchaeo- logical record is poorly known and the historic record is internally contradictory

From the perspective of dynamic biogeographic histories provided by zooarchaeology one thing is abundantly clear the definitions of native and exotic species used by the NPS to implement its policies are contradictory Exotic species occur in a given place as a result of actions by humans whereas native species are those which presently occur or once did occ~crprior to some human influence in a place as the result of natural ecological processes The italicized phrase is where the contradiction resides because it indicates that i f for example mountain goats occurred at any tirne in the past in the Olympic Mountains

118 R Lee Lyman

- or bison in Wrangell-St Elias - then by definition the extant individuals represcl~i a native species But the introduced species are also by definition exotic having been transplanted from northwestern Canada and southeastern Alaska Olympic Nationai Parks mountain goats are of a different genetic stock and thus are a form distinct fromi mountain goats that may have been in the Olympic Mountains 200 years ago Rccogr-iizing this contradiction some years ago Y argued that if the prehistoric remains of rnonirrtain goats should be found in the Olympic Mountains then the NPS should rethink policy issues (Lyman I9RWa 22) Some NPS biologists have only recently come to the same conclusion (Houston and Schreiner 1995) And given the discussions above of appropriata forms (ecotypes subspecies species) and the benefit of hindsight I would argue that the policies and definitions of key terms should also be reworded to reflect the reality of wildlife management practice (Anonymous 1992 OBrien and Mayr 1991) Thus given their perspective of time depth zooarchaeologists not only can contribute significant paleobiological information to wildlife management concerns but perhaps they can also help write workable policies (see Wagner et al 1995 for other examples) Paleobiological information provided by zooarchaelogical research may also help wildlife managers plals for the future as the final example demonstrates

Biological preserves and planning for the future

Thus far the examples of zooarchaeology applied to wildlife management concerns have focused on the use of information on prehistoric faunal dynamics to inform management decisions That same kind of information can also be used to make broad predictions about the future effects of certain kinds of decisions For instance as alluded to in the first example presented above the spatial boundaries of parks and wildlife preserves may Piavc significant implications for the long-term survival of populations found withirl those boundaries In an important contribution to the role of planning for the future on tllc trasis of the past Graham (1988) notes that the boundaries of many parks and preserves are defined based on modern climatic and environmental conditions However tkrose conditions as we all know change and the probability of significant change increases as the length of time considered increases Graham underscores the [act that much planning lo preserve terrestrial biological diversity operates under the assumption that biotas or communities of organisms tend to respond to environmental change as intact units The

paleobiological record indicates however that individual species respond to environ- mental changes by migrating in different directions at different rates during different times (Graham 1988 392) Such individualistic responses to environmental change mean that the biological preserves of today are artifacts of the time when they were identified and created Thus Graham (1988 393) argues that constructing future reserves as habitat islands without avenues of migration [corridors] would probably result in [extirpation of bounded populations of managed species]

Graham (1988) thus suggests that as in the example of the Great Basins mountaintop mammals migration corridors are critical to organism survival and that preserves must be located on the landscape in such a rnanner as to be conducive to inter-areal migration I suggest following my earlier discussion that knowing where ihose corridors were located

The relevance of faunal analjgtsis to wildlife management 119

in the past surely will help in locating future preserves And as Graham and Graham (1994) indicate detailed knowledge of the historical and ecological biogeography of a taxon could prove critical to reconstructing appropriate habitats for reintroduced populations of the taxon Such knowledge can often come only from the fossil record

Extinction -of populations of species of lineages - is a natural fact of life (eg Raup 1984) But ignorance resulted in the loss of the passenger pigeon (Ectopistes migratorius) from North America the Great Auk (Pingninns impennis) from the North Atlantic and the dodo (Raphus cuci~llatus) from Mauritius in the Indian Ocean Our often stumbling attempts at wildlife management may due to ignorance ultimately lead to other such losses (eg Graham 1988 Hess 1993 Kay 1994) Ihe international move to maintain biological diversity - not just the taxonomic richness of biotic communities but the particular structure of those communities - seems to be in the correct direction if we are to maintain some quality of life and wilderness Understanding the loss of diversity often demands zooarchaeological and related data (eg Culotta 1995 Steadman 1995) and will help us avoid future inadvertent and unintentional losses Diamonds (1984) Rosetta Stone of using knowledge of modern extinctions to understand prehistoric extinctions could and perhaps should be turned on its head

Discussion

Nearly eighty years ago archaeologist W J Wintemberg (1919) noted how zooarchaeo- logical research could contribute to zoological knowledge and indirectly wildlife management His comments were repeated and elaborated thirty years later by biologist R M Gilmore (1949) Their remarks have largely gone unheeded perhaps because like Legge (1978 129 130) many of us see our job as aiming to contribute to the sum of the archaeological knowledge and that bone data are archaeological data I agree with Legge but also with Wintemberg and Gilmore we should be more than just consumers of wildlife data (cf Will 1982)

The potential topics under the broad umbrella of wildlife management to which zooarchaeological research might contribute are numerous I have touched on only a fevv of the more readily identifiable ones above There are to be sure others of some significance For example the modern fisheries industry of the eastern Pacific Ocean has been disrupted several times in the recent past by major changes in atmospheric circulation patterns causing alteration of ocean currents These events - variously referred to as El Nifio or Southern Oscillation events -appear to have some periodicity (Rasmusson 1985) but the historic record is of limited utility in plotting the precise nature of that periodicity These events clearly influence much of the oceanic biota (eg Arnold and Tissot 1993 Trillmich and Limberger 1985) Given sufficient temporal resolution the zooarchaeologi- cal record thus may eventually provide the time depth necessary to clarify the periodicity and magnitude of El Nifio events and thus allow more accurate predictions of future events

Zooarchaeological research has also shown that humans have long had a hand in modifying ecosystems Based on his extensive research on the prehistoric extinctions of Pacific Island bird populations and species Steadman (1995 1130) concludes that humans

tilay find Yt difficult to manage scarce resources in a sustainable way That humans contributed to these extinctions seems clear (cg Weisler and Cargett 1993) and thus Steadmans conclusion has the proper ominous tone But zooarchaeological research cair also perhaps help us out of such situations The wapiti (Cervns elaphus) of Yellowstone National Park in the United States are an example Herc nearly 60000 wapiti are managed and to a significant degree protected from human predation However historic and zooarchaeological research indicates that this population is a function of modern wildlife management historic records virtually vithout exception fail to mention wapiti i l l

the area and zooarchaeological data indicate this large ungulate was very riirc there during the last several thousand years (Kay 1994) Clearly if the Yellowstone ecosSsten1 is to be managed and maintained as a pristine wilderness in perpetuity (a laudable but in reality unattainable goal see Wager et al 1995) things must change before the ecosystem is irreparably altered A virtually identical situation exists in Rocky Mountain National Park sotile 550 km southwest of Yellowstone (Hcss 1993) where again zooarchaeologicai and other data from prehistoric times might help rcsolve debates

The few examples of how zooarchaeological research can be applied to t i ioder~~ problems I have outlined are rather specific to a place andlor to a taxon I do not perccive that to bc a weakness of an applied zooarchaeology but rather as an intellectual challenge Broadening the scope of zooarchaeology in the manner in which 1 have discussed will not be easy but i t will be beneficial to our Suture not only from the perspective of contributing to ensuring the preservation of biological diversity for future generations but also as 1conclude below from the perspective of our discipline which tilight otherwise become increasingly perceived as the pursuit of esoteric knotvledge of little practical use

One might argue that the discussion here could be expanded by suggesting that the detection of animal residues such as blood and hair (eg Loy 1993) could also providr information on the taxonomic identity of species native to the locale where the tools were collected (eg Hyland et al 1990) I doubt however that such a study would convince wildlife managers or all zooarchaeologists that a species was native given that stone tools were often transported significant distances by their tiiakers (eg Meltzer 1995) 1 beiievc multiple instances of such residues from multiple sites would be necessary to build 21

strong argument A partial skeleton or selected isolated bones and teeth of the species in que5tion recovered from a s~ngle ~ t e uould probably be muclr more conblnclng (Lyman 1994b)

I have attempted here to describe the structure of an applied zooarchaeology This is not a plea to legitimize what some of us do for fun andlor a living Rather i t is an attempt to broaden the scope of zooarchaeological research 1 am all for research for the sake of research if there is a problem in need of solving or a question in need of an answer But B am also concerned with the future of zooarchaeology Too many of our new doctoral str~dcnts are finding that the job market is closed and too marly of us arc finding it increasingly difficult to secure funding for research Perhaps a developed applied zooarchaeology wil

The relevance o f faunal analysis t o wildlife rnanagement 121

provide a new job market and new sources of funding It seems self-evident to me that if a national park for example has a staff archaeologist who deals with the archaeological resources on park lands and has a staff biologist who deals with biological resources on park lands then a marriage of the two would benefit both However given the increasing specialization of both archaeology and biology a viable marriage may not be possible Ihc obvious alternative is a park staff zooarchaeologist who has a deep knowledge ancl understanding of both disciplines and has been trained to integrate the two

New sources of funding may also become available if we can convince the wildlife management community that because its decisions and policies are typically aimed at tht future knowing something of the past can result in wise decisions That knowledge will of course come with a price tag The cost we may encounter is that members of the wildlife management community -while they may well have a solid knowledge of the time depth of evolutionary processes- often tend to view biotas as static ever enduring entities That is biotas have no past or at least their history is insignificant because they are being managed for the future While we certainly cannot predict exactly when the next ice age or glacial period will begin we can I think argue convincingly that climates and environments will change and knowing this we can use the prehistoric record to test our predictions about how certain kinds of changes may affect biotas of the future And confirmed - or even rejected or falsified -predictions would be the strong selling point Federal land-managing agencies whose charge includes wildlife management might well pay a zooarchaeologist to help them make wise management decisions The other selling point is that a less desirable kind of price -ecosystem destruction and loss of biodiversity -might otherwise have to be paid

Acknowledgements

I thank Donald K Grayson Adrian Lister Michael J OBrien and Ken Ihornas for comments on earlier versions of the manuscript

Departtnent o f Anthropology University o f Missouri-Columbia

References

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Anunscn C S and Anunsen R 1993 Rcsponsc to Sclleffer Conservation Biology 7 054-7

Arnold J E andTissot B N 1993 Measurcmcnt of significant rnarine palcotcmperature variation using black abalone shells from prehistoric middens Quaternurj~ Iesearch 39 3901

Brewer D J 1992 Zooarchacology method tlleory and goals In Archaeological Metl~od and Theory Vol 4 (ed M B Schiffer) Tucson University of Arizona Press pp 195-244

Brown J H 1971 Mammals on mountaintops nonequilibrium insular biogcograplly American Vr~turulist105 467-78

Brown J PI 1978 The theory of insular biogeography and the distribution of boreal mammals and birds In Intermountain Biogeogrur~hy A Synzposium (eds KT Harper and J L Rcvcal) Great Basin Natural Memoir 2 pp 209-28

Brown J H 1986 Two decades of interaction between the MacArthiu-Wilson model and tile complexities of mammalian distributions Biological Journal d t h c Lirlnearl Society 28 23151

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Fisller J W Jr 1995 Bone surface modifications in 7ooarchaeology Journal ofArchacologica1 Method rind Theory 2 7-68

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Graham RW 1988 The role of climatic change in the design of biological reserves tho palcoccological perspccti~rc for conser~ration biology Corltervution Biologjl 2 391-4

Graham RW and Graham M A 1994 Late Quaternary distribution of Marter in North America In Mcrrten~ Sublcs urtd Fislrcrs Biology and Conservation (eds S W Ruskirk 14 S Harestad M 6Raphael and RA Powell) Itllaca NY Corncll University Prcss pp 2 6 5 8

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Green J S and Flindcrs J T 1980b Habitat and dietary relationships of the pygmy rabbit dournnl Kringe Marlrigernent 33 13amp42

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Flestcr F E 1991 The U S National Park Scrvicc experience with exotic species Nriturril Areri ~ Jourr~ril1 1 127-8

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Ilouston D B and Schrciner E 6 1995 Alien specie in national parks drawing lines in space and time Conservation Biology 9 204-9

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I-Iudson J (cd) 1993 From Bones to Bc~lzavior Etlznotrrchaeologicul and E~plcrimcntill Coiltri- butiorls ro rhe Intcrprc~tation of Fblrr~al liernairls Carbondalc Southern Illinois University Center for Archaeological Investigations Occasional Papcr No 21

T h e relevance of faunal analysis t o wildlife management 123

Hyland D C Tersak J M Adovasio J M and Sicgal M I 1990 Identification of thc speciesof origin of residual blood on lithic material Americntl Antiquity 55 104-12

Jameson R J Kenyon K W Johnson A M and Wight H M 1982 History and status of translocated sea otter populations in North America Wildlife Society Bulletin 10 100-7

Jameson R J Kenyon K W Jeffries S and VanBlaricom G R 1986 Status of a I ranslocatcd sea otter population and its habitat in Washington The Murrelet 67 84-7

Kay C E 1994 Aboriginal overkill the role of Native Americans in structur~ng western ecosystems Hurnnt~ Nrrrure 5 359-98

Kenyon K W 1969 The sea otter In the eastern Pacific Ocean Worth Amer~crrtz Firuna 68

Korn H 1994 Genetic demographic spatial environmental and catastrophic effects on the survival probability of small populations of mammals In Mitlimum Animrrl Popu1rrrior1s (ed H Remmert) Berlin Springer-Verlag pp 3)-49

Lcggc A J 1978 Archacozoology - or zooarchacology In Research Prohlems in Zoourchueology (eds D R Brothwell K D Thomas and J Clutton-Brock) Univcrsity of London Institute of Archaeology Occasional Publications No 3 pp 129-32

Leopold A S Cain S A Cottam C IM Gabrielson I N and Kimhall T L 1963 Wildlife management in the national parks Trunsuctions of the Twenty-Eighth North American Wildlife und Nutural Resoirrce~ Cd70nferencepp 2945

Loy T H 1993 The artifact as site an example of the biomolccular analysis of organic residues on prehistoric tools World Archaeology 25 44-63

Lyman R L 1988a Significance for wildlife management of the late Quaternary biogeography of mountain goats (Oreurnnos urnericunus) in the Pacific Northwest USA Arctic and Alpine Reseurch 20 13-23

Lyman R L 1988b Zoogeography of Oregon coast marine mammals the last 3000 years Murine Murnrnul Science 4 247-64

Lyman R L 1991 Late Quaternary biogeography of the pygmy rabbit (Brachylagirs iduhoensis) in eastern Washington Jo~lrnalof Mammalogy 72 11amp17

Lyman R L 1994a The Olympic Mountain goat controversy a different perspective Conservation Biology 8 898-901

Lyman R L 1994b Vertebrute Taphonomy Cambridge Cambridge University Press

Lyman R L 1995a Determining when rare (zoo)archacological phenomena are truly absent Journal of Archueologicul Method unrl Theory 2 369-424

Lyman R L 199 Inaccurate data and the Olympic National Park mountain goat controversy North wect Science 69 234-8

McAllister K and Allen H 1993 Stutuc of the Pygmy Rabbit (Brachylagus idahoensis) in Wushington Olympia Washington Department of Wildlife Wildlife Management Division

McAllister K and Allen H 1994 Wushington Stute Draft Recovery Plun for the Pygmy Rabbit (Brachylagus idahocnsis) Olympia Washington Department of Wildlife Wildlife Management Division

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MacAskie I 1987 Updated status of the sea otter Enhydru lutris in Canada Cunudiun Field- Nuturcrlist 101 279-83

McDonald J N 1981 North American Bison Their Clascicution and Evolution Berkeley University of California Press

Mcltzcr D J 1995 Clocking the first Americans Annual Review of Anthropology 24 71-45

124 R Lee E y n ~ u n

OBricn S J and Mays E 1991 Burcauc1atic mischief rccogni~ing enciangcrcd spccics and subspecies Scicncc 251 1 187--8

Ogden A 1933 Russian sea ottcr anal seal hunting on the Califorilia coast Caij(~rrrirrNisroricci1 Qmcrrrerly 12 29-5 1

Olscn S I and Olscn I W 1981 A conin-icnt on nomcnclaturc in faunal studies iui~cric~un Antiquity 46 192-4

Peck J M Miclucllc 14 C and Wright M 61987 Arc bison cuotic in the Wrangcll-St Elia National Park and Pre~cr~c Envivnnnlcrzrcrl iWrlrlcrgcineilr I I l(195 3

Quinn J Iand ifarrison S P 1988 Effects of habitat fragmentation anhi isolation on spci-is richness evidence from biogeographic patterns Oecologiii 75 132-40

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Raup D M 1984 Death of spccics I n Errirrcrions (etl M 14 Nitccllti) dlhicago University of Chicago Press pp 1-19

IcnsbcrgcrJ M and Barnosky A I) 1993 Short-term fluctuations in slniill mammals ot the lato Pleistocene from eastern Washington Wn iWorphologicrrl Chr~rigc irr C)iir~t~rricrr~ Mrrinnzcrls (J ~ i i~ t l~ Alzcrica (eds R A Martin and 4 Igt 131rnosky) Qambridgc Ciimbsidgc University Prcss pp 299-342

Richards M Smallcy KSykcs R and Iledges BP 1993 iir-ch21eology and genetics anaiysirig DNA from skeletal remains Worlrl~lrchtrcolog~5 18-28

Richards M B Sykcs B C and Hedges MF M 1995 Authenticating D N A extriicted Irorri ancient skeletal remains Iourr7crl of Arcl~aeologicril Scicncc 22 2W-9

Faohison ND 1978 Zooarchaeology its hiirto~y and dcvclopnirn~7r7rrcssrc Anthropologic~ul A~orirition Miscellancotrs Pripcr 2 1-2

Rocst A 1 1973 Subspecies of the sea ottcs Er7llytira Iiitris Io5 ii7gcles Coirnty Nnturcrl tiisrory Mltscirrn (ontributior~s in Scicncr 252

Rocst A 1 1979 A re-evaluation oi sea o t t c ~ taxonomy Paper prcscntcci at the Sea Ottcr Workshop Santa Barbara Cilifornia

Scheffer V B 1993 Tlie Olympic Mo~rntain goat conrrovcrs) a pcr5pectivc Coriscrvtrtior Biology 7 916-19

Sclicffcr V B and Wilkc F 1950 Validity of the subspccics E~llzydrii 1iltris r7crcic the southcrn s ix ottcr Joltvnill qf the Wcr~I7ir7gton Accrtlcuny 01 S~icr~cr402672

Simons I) R 1083 Interactions lsctween Calilornin corrdors and humans in prehistoric far wcstern North America In Vcrltcrrr Biology und ~tlunuget~ierlt (cds S KWilbur and J A Jackson) Berkeley University or Ciliforni~i Press pp 470-91

Steadman D xi 1995 Prehistoric cstinctions of Pacific isiarld birds biodiversity nlccts zooarchacology Scieilce 267 1 123-3 1

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Springer-Verliig

Wagner F 11 Forest RC i i l l R R Mcdullough 14 R Pcltoil M MPorTcr W F and Salwasscr 13 1995 WildlifePolicies in the U S Nirrionul Purks Washington DC Island Prcsi

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116 R Lee Lytriurz

between late Holocene floral and faunal communities and late Pleistocene commaanities (Emslie 1987)

In the absence of historic data or even with it (see below) zooarchaeological infor- mation may be critical to ensuring that the correct forms of animals are transplanted The wrong forms may be i l l suited to their new habitats or they may be well adapted and pro- liferate in their new homes Of course proliferation of artificially introduced species may on the one hand be beneficial and what is desired On the other hand as H think many of us are aware it may not be desired And the latter brings us to another kind of use that can be made of zooarchaeological data when wildlife management is the issue

Identifying exotic animals

Recalling the TUCNs guidelines for reintroductions if a particular form of organism (ecotype subspecies or species) is exotic to a park or wildlife preserve it map be necessary to remove it from the area in order to preserve the natural status of the ecosystem and the biota of the area Thus we must have a solid definition of exotic ipecieThe United Stttes National Park Service (NPS) defines an exotic species as one that occurs in a given place ts a result of direct or indirect deliberate or accidental actions by humans (not including de- liberate reintroductions) (Hester 1991 127) In contrast native species are those which presently occur or once did occur prior to sonic human influence in a given place area or region as the result of ecological processes that operate and have operatcd without signifi cant direct or indirect deliberate or accidental alteration by humans (NPS documents cited in Lyman 1988a 21 see also Wagner et al 1995) Clearly then if wildlife managcrs are to manage native ecosystems variously referred to as pristine ecosystems or wilder- ness -by exclusion or removal of exotic species and reintroduction of artiticially extirpated native species then they must have that baseline list of native species In the United Stares that list is derived from the earliest historical documents for the area included within a national park (eg Houston and Schreiner 1995 Lgteopold et al 1963see also Hoerr 1993) The ethnocentricity of such a procedure is as might be imagined unavoidable from a przc tical standpoint More importantly historical documents arc sometimes incomplete and are sometimes simply inaccurate Ihose written documents can of course be sup plemented with another kind of document - the zooarchaeological record Here I men tion two examples of such supplementation

Bison (Bison bison) were artificially introduced to a portion of the state of Alaska that was to subsequently become Wrangell-St Elias National Park and Preserve (Peek et al 1987) By definition these introduced bison should be considered exotic and are so corl-sidered by the NPS (Houston and Schreiner 1995) However zooarchaeological evidence indicates that bison were present in the area until at least 500 years ago or slightly later (reviewed in Peek et al 1987) Historic records suggest bison wcre not prcsent when the first white men visited the area in the middle and late nineteenth ccntury but non- fossilized bison skulls were collected from the arca early in the twentieth century (Peek et al 1987) Given the paucity of late Holocene zooarchaeological data in thc area and the unknown nature of the relevant fossilization processes it is unclear if bison were present in Alaska between about 450 years ago and the carly twentieth century (McDonaald 1981)

The relevance of faunal analyl to wrldllfe rnanugerrzent 117

The NPS considers the bison of Wrangell-St Elias National Park to be exotic due to the lack of evidence that bison were present when the tirst white explorers passed through the area (Houston and Schreiner 1995) As well NPS biologists note that the introduced forrn of bison was of an incorrect - non-native -genetic stock (a distinct subspecies or ecotype) given current beliefs about bison taxonomy For this reason alone perhaps the extant bison of Wrangell-St Elias should be removed However the zooarchaeological record is so poorly known that one cannot conclusively demonstrate that native bison were locally extinct by the beginning of the twentieth century This in turn raises the possibility that extant bison are hybrids of native and introduced genetic stocks Whether or not this possibility is in fact reality is irrelevant What is critical here is that only more zooarchaeological and paleontological research will establish first the timing of local extirpation of native Alaskan bison populations and second if all Alaskan populations were extinct when transplanting occurred in the middle of the twentieth century Once the facts are determined we will have strong bases for making a decision regarding the ultimate fate of the bison presently living in Wrangell-St Elias National Park and Preserve

I am much more familiar - and intimately involved -with a virtually identical situation that exists in western Washington state Here mountain goats (Oreamnos urnericanlls) were introduced in the 1920s to the Olympic Mountains A decade later in 1938 these mountains were designated Olympic National Park The present position of the NPS is that the mountain goats are exotic and plans are in mid-1995 under way to eradicate them The plan is controversial (Anunsen and Anunsen 1993 Scheffer 1993) as is the evidence mustered to support it (compare Houston et al 119941 with Lyman 11991a 1995a 1995b1) This is not the place to review that evidence in detail (see references cited) It is relevant however to note the significance of zooarchaeological evidence for final resolution of the controversy

The historical evidence concerning the pre-1920s presence of mountain goats in the Olympic Mountains is ambiguous Some documents indicate goats were absent while others indicate goats were present (Lyman 1994a 1995b) In fact the only way to demonstrate that mountain goats were present or absent from the Olympic Mountains prior to their 1920s introduction requires paleontological andor zooarchaeological data At present there are no paleontological data of any kind for the area and the zooarchaeological record is woefully inadequate and thus cannot serve as a basis for a management decision (Lyman 1995a) Park biologists note that the purpose of national parks in the United States is not to attempt to recreate late-Pleistocene or early Holocene biotas [The NPS] is primarily concerned with historic post-Columbian specres distri- butions (Houston and Schreiner 1995 207) The facts remain however the zooarchaeo- logical record is poorly known and the historic record is internally contradictory

From the perspective of dynamic biogeographic histories provided by zooarchaeology one thing is abundantly clear the definitions of native and exotic species used by the NPS to implement its policies are contradictory Exotic species occur in a given place as a result of actions by humans whereas native species are those which presently occur or once did occ~crprior to some human influence in a place as the result of natural ecological processes The italicized phrase is where the contradiction resides because it indicates that i f for example mountain goats occurred at any tirne in the past in the Olympic Mountains

118 R Lee Lyman

- or bison in Wrangell-St Elias - then by definition the extant individuals represcl~i a native species But the introduced species are also by definition exotic having been transplanted from northwestern Canada and southeastern Alaska Olympic Nationai Parks mountain goats are of a different genetic stock and thus are a form distinct fromi mountain goats that may have been in the Olympic Mountains 200 years ago Rccogr-iizing this contradiction some years ago Y argued that if the prehistoric remains of rnonirrtain goats should be found in the Olympic Mountains then the NPS should rethink policy issues (Lyman I9RWa 22) Some NPS biologists have only recently come to the same conclusion (Houston and Schreiner 1995) And given the discussions above of appropriata forms (ecotypes subspecies species) and the benefit of hindsight I would argue that the policies and definitions of key terms should also be reworded to reflect the reality of wildlife management practice (Anonymous 1992 OBrien and Mayr 1991) Thus given their perspective of time depth zooarchaeologists not only can contribute significant paleobiological information to wildlife management concerns but perhaps they can also help write workable policies (see Wagner et al 1995 for other examples) Paleobiological information provided by zooarchaelogical research may also help wildlife managers plals for the future as the final example demonstrates

Biological preserves and planning for the future

Thus far the examples of zooarchaeology applied to wildlife management concerns have focused on the use of information on prehistoric faunal dynamics to inform management decisions That same kind of information can also be used to make broad predictions about the future effects of certain kinds of decisions For instance as alluded to in the first example presented above the spatial boundaries of parks and wildlife preserves may Piavc significant implications for the long-term survival of populations found withirl those boundaries In an important contribution to the role of planning for the future on tllc trasis of the past Graham (1988) notes that the boundaries of many parks and preserves are defined based on modern climatic and environmental conditions However tkrose conditions as we all know change and the probability of significant change increases as the length of time considered increases Graham underscores the [act that much planning lo preserve terrestrial biological diversity operates under the assumption that biotas or communities of organisms tend to respond to environmental change as intact units The

paleobiological record indicates however that individual species respond to environ- mental changes by migrating in different directions at different rates during different times (Graham 1988 392) Such individualistic responses to environmental change mean that the biological preserves of today are artifacts of the time when they were identified and created Thus Graham (1988 393) argues that constructing future reserves as habitat islands without avenues of migration [corridors] would probably result in [extirpation of bounded populations of managed species]

Graham (1988) thus suggests that as in the example of the Great Basins mountaintop mammals migration corridors are critical to organism survival and that preserves must be located on the landscape in such a rnanner as to be conducive to inter-areal migration I suggest following my earlier discussion that knowing where ihose corridors were located

The relevance of faunal analjgtsis to wildlife management 119

in the past surely will help in locating future preserves And as Graham and Graham (1994) indicate detailed knowledge of the historical and ecological biogeography of a taxon could prove critical to reconstructing appropriate habitats for reintroduced populations of the taxon Such knowledge can often come only from the fossil record

Extinction -of populations of species of lineages - is a natural fact of life (eg Raup 1984) But ignorance resulted in the loss of the passenger pigeon (Ectopistes migratorius) from North America the Great Auk (Pingninns impennis) from the North Atlantic and the dodo (Raphus cuci~llatus) from Mauritius in the Indian Ocean Our often stumbling attempts at wildlife management may due to ignorance ultimately lead to other such losses (eg Graham 1988 Hess 1993 Kay 1994) Ihe international move to maintain biological diversity - not just the taxonomic richness of biotic communities but the particular structure of those communities - seems to be in the correct direction if we are to maintain some quality of life and wilderness Understanding the loss of diversity often demands zooarchaeological and related data (eg Culotta 1995 Steadman 1995) and will help us avoid future inadvertent and unintentional losses Diamonds (1984) Rosetta Stone of using knowledge of modern extinctions to understand prehistoric extinctions could and perhaps should be turned on its head

Discussion

Nearly eighty years ago archaeologist W J Wintemberg (1919) noted how zooarchaeo- logical research could contribute to zoological knowledge and indirectly wildlife management His comments were repeated and elaborated thirty years later by biologist R M Gilmore (1949) Their remarks have largely gone unheeded perhaps because like Legge (1978 129 130) many of us see our job as aiming to contribute to the sum of the archaeological knowledge and that bone data are archaeological data I agree with Legge but also with Wintemberg and Gilmore we should be more than just consumers of wildlife data (cf Will 1982)

The potential topics under the broad umbrella of wildlife management to which zooarchaeological research might contribute are numerous I have touched on only a fevv of the more readily identifiable ones above There are to be sure others of some significance For example the modern fisheries industry of the eastern Pacific Ocean has been disrupted several times in the recent past by major changes in atmospheric circulation patterns causing alteration of ocean currents These events - variously referred to as El Nifio or Southern Oscillation events -appear to have some periodicity (Rasmusson 1985) but the historic record is of limited utility in plotting the precise nature of that periodicity These events clearly influence much of the oceanic biota (eg Arnold and Tissot 1993 Trillmich and Limberger 1985) Given sufficient temporal resolution the zooarchaeologi- cal record thus may eventually provide the time depth necessary to clarify the periodicity and magnitude of El Nifio events and thus allow more accurate predictions of future events

Zooarchaeological research has also shown that humans have long had a hand in modifying ecosystems Based on his extensive research on the prehistoric extinctions of Pacific Island bird populations and species Steadman (1995 1130) concludes that humans

tilay find Yt difficult to manage scarce resources in a sustainable way That humans contributed to these extinctions seems clear (cg Weisler and Cargett 1993) and thus Steadmans conclusion has the proper ominous tone But zooarchaeological research cair also perhaps help us out of such situations The wapiti (Cervns elaphus) of Yellowstone National Park in the United States are an example Herc nearly 60000 wapiti are managed and to a significant degree protected from human predation However historic and zooarchaeological research indicates that this population is a function of modern wildlife management historic records virtually vithout exception fail to mention wapiti i l l

the area and zooarchaeological data indicate this large ungulate was very riirc there during the last several thousand years (Kay 1994) Clearly if the Yellowstone ecosSsten1 is to be managed and maintained as a pristine wilderness in perpetuity (a laudable but in reality unattainable goal see Wager et al 1995) things must change before the ecosystem is irreparably altered A virtually identical situation exists in Rocky Mountain National Park sotile 550 km southwest of Yellowstone (Hcss 1993) where again zooarchaeologicai and other data from prehistoric times might help rcsolve debates

The few examples of how zooarchaeological research can be applied to t i ioder~~ problems I have outlined are rather specific to a place andlor to a taxon I do not perccive that to bc a weakness of an applied zooarchaeology but rather as an intellectual challenge Broadening the scope of zooarchaeology in the manner in which 1 have discussed will not be easy but i t will be beneficial to our Suture not only from the perspective of contributing to ensuring the preservation of biological diversity for future generations but also as 1conclude below from the perspective of our discipline which tilight otherwise become increasingly perceived as the pursuit of esoteric knotvledge of little practical use

One might argue that the discussion here could be expanded by suggesting that the detection of animal residues such as blood and hair (eg Loy 1993) could also providr information on the taxonomic identity of species native to the locale where the tools were collected (eg Hyland et al 1990) I doubt however that such a study would convince wildlife managers or all zooarchaeologists that a species was native given that stone tools were often transported significant distances by their tiiakers (eg Meltzer 1995) 1 beiievc multiple instances of such residues from multiple sites would be necessary to build 21

strong argument A partial skeleton or selected isolated bones and teeth of the species in que5tion recovered from a s~ngle ~ t e uould probably be muclr more conblnclng (Lyman 1994b)

I have attempted here to describe the structure of an applied zooarchaeology This is not a plea to legitimize what some of us do for fun andlor a living Rather i t is an attempt to broaden the scope of zooarchaeological research 1 am all for research for the sake of research if there is a problem in need of solving or a question in need of an answer But B am also concerned with the future of zooarchaeology Too many of our new doctoral str~dcnts are finding that the job market is closed and too marly of us arc finding it increasingly difficult to secure funding for research Perhaps a developed applied zooarchaeology wil

The relevance o f faunal analysis t o wildlife rnanagement 121

provide a new job market and new sources of funding It seems self-evident to me that if a national park for example has a staff archaeologist who deals with the archaeological resources on park lands and has a staff biologist who deals with biological resources on park lands then a marriage of the two would benefit both However given the increasing specialization of both archaeology and biology a viable marriage may not be possible Ihc obvious alternative is a park staff zooarchaeologist who has a deep knowledge ancl understanding of both disciplines and has been trained to integrate the two

New sources of funding may also become available if we can convince the wildlife management community that because its decisions and policies are typically aimed at tht future knowing something of the past can result in wise decisions That knowledge will of course come with a price tag The cost we may encounter is that members of the wildlife management community -while they may well have a solid knowledge of the time depth of evolutionary processes- often tend to view biotas as static ever enduring entities That is biotas have no past or at least their history is insignificant because they are being managed for the future While we certainly cannot predict exactly when the next ice age or glacial period will begin we can I think argue convincingly that climates and environments will change and knowing this we can use the prehistoric record to test our predictions about how certain kinds of changes may affect biotas of the future And confirmed - or even rejected or falsified -predictions would be the strong selling point Federal land-managing agencies whose charge includes wildlife management might well pay a zooarchaeologist to help them make wise management decisions The other selling point is that a less desirable kind of price -ecosystem destruction and loss of biodiversity -might otherwise have to be paid

Acknowledgements

I thank Donald K Grayson Adrian Lister Michael J OBrien and Ken Ihornas for comments on earlier versions of the manuscript

Departtnent o f Anthropology University o f Missouri-Columbia

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Brown J H 1986 Two decades of interaction between the MacArthiu-Wilson model and tile complexities of mammalian distributions Biological Journal d t h c Lirlnearl Society 28 23151

Cookc H B S 1976 Suidac from Plio-Pleistocene strata of the Rudolf Basin In Earlicst Mair a ~ r d Environmcrlts in the Lakc Rlrdolf Basin Stratigraphy Paleoecology and Evolutioir (cds Y Coppcns F C Flowcll G LI Isaac and RE F Lcakey) Chicago Univcrsity of Cllicago Press pp 25 1-63

Culotta E 1995 Many suspects to blame in Madagascar cxtinctions Science 268 1568-9

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Ilouston D B and Schrciner E 6 1995 Alien specie in national parks drawing lines in space and time Conservation Biology 9 204-9

IHouston D B Schrciner E 6and Moorhcad R R 1994 Mourltairl Goats in Olympic Nutionul Park Biologjl and Marlugernerlt of an Introduced Species U S National Park Scrvicc Scientific Monograph NPSINROLYMINRSM-9325

I-Iudson J (cd) 1993 From Bones to Bc~lzavior Etlznotrrchaeologicul and E~plcrimcntill Coiltri- butiorls ro rhe Intcrprc~tation of Fblrr~al liernairls Carbondalc Southern Illinois University Center for Archaeological Investigations Occasional Papcr No 21

T h e relevance of faunal analysis t o wildlife management 123

Hyland D C Tersak J M Adovasio J M and Sicgal M I 1990 Identification of thc speciesof origin of residual blood on lithic material Americntl Antiquity 55 104-12

Jameson R J Kenyon K W Johnson A M and Wight H M 1982 History and status of translocated sea otter populations in North America Wildlife Society Bulletin 10 100-7

Jameson R J Kenyon K W Jeffries S and VanBlaricom G R 1986 Status of a I ranslocatcd sea otter population and its habitat in Washington The Murrelet 67 84-7

Kay C E 1994 Aboriginal overkill the role of Native Americans in structur~ng western ecosystems Hurnnt~ Nrrrure 5 359-98

Kenyon K W 1969 The sea otter In the eastern Pacific Ocean Worth Amer~crrtz Firuna 68

Korn H 1994 Genetic demographic spatial environmental and catastrophic effects on the survival probability of small populations of mammals In Mitlimum Animrrl Popu1rrrior1s (ed H Remmert) Berlin Springer-Verlag pp 3)-49

Lcggc A J 1978 Archacozoology - or zooarchacology In Research Prohlems in Zoourchueology (eds D R Brothwell K D Thomas and J Clutton-Brock) Univcrsity of London Institute of Archaeology Occasional Publications No 3 pp 129-32

Leopold A S Cain S A Cottam C IM Gabrielson I N and Kimhall T L 1963 Wildlife management in the national parks Trunsuctions of the Twenty-Eighth North American Wildlife und Nutural Resoirrce~ Cd70nferencepp 2945

Loy T H 1993 The artifact as site an example of the biomolccular analysis of organic residues on prehistoric tools World Archaeology 25 44-63

Lyman R L 1988a Significance for wildlife management of the late Quaternary biogeography of mountain goats (Oreurnnos urnericunus) in the Pacific Northwest USA Arctic and Alpine Reseurch 20 13-23

Lyman R L 1988b Zoogeography of Oregon coast marine mammals the last 3000 years Murine Murnrnul Science 4 247-64

Lyman R L 1991 Late Quaternary biogeography of the pygmy rabbit (Brachylagirs iduhoensis) in eastern Washington Jo~lrnalof Mammalogy 72 11amp17

Lyman R L 1994a The Olympic Mountain goat controversy a different perspective Conservation Biology 8 898-901

Lyman R L 1994b Vertebrute Taphonomy Cambridge Cambridge University Press

Lyman R L 1995a Determining when rare (zoo)archacological phenomena are truly absent Journal of Archueologicul Method unrl Theory 2 369-424

Lyman R L 199 Inaccurate data and the Olympic National Park mountain goat controversy North wect Science 69 234-8

McAllister K and Allen H 1993 Stutuc of the Pygmy Rabbit (Brachylagus idahoensis) in Wushington Olympia Washington Department of Wildlife Wildlife Management Division

McAllister K and Allen H 1994 Wushington Stute Draft Recovery Plun for the Pygmy Rabbit (Brachylagus idahocnsis) Olympia Washington Department of Wildlife Wildlife Management Division

MacArthur R H and Wilson E 0 1967 The Theory of Islunci Biogeography Princeton NJ Princeton University Press

MacAskie I 1987 Updated status of the sea otter Enhydru lutris in Canada Cunudiun Field- Nuturcrlist 101 279-83

McDonald J N 1981 North American Bison Their Clascicution and Evolution Berkeley University of California Press

Mcltzcr D J 1995 Clocking the first Americans Annual Review of Anthropology 24 71-45

124 R Lee E y n ~ u n

OBricn S J and Mays E 1991 Burcauc1atic mischief rccogni~ing enciangcrcd spccics and subspecies Scicncc 251 1 187--8

Ogden A 1933 Russian sea ottcr anal seal hunting on the Califorilia coast Caij(~rrrirrNisroricci1 Qmcrrrerly 12 29-5 1

Olscn S I and Olscn I W 1981 A conin-icnt on nomcnclaturc in faunal studies iui~cric~un Antiquity 46 192-4

Peck J M Miclucllc 14 C and Wright M 61987 Arc bison cuotic in the Wrangcll-St Elia National Park and Pre~cr~c Envivnnnlcrzrcrl iWrlrlcrgcineilr I I l(195 3

Quinn J Iand ifarrison S P 1988 Effects of habitat fragmentation anhi isolation on spci-is richness evidence from biogeographic patterns Oecologiii 75 132-40

Rasmusson E M 1985 El Nino and asiations in climate Atzcricr~z Scic3nritt 73 168-77

Raup D M 1984 Death of spccics I n Errirrcrions (etl M 14 Nitccllti) dlhicago University of Chicago Press pp 1-19

IcnsbcrgcrJ M and Barnosky A I) 1993 Short-term fluctuations in slniill mammals ot the lato Pleistocene from eastern Washington Wn iWorphologicrrl Chr~rigc irr C)iir~t~rricrr~ Mrrinnzcrls (J ~ i i~ t l~ Alzcrica (eds R A Martin and 4 Igt 131rnosky) Qambridgc Ciimbsidgc University Prcss pp 299-342

Richards M Smallcy KSykcs R and Iledges BP 1993 iir-ch21eology and genetics anaiysirig DNA from skeletal remains Worlrl~lrchtrcolog~5 18-28

Richards M B Sykcs B C and Hedges MF M 1995 Authenticating D N A extriicted Irorri ancient skeletal remains Iourr7crl of Arcl~aeologicril Scicncc 22 2W-9

Faohison ND 1978 Zooarchaeology its hiirto~y and dcvclopnirn~7r7rrcssrc Anthropologic~ul A~orirition Miscellancotrs Pripcr 2 1-2

Rocst A 1 1973 Subspecies of the sea ottcs Er7llytira Iiitris Io5 ii7gcles Coirnty Nnturcrl tiisrory Mltscirrn (ontributior~s in Scicncr 252

Rocst A 1 1979 A re-evaluation oi sea o t t c ~ taxonomy Paper prcscntcci at the Sea Ottcr Workshop Santa Barbara Cilifornia

Scheffer V B 1993 Tlie Olympic Mo~rntain goat conrrovcrs) a pcr5pectivc Coriscrvtrtior Biology 7 916-19

Sclicffcr V B and Wilkc F 1950 Validity of the subspccics E~llzydrii 1iltris r7crcic the southcrn s ix ottcr Joltvnill qf the Wcr~I7ir7gton Accrtlcuny 01 S~icr~cr402672

Simons I) R 1083 Interactions lsctween Calilornin corrdors and humans in prehistoric far wcstern North America In Vcrltcrrr Biology und ~tlunuget~ierlt (cds S KWilbur and J A Jackson) Berkeley University or Ciliforni~i Press pp 470-91

Steadman D xi 1995 Prehistoric cstinctions of Pacific isiarld birds biodiversity nlccts zooarchacology Scieilce 267 1 123-3 1

Trillrnich F and Limbcrger D 1985 Drastic effects of 871 Nirio on Ga1apigos piniiipcd Oecologiu 67 19-22

VanBlaricorn 6Rand Estcs 9 A (cds) 1988 l h p Bot~irnlrniij bcologj oj Seu O t r e r ~ (Berlin

Springer-Verliig

Wagner F 11 Forest RC i i l l R R Mcdullough 14 R Pcltoil M MPorTcr W F and Salwasscr 13 1995 WildlifePolicies in the U S Nirrionul Purks Washington DC Island Prcsi

Weisler M I and Gargett R H 1993 Pacific island avian cstinctions the taphonorny or human predation Arclzucoloyj in Occuniu 28 85-03

Wendell F E flardy R A An-rc5 1 land Burgi K T 1986 Tcrnporalanci spatial patterns in

The relevance of faunal analyl to wrldllfe rnanugerrzent 117

The NPS considers the bison of Wrangell-St Elias National Park to be exotic due to the lack of evidence that bison were present when the tirst white explorers passed through the area (Houston and Schreiner 1995) As well NPS biologists note that the introduced forrn of bison was of an incorrect - non-native -genetic stock (a distinct subspecies or ecotype) given current beliefs about bison taxonomy For this reason alone perhaps the extant bison of Wrangell-St Elias should be removed However the zooarchaeological record is so poorly known that one cannot conclusively demonstrate that native bison were locally extinct by the beginning of the twentieth century This in turn raises the possibility that extant bison are hybrids of native and introduced genetic stocks Whether or not this possibility is in fact reality is irrelevant What is critical here is that only more zooarchaeological and paleontological research will establish first the timing of local extirpation of native Alaskan bison populations and second if all Alaskan populations were extinct when transplanting occurred in the middle of the twentieth century Once the facts are determined we will have strong bases for making a decision regarding the ultimate fate of the bison presently living in Wrangell-St Elias National Park and Preserve

I am much more familiar - and intimately involved -with a virtually identical situation that exists in western Washington state Here mountain goats (Oreamnos urnericanlls) were introduced in the 1920s to the Olympic Mountains A decade later in 1938 these mountains were designated Olympic National Park The present position of the NPS is that the mountain goats are exotic and plans are in mid-1995 under way to eradicate them The plan is controversial (Anunsen and Anunsen 1993 Scheffer 1993) as is the evidence mustered to support it (compare Houston et al 119941 with Lyman 11991a 1995a 1995b1) This is not the place to review that evidence in detail (see references cited) It is relevant however to note the significance of zooarchaeological evidence for final resolution of the controversy

The historical evidence concerning the pre-1920s presence of mountain goats in the Olympic Mountains is ambiguous Some documents indicate goats were absent while others indicate goats were present (Lyman 1994a 1995b) In fact the only way to demonstrate that mountain goats were present or absent from the Olympic Mountains prior to their 1920s introduction requires paleontological andor zooarchaeological data At present there are no paleontological data of any kind for the area and the zooarchaeological record is woefully inadequate and thus cannot serve as a basis for a management decision (Lyman 1995a) Park biologists note that the purpose of national parks in the United States is not to attempt to recreate late-Pleistocene or early Holocene biotas [The NPS] is primarily concerned with historic post-Columbian specres distri- butions (Houston and Schreiner 1995 207) The facts remain however the zooarchaeo- logical record is poorly known and the historic record is internally contradictory

From the perspective of dynamic biogeographic histories provided by zooarchaeology one thing is abundantly clear the definitions of native and exotic species used by the NPS to implement its policies are contradictory Exotic species occur in a given place as a result of actions by humans whereas native species are those which presently occur or once did occ~crprior to some human influence in a place as the result of natural ecological processes The italicized phrase is where the contradiction resides because it indicates that i f for example mountain goats occurred at any tirne in the past in the Olympic Mountains

118 R Lee Lyman

- or bison in Wrangell-St Elias - then by definition the extant individuals represcl~i a native species But the introduced species are also by definition exotic having been transplanted from northwestern Canada and southeastern Alaska Olympic Nationai Parks mountain goats are of a different genetic stock and thus are a form distinct fromi mountain goats that may have been in the Olympic Mountains 200 years ago Rccogr-iizing this contradiction some years ago Y argued that if the prehistoric remains of rnonirrtain goats should be found in the Olympic Mountains then the NPS should rethink policy issues (Lyman I9RWa 22) Some NPS biologists have only recently come to the same conclusion (Houston and Schreiner 1995) And given the discussions above of appropriata forms (ecotypes subspecies species) and the benefit of hindsight I would argue that the policies and definitions of key terms should also be reworded to reflect the reality of wildlife management practice (Anonymous 1992 OBrien and Mayr 1991) Thus given their perspective of time depth zooarchaeologists not only can contribute significant paleobiological information to wildlife management concerns but perhaps they can also help write workable policies (see Wagner et al 1995 for other examples) Paleobiological information provided by zooarchaelogical research may also help wildlife managers plals for the future as the final example demonstrates

Biological preserves and planning for the future

Thus far the examples of zooarchaeology applied to wildlife management concerns have focused on the use of information on prehistoric faunal dynamics to inform management decisions That same kind of information can also be used to make broad predictions about the future effects of certain kinds of decisions For instance as alluded to in the first example presented above the spatial boundaries of parks and wildlife preserves may Piavc significant implications for the long-term survival of populations found withirl those boundaries In an important contribution to the role of planning for the future on tllc trasis of the past Graham (1988) notes that the boundaries of many parks and preserves are defined based on modern climatic and environmental conditions However tkrose conditions as we all know change and the probability of significant change increases as the length of time considered increases Graham underscores the [act that much planning lo preserve terrestrial biological diversity operates under the assumption that biotas or communities of organisms tend to respond to environmental change as intact units The

paleobiological record indicates however that individual species respond to environ- mental changes by migrating in different directions at different rates during different times (Graham 1988 392) Such individualistic responses to environmental change mean that the biological preserves of today are artifacts of the time when they were identified and created Thus Graham (1988 393) argues that constructing future reserves as habitat islands without avenues of migration [corridors] would probably result in [extirpation of bounded populations of managed species]

Graham (1988) thus suggests that as in the example of the Great Basins mountaintop mammals migration corridors are critical to organism survival and that preserves must be located on the landscape in such a rnanner as to be conducive to inter-areal migration I suggest following my earlier discussion that knowing where ihose corridors were located

The relevance of faunal analjgtsis to wildlife management 119

in the past surely will help in locating future preserves And as Graham and Graham (1994) indicate detailed knowledge of the historical and ecological biogeography of a taxon could prove critical to reconstructing appropriate habitats for reintroduced populations of the taxon Such knowledge can often come only from the fossil record

Extinction -of populations of species of lineages - is a natural fact of life (eg Raup 1984) But ignorance resulted in the loss of the passenger pigeon (Ectopistes migratorius) from North America the Great Auk (Pingninns impennis) from the North Atlantic and the dodo (Raphus cuci~llatus) from Mauritius in the Indian Ocean Our often stumbling attempts at wildlife management may due to ignorance ultimately lead to other such losses (eg Graham 1988 Hess 1993 Kay 1994) Ihe international move to maintain biological diversity - not just the taxonomic richness of biotic communities but the particular structure of those communities - seems to be in the correct direction if we are to maintain some quality of life and wilderness Understanding the loss of diversity often demands zooarchaeological and related data (eg Culotta 1995 Steadman 1995) and will help us avoid future inadvertent and unintentional losses Diamonds (1984) Rosetta Stone of using knowledge of modern extinctions to understand prehistoric extinctions could and perhaps should be turned on its head

Discussion

Nearly eighty years ago archaeologist W J Wintemberg (1919) noted how zooarchaeo- logical research could contribute to zoological knowledge and indirectly wildlife management His comments were repeated and elaborated thirty years later by biologist R M Gilmore (1949) Their remarks have largely gone unheeded perhaps because like Legge (1978 129 130) many of us see our job as aiming to contribute to the sum of the archaeological knowledge and that bone data are archaeological data I agree with Legge but also with Wintemberg and Gilmore we should be more than just consumers of wildlife data (cf Will 1982)

The potential topics under the broad umbrella of wildlife management to which zooarchaeological research might contribute are numerous I have touched on only a fevv of the more readily identifiable ones above There are to be sure others of some significance For example the modern fisheries industry of the eastern Pacific Ocean has been disrupted several times in the recent past by major changes in atmospheric circulation patterns causing alteration of ocean currents These events - variously referred to as El Nifio or Southern Oscillation events -appear to have some periodicity (Rasmusson 1985) but the historic record is of limited utility in plotting the precise nature of that periodicity These events clearly influence much of the oceanic biota (eg Arnold and Tissot 1993 Trillmich and Limberger 1985) Given sufficient temporal resolution the zooarchaeologi- cal record thus may eventually provide the time depth necessary to clarify the periodicity and magnitude of El Nifio events and thus allow more accurate predictions of future events

Zooarchaeological research has also shown that humans have long had a hand in modifying ecosystems Based on his extensive research on the prehistoric extinctions of Pacific Island bird populations and species Steadman (1995 1130) concludes that humans

tilay find Yt difficult to manage scarce resources in a sustainable way That humans contributed to these extinctions seems clear (cg Weisler and Cargett 1993) and thus Steadmans conclusion has the proper ominous tone But zooarchaeological research cair also perhaps help us out of such situations The wapiti (Cervns elaphus) of Yellowstone National Park in the United States are an example Herc nearly 60000 wapiti are managed and to a significant degree protected from human predation However historic and zooarchaeological research indicates that this population is a function of modern wildlife management historic records virtually vithout exception fail to mention wapiti i l l

the area and zooarchaeological data indicate this large ungulate was very riirc there during the last several thousand years (Kay 1994) Clearly if the Yellowstone ecosSsten1 is to be managed and maintained as a pristine wilderness in perpetuity (a laudable but in reality unattainable goal see Wager et al 1995) things must change before the ecosystem is irreparably altered A virtually identical situation exists in Rocky Mountain National Park sotile 550 km southwest of Yellowstone (Hcss 1993) where again zooarchaeologicai and other data from prehistoric times might help rcsolve debates

The few examples of how zooarchaeological research can be applied to t i ioder~~ problems I have outlined are rather specific to a place andlor to a taxon I do not perccive that to bc a weakness of an applied zooarchaeology but rather as an intellectual challenge Broadening the scope of zooarchaeology in the manner in which 1 have discussed will not be easy but i t will be beneficial to our Suture not only from the perspective of contributing to ensuring the preservation of biological diversity for future generations but also as 1conclude below from the perspective of our discipline which tilight otherwise become increasingly perceived as the pursuit of esoteric knotvledge of little practical use

One might argue that the discussion here could be expanded by suggesting that the detection of animal residues such as blood and hair (eg Loy 1993) could also providr information on the taxonomic identity of species native to the locale where the tools were collected (eg Hyland et al 1990) I doubt however that such a study would convince wildlife managers or all zooarchaeologists that a species was native given that stone tools were often transported significant distances by their tiiakers (eg Meltzer 1995) 1 beiievc multiple instances of such residues from multiple sites would be necessary to build 21

strong argument A partial skeleton or selected isolated bones and teeth of the species in que5tion recovered from a s~ngle ~ t e uould probably be muclr more conblnclng (Lyman 1994b)

I have attempted here to describe the structure of an applied zooarchaeology This is not a plea to legitimize what some of us do for fun andlor a living Rather i t is an attempt to broaden the scope of zooarchaeological research 1 am all for research for the sake of research if there is a problem in need of solving or a question in need of an answer But B am also concerned with the future of zooarchaeology Too many of our new doctoral str~dcnts are finding that the job market is closed and too marly of us arc finding it increasingly difficult to secure funding for research Perhaps a developed applied zooarchaeology wil

The relevance o f faunal analysis t o wildlife rnanagement 121

provide a new job market and new sources of funding It seems self-evident to me that if a national park for example has a staff archaeologist who deals with the archaeological resources on park lands and has a staff biologist who deals with biological resources on park lands then a marriage of the two would benefit both However given the increasing specialization of both archaeology and biology a viable marriage may not be possible Ihc obvious alternative is a park staff zooarchaeologist who has a deep knowledge ancl understanding of both disciplines and has been trained to integrate the two

New sources of funding may also become available if we can convince the wildlife management community that because its decisions and policies are typically aimed at tht future knowing something of the past can result in wise decisions That knowledge will of course come with a price tag The cost we may encounter is that members of the wildlife management community -while they may well have a solid knowledge of the time depth of evolutionary processes- often tend to view biotas as static ever enduring entities That is biotas have no past or at least their history is insignificant because they are being managed for the future While we certainly cannot predict exactly when the next ice age or glacial period will begin we can I think argue convincingly that climates and environments will change and knowing this we can use the prehistoric record to test our predictions about how certain kinds of changes may affect biotas of the future And confirmed - or even rejected or falsified -predictions would be the strong selling point Federal land-managing agencies whose charge includes wildlife management might well pay a zooarchaeologist to help them make wise management decisions The other selling point is that a less desirable kind of price -ecosystem destruction and loss of biodiversity -might otherwise have to be paid

Acknowledgements

I thank Donald K Grayson Adrian Lister Michael J OBrien and Ken Ihornas for comments on earlier versions of the manuscript

Departtnent o f Anthropology University o f Missouri-Columbia

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I-Iudson J (cd) 1993 From Bones to Bc~lzavior Etlznotrrchaeologicul and E~plcrimcntill Coiltri- butiorls ro rhe Intcrprc~tation of Fblrr~al liernairls Carbondalc Southern Illinois University Center for Archaeological Investigations Occasional Papcr No 21

T h e relevance of faunal analysis t o wildlife management 123

Hyland D C Tersak J M Adovasio J M and Sicgal M I 1990 Identification of thc speciesof origin of residual blood on lithic material Americntl Antiquity 55 104-12

Jameson R J Kenyon K W Johnson A M and Wight H M 1982 History and status of translocated sea otter populations in North America Wildlife Society Bulletin 10 100-7

Jameson R J Kenyon K W Jeffries S and VanBlaricom G R 1986 Status of a I ranslocatcd sea otter population and its habitat in Washington The Murrelet 67 84-7

Kay C E 1994 Aboriginal overkill the role of Native Americans in structur~ng western ecosystems Hurnnt~ Nrrrure 5 359-98

Kenyon K W 1969 The sea otter In the eastern Pacific Ocean Worth Amer~crrtz Firuna 68

Korn H 1994 Genetic demographic spatial environmental and catastrophic effects on the survival probability of small populations of mammals In Mitlimum Animrrl Popu1rrrior1s (ed H Remmert) Berlin Springer-Verlag pp 3)-49

Lcggc A J 1978 Archacozoology - or zooarchacology In Research Prohlems in Zoourchueology (eds D R Brothwell K D Thomas and J Clutton-Brock) Univcrsity of London Institute of Archaeology Occasional Publications No 3 pp 129-32

Leopold A S Cain S A Cottam C IM Gabrielson I N and Kimhall T L 1963 Wildlife management in the national parks Trunsuctions of the Twenty-Eighth North American Wildlife und Nutural Resoirrce~ Cd70nferencepp 2945

Loy T H 1993 The artifact as site an example of the biomolccular analysis of organic residues on prehistoric tools World Archaeology 25 44-63

Lyman R L 1988a Significance for wildlife management of the late Quaternary biogeography of mountain goats (Oreurnnos urnericunus) in the Pacific Northwest USA Arctic and Alpine Reseurch 20 13-23

Lyman R L 1988b Zoogeography of Oregon coast marine mammals the last 3000 years Murine Murnrnul Science 4 247-64

Lyman R L 1991 Late Quaternary biogeography of the pygmy rabbit (Brachylagirs iduhoensis) in eastern Washington Jo~lrnalof Mammalogy 72 11amp17

Lyman R L 1994a The Olympic Mountain goat controversy a different perspective Conservation Biology 8 898-901

Lyman R L 1994b Vertebrute Taphonomy Cambridge Cambridge University Press

Lyman R L 1995a Determining when rare (zoo)archacological phenomena are truly absent Journal of Archueologicul Method unrl Theory 2 369-424

Lyman R L 199 Inaccurate data and the Olympic National Park mountain goat controversy North wect Science 69 234-8

McAllister K and Allen H 1993 Stutuc of the Pygmy Rabbit (Brachylagus idahoensis) in Wushington Olympia Washington Department of Wildlife Wildlife Management Division

McAllister K and Allen H 1994 Wushington Stute Draft Recovery Plun for the Pygmy Rabbit (Brachylagus idahocnsis) Olympia Washington Department of Wildlife Wildlife Management Division

MacArthur R H and Wilson E 0 1967 The Theory of Islunci Biogeography Princeton NJ Princeton University Press

MacAskie I 1987 Updated status of the sea otter Enhydru lutris in Canada Cunudiun Field- Nuturcrlist 101 279-83

McDonald J N 1981 North American Bison Their Clascicution and Evolution Berkeley University of California Press

Mcltzcr D J 1995 Clocking the first Americans Annual Review of Anthropology 24 71-45

124 R Lee E y n ~ u n

OBricn S J and Mays E 1991 Burcauc1atic mischief rccogni~ing enciangcrcd spccics and subspecies Scicncc 251 1 187--8

Ogden A 1933 Russian sea ottcr anal seal hunting on the Califorilia coast Caij(~rrrirrNisroricci1 Qmcrrrerly 12 29-5 1

Olscn S I and Olscn I W 1981 A conin-icnt on nomcnclaturc in faunal studies iui~cric~un Antiquity 46 192-4

Peck J M Miclucllc 14 C and Wright M 61987 Arc bison cuotic in the Wrangcll-St Elia National Park and Pre~cr~c Envivnnnlcrzrcrl iWrlrlcrgcineilr I I l(195 3

Quinn J Iand ifarrison S P 1988 Effects of habitat fragmentation anhi isolation on spci-is richness evidence from biogeographic patterns Oecologiii 75 132-40

Rasmusson E M 1985 El Nino and asiations in climate Atzcricr~z Scic3nritt 73 168-77

Raup D M 1984 Death of spccics I n Errirrcrions (etl M 14 Nitccllti) dlhicago University of Chicago Press pp 1-19

IcnsbcrgcrJ M and Barnosky A I) 1993 Short-term fluctuations in slniill mammals ot the lato Pleistocene from eastern Washington Wn iWorphologicrrl Chr~rigc irr C)iir~t~rricrr~ Mrrinnzcrls (J ~ i i~ t l~ Alzcrica (eds R A Martin and 4 Igt 131rnosky) Qambridgc Ciimbsidgc University Prcss pp 299-342

Richards M Smallcy KSykcs R and Iledges BP 1993 iir-ch21eology and genetics anaiysirig DNA from skeletal remains Worlrl~lrchtrcolog~5 18-28

Richards M B Sykcs B C and Hedges MF M 1995 Authenticating D N A extriicted Irorri ancient skeletal remains Iourr7crl of Arcl~aeologicril Scicncc 22 2W-9

Faohison ND 1978 Zooarchaeology its hiirto~y and dcvclopnirn~7r7rrcssrc Anthropologic~ul A~orirition Miscellancotrs Pripcr 2 1-2

Rocst A 1 1973 Subspecies of the sea ottcs Er7llytira Iiitris Io5 ii7gcles Coirnty Nnturcrl tiisrory Mltscirrn (ontributior~s in Scicncr 252

Rocst A 1 1979 A re-evaluation oi sea o t t c ~ taxonomy Paper prcscntcci at the Sea Ottcr Workshop Santa Barbara Cilifornia

Scheffer V B 1993 Tlie Olympic Mo~rntain goat conrrovcrs) a pcr5pectivc Coriscrvtrtior Biology 7 916-19

Sclicffcr V B and Wilkc F 1950 Validity of the subspccics E~llzydrii 1iltris r7crcic the southcrn s ix ottcr Joltvnill qf the Wcr~I7ir7gton Accrtlcuny 01 S~icr~cr402672

Simons I) R 1083 Interactions lsctween Calilornin corrdors and humans in prehistoric far wcstern North America In Vcrltcrrr Biology und ~tlunuget~ierlt (cds S KWilbur and J A Jackson) Berkeley University or Ciliforni~i Press pp 470-91

Steadman D xi 1995 Prehistoric cstinctions of Pacific isiarld birds biodiversity nlccts zooarchacology Scieilce 267 1 123-3 1

Trillrnich F and Limbcrger D 1985 Drastic effects of 871 Nirio on Ga1apigos piniiipcd Oecologiu 67 19-22

VanBlaricorn 6Rand Estcs 9 A (cds) 1988 l h p Bot~irnlrniij bcologj oj Seu O t r e r ~ (Berlin

Springer-Verliig

Wagner F 11 Forest RC i i l l R R Mcdullough 14 R Pcltoil M MPorTcr W F and Salwasscr 13 1995 WildlifePolicies in the U S Nirrionul Purks Washington DC Island Prcsi

Weisler M I and Gargett R H 1993 Pacific island avian cstinctions the taphonorny or human predation Arclzucoloyj in Occuniu 28 85-03

Wendell F E flardy R A An-rc5 1 land Burgi K T 1986 Tcrnporalanci spatial patterns in

118 R Lee Lyman

- or bison in Wrangell-St Elias - then by definition the extant individuals represcl~i a native species But the introduced species are also by definition exotic having been transplanted from northwestern Canada and southeastern Alaska Olympic Nationai Parks mountain goats are of a different genetic stock and thus are a form distinct fromi mountain goats that may have been in the Olympic Mountains 200 years ago Rccogr-iizing this contradiction some years ago Y argued that if the prehistoric remains of rnonirrtain goats should be found in the Olympic Mountains then the NPS should rethink policy issues (Lyman I9RWa 22) Some NPS biologists have only recently come to the same conclusion (Houston and Schreiner 1995) And given the discussions above of appropriata forms (ecotypes subspecies species) and the benefit of hindsight I would argue that the policies and definitions of key terms should also be reworded to reflect the reality of wildlife management practice (Anonymous 1992 OBrien and Mayr 1991) Thus given their perspective of time depth zooarchaeologists not only can contribute significant paleobiological information to wildlife management concerns but perhaps they can also help write workable policies (see Wagner et al 1995 for other examples) Paleobiological information provided by zooarchaelogical research may also help wildlife managers plals for the future as the final example demonstrates

Biological preserves and planning for the future

Thus far the examples of zooarchaeology applied to wildlife management concerns have focused on the use of information on prehistoric faunal dynamics to inform management decisions That same kind of information can also be used to make broad predictions about the future effects of certain kinds of decisions For instance as alluded to in the first example presented above the spatial boundaries of parks and wildlife preserves may Piavc significant implications for the long-term survival of populations found withirl those boundaries In an important contribution to the role of planning for the future on tllc trasis of the past Graham (1988) notes that the boundaries of many parks and preserves are defined based on modern climatic and environmental conditions However tkrose conditions as we all know change and the probability of significant change increases as the length of time considered increases Graham underscores the [act that much planning lo preserve terrestrial biological diversity operates under the assumption that biotas or communities of organisms tend to respond to environmental change as intact units The

paleobiological record indicates however that individual species respond to environ- mental changes by migrating in different directions at different rates during different times (Graham 1988 392) Such individualistic responses to environmental change mean that the biological preserves of today are artifacts of the time when they were identified and created Thus Graham (1988 393) argues that constructing future reserves as habitat islands without avenues of migration [corridors] would probably result in [extirpation of bounded populations of managed species]

Graham (1988) thus suggests that as in the example of the Great Basins mountaintop mammals migration corridors are critical to organism survival and that preserves must be located on the landscape in such a rnanner as to be conducive to inter-areal migration I suggest following my earlier discussion that knowing where ihose corridors were located

The relevance of faunal analjgtsis to wildlife management 119

in the past surely will help in locating future preserves And as Graham and Graham (1994) indicate detailed knowledge of the historical and ecological biogeography of a taxon could prove critical to reconstructing appropriate habitats for reintroduced populations of the taxon Such knowledge can often come only from the fossil record

Extinction -of populations of species of lineages - is a natural fact of life (eg Raup 1984) But ignorance resulted in the loss of the passenger pigeon (Ectopistes migratorius) from North America the Great Auk (Pingninns impennis) from the North Atlantic and the dodo (Raphus cuci~llatus) from Mauritius in the Indian Ocean Our often stumbling attempts at wildlife management may due to ignorance ultimately lead to other such losses (eg Graham 1988 Hess 1993 Kay 1994) Ihe international move to maintain biological diversity - not just the taxonomic richness of biotic communities but the particular structure of those communities - seems to be in the correct direction if we are to maintain some quality of life and wilderness Understanding the loss of diversity often demands zooarchaeological and related data (eg Culotta 1995 Steadman 1995) and will help us avoid future inadvertent and unintentional losses Diamonds (1984) Rosetta Stone of using knowledge of modern extinctions to understand prehistoric extinctions could and perhaps should be turned on its head

Discussion

Nearly eighty years ago archaeologist W J Wintemberg (1919) noted how zooarchaeo- logical research could contribute to zoological knowledge and indirectly wildlife management His comments were repeated and elaborated thirty years later by biologist R M Gilmore (1949) Their remarks have largely gone unheeded perhaps because like Legge (1978 129 130) many of us see our job as aiming to contribute to the sum of the archaeological knowledge and that bone data are archaeological data I agree with Legge but also with Wintemberg and Gilmore we should be more than just consumers of wildlife data (cf Will 1982)

The potential topics under the broad umbrella of wildlife management to which zooarchaeological research might contribute are numerous I have touched on only a fevv of the more readily identifiable ones above There are to be sure others of some significance For example the modern fisheries industry of the eastern Pacific Ocean has been disrupted several times in the recent past by major changes in atmospheric circulation patterns causing alteration of ocean currents These events - variously referred to as El Nifio or Southern Oscillation events -appear to have some periodicity (Rasmusson 1985) but the historic record is of limited utility in plotting the precise nature of that periodicity These events clearly influence much of the oceanic biota (eg Arnold and Tissot 1993 Trillmich and Limberger 1985) Given sufficient temporal resolution the zooarchaeologi- cal record thus may eventually provide the time depth necessary to clarify the periodicity and magnitude of El Nifio events and thus allow more accurate predictions of future events

Zooarchaeological research has also shown that humans have long had a hand in modifying ecosystems Based on his extensive research on the prehistoric extinctions of Pacific Island bird populations and species Steadman (1995 1130) concludes that humans

tilay find Yt difficult to manage scarce resources in a sustainable way That humans contributed to these extinctions seems clear (cg Weisler and Cargett 1993) and thus Steadmans conclusion has the proper ominous tone But zooarchaeological research cair also perhaps help us out of such situations The wapiti (Cervns elaphus) of Yellowstone National Park in the United States are an example Herc nearly 60000 wapiti are managed and to a significant degree protected from human predation However historic and zooarchaeological research indicates that this population is a function of modern wildlife management historic records virtually vithout exception fail to mention wapiti i l l

the area and zooarchaeological data indicate this large ungulate was very riirc there during the last several thousand years (Kay 1994) Clearly if the Yellowstone ecosSsten1 is to be managed and maintained as a pristine wilderness in perpetuity (a laudable but in reality unattainable goal see Wager et al 1995) things must change before the ecosystem is irreparably altered A virtually identical situation exists in Rocky Mountain National Park sotile 550 km southwest of Yellowstone (Hcss 1993) where again zooarchaeologicai and other data from prehistoric times might help rcsolve debates

The few examples of how zooarchaeological research can be applied to t i ioder~~ problems I have outlined are rather specific to a place andlor to a taxon I do not perccive that to bc a weakness of an applied zooarchaeology but rather as an intellectual challenge Broadening the scope of zooarchaeology in the manner in which 1 have discussed will not be easy but i t will be beneficial to our Suture not only from the perspective of contributing to ensuring the preservation of biological diversity for future generations but also as 1conclude below from the perspective of our discipline which tilight otherwise become increasingly perceived as the pursuit of esoteric knotvledge of little practical use

One might argue that the discussion here could be expanded by suggesting that the detection of animal residues such as blood and hair (eg Loy 1993) could also providr information on the taxonomic identity of species native to the locale where the tools were collected (eg Hyland et al 1990) I doubt however that such a study would convince wildlife managers or all zooarchaeologists that a species was native given that stone tools were often transported significant distances by their tiiakers (eg Meltzer 1995) 1 beiievc multiple instances of such residues from multiple sites would be necessary to build 21

strong argument A partial skeleton or selected isolated bones and teeth of the species in que5tion recovered from a s~ngle ~ t e uould probably be muclr more conblnclng (Lyman 1994b)

I have attempted here to describe the structure of an applied zooarchaeology This is not a plea to legitimize what some of us do for fun andlor a living Rather i t is an attempt to broaden the scope of zooarchaeological research 1 am all for research for the sake of research if there is a problem in need of solving or a question in need of an answer But B am also concerned with the future of zooarchaeology Too many of our new doctoral str~dcnts are finding that the job market is closed and too marly of us arc finding it increasingly difficult to secure funding for research Perhaps a developed applied zooarchaeology wil

The relevance o f faunal analysis t o wildlife rnanagement 121

provide a new job market and new sources of funding It seems self-evident to me that if a national park for example has a staff archaeologist who deals with the archaeological resources on park lands and has a staff biologist who deals with biological resources on park lands then a marriage of the two would benefit both However given the increasing specialization of both archaeology and biology a viable marriage may not be possible Ihc obvious alternative is a park staff zooarchaeologist who has a deep knowledge ancl understanding of both disciplines and has been trained to integrate the two

New sources of funding may also become available if we can convince the wildlife management community that because its decisions and policies are typically aimed at tht future knowing something of the past can result in wise decisions That knowledge will of course come with a price tag The cost we may encounter is that members of the wildlife management community -while they may well have a solid knowledge of the time depth of evolutionary processes- often tend to view biotas as static ever enduring entities That is biotas have no past or at least their history is insignificant because they are being managed for the future While we certainly cannot predict exactly when the next ice age or glacial period will begin we can I think argue convincingly that climates and environments will change and knowing this we can use the prehistoric record to test our predictions about how certain kinds of changes may affect biotas of the future And confirmed - or even rejected or falsified -predictions would be the strong selling point Federal land-managing agencies whose charge includes wildlife management might well pay a zooarchaeologist to help them make wise management decisions The other selling point is that a less desirable kind of price -ecosystem destruction and loss of biodiversity -might otherwise have to be paid

Acknowledgements

I thank Donald K Grayson Adrian Lister Michael J OBrien and Ken Ihornas for comments on earlier versions of the manuscript

Departtnent o f Anthropology University o f Missouri-Columbia

References

Anonymous 1992 Draft guidclincs for rcintroductions hZws1etttr of the ILJCN feintvorirlctiorr Specinlist Group 4 2-3

Anunscn C S and Anunsen R 1993 Rcsponsc to Sclleffer Conservation Biology 7 054-7

Arnold J E andTissot B N 1993 Measurcmcnt of significant rnarine palcotcmperature variation using black abalone shells from prehistoric middens Quaternurj~ Iesearch 39 3901

Brewer D J 1992 Zooarchacology method tlleory and goals In Archaeological Metl~od and Theory Vol 4 (ed M B Schiffer) Tucson University of Arizona Press pp 195-244

Brown J H 1971 Mammals on mountaintops nonequilibrium insular biogcograplly American Vr~turulist105 467-78

Brown J PI 1978 The theory of insular biogeography and the distribution of boreal mammals and birds In Intermountain Biogeogrur~hy A Synzposium (eds KT Harper and J L Rcvcal) Great Basin Natural Memoir 2 pp 209-28

Brown J H 1986 Two decades of interaction between the MacArthiu-Wilson model and tile complexities of mammalian distributions Biological Journal d t h c Lirlnearl Society 28 23151

Cookc H B S 1976 Suidac from Plio-Pleistocene strata of the Rudolf Basin In Earlicst Mair a ~ r d Environmcrlts in the Lakc Rlrdolf Basin Stratigraphy Paleoecology and Evolutioir (cds Y Coppcns F C Flowcll G LI Isaac and RE F Lcakey) Chicago Univcrsity of Cllicago Press pp 25 1-63

Culotta E 1995 Many suspects to blame in Madagascar cxtinctions Science 268 1568-9

Davis J and Lidicker W Z Jr 1975 Tlle taxonomic status of tllc soutl~ern sea otter P r o c ~ ~ d i i z g oJtlze Cr~lijorrliri Acriderny o f Sciences 40(14) 429-37

Diamond J M 1984 Historic extinctions a Rosetta Stone for understanding prcllistoric extinctions in Quuterrlury Extir1ctions A Prehistoric Revolution (eds P S Martin and R 6Klcin) Tucson University of Arizona Prcss pp 824-62

Emslic S D 1987 Age and diet of fossil California condors in Grand Canyon Arizona Science 237 768-70

Fisller J W Jr 1995 Bone surface modifications in 7ooarchaeology Journal ofArchacologica1 Method rind Theory 2 7-68

Gifford-Gonzalez D 1991 Bones arc not enough analogues knowledge and intcrprctivc strategies in zooarchaeology Journal ofAntlzropologica1 Archrreologj 10 215-54

Gilmorc R M 1949 Tllc identification and value of mammal bones from archeological excavations Journal of Mummulogy 30 163-9

Graham RW 1988 The role of climatic change in the design of biological reserves tho palcoccological perspccti~rc for conser~ration biology Corltervution Biologjl 2 391-4

Graham RW and Graham M A 1994 Late Quaternary distribution of Marter in North America In Mcrrten~ Sublcs urtd Fislrcrs Biology and Conservation (eds S W Ruskirk 14 S Harestad M 6Raphael and RA Powell) Itllaca NY Corncll University Prcss pp 2 6 5 8

Grayson D M 1991 Tllc biogeographic history of small mammals in the Great Basin observations on thc last 20000 years Journal of Mammalogy 68 359-75

Green J S and Flinders J T 1980a Brachylrig~s idahoerisis Manznzalian Specics 125 1--4

Green J S and Flindcrs J T 1980b Habitat and dietary relationships of the pygmy rabbit dournnl Kringe Marlrigernent 33 13amp42

I3css K J r 1993 Rocky Times in Rocky Mourltairl hcrtiorzril Park Niwot Colo University Prcss of Colorado

Flestcr F E 1991 The U S National Park Scrvicc experience with exotic species Nriturril Areri ~ Jourr~ril1 1 127-8

Hoerr W 1993 The concept of naturalness in environmental tliscoursc Natural Areas Journal 1329-32

Ilouston D B and Schrciner E 6 1995 Alien specie in national parks drawing lines in space and time Conservation Biology 9 204-9

IHouston D B Schrciner E 6and Moorhcad R R 1994 Mourltairl Goats in Olympic Nutionul Park Biologjl and Marlugernerlt of an Introduced Species U S National Park Scrvicc Scientific Monograph NPSINROLYMINRSM-9325

I-Iudson J (cd) 1993 From Bones to Bc~lzavior Etlznotrrchaeologicul and E~plcrimcntill Coiltri- butiorls ro rhe Intcrprc~tation of Fblrr~al liernairls Carbondalc Southern Illinois University Center for Archaeological Investigations Occasional Papcr No 21

T h e relevance of faunal analysis t o wildlife management 123

Hyland D C Tersak J M Adovasio J M and Sicgal M I 1990 Identification of thc speciesof origin of residual blood on lithic material Americntl Antiquity 55 104-12

Jameson R J Kenyon K W Johnson A M and Wight H M 1982 History and status of translocated sea otter populations in North America Wildlife Society Bulletin 10 100-7

Jameson R J Kenyon K W Jeffries S and VanBlaricom G R 1986 Status of a I ranslocatcd sea otter population and its habitat in Washington The Murrelet 67 84-7

Kay C E 1994 Aboriginal overkill the role of Native Americans in structur~ng western ecosystems Hurnnt~ Nrrrure 5 359-98

Kenyon K W 1969 The sea otter In the eastern Pacific Ocean Worth Amer~crrtz Firuna 68

Korn H 1994 Genetic demographic spatial environmental and catastrophic effects on the survival probability of small populations of mammals In Mitlimum Animrrl Popu1rrrior1s (ed H Remmert) Berlin Springer-Verlag pp 3)-49

Lcggc A J 1978 Archacozoology - or zooarchacology In Research Prohlems in Zoourchueology (eds D R Brothwell K D Thomas and J Clutton-Brock) Univcrsity of London Institute of Archaeology Occasional Publications No 3 pp 129-32

Leopold A S Cain S A Cottam C IM Gabrielson I N and Kimhall T L 1963 Wildlife management in the national parks Trunsuctions of the Twenty-Eighth North American Wildlife und Nutural Resoirrce~ Cd70nferencepp 2945

Loy T H 1993 The artifact as site an example of the biomolccular analysis of organic residues on prehistoric tools World Archaeology 25 44-63

Lyman R L 1988a Significance for wildlife management of the late Quaternary biogeography of mountain goats (Oreurnnos urnericunus) in the Pacific Northwest USA Arctic and Alpine Reseurch 20 13-23

Lyman R L 1988b Zoogeography of Oregon coast marine mammals the last 3000 years Murine Murnrnul Science 4 247-64

Lyman R L 1991 Late Quaternary biogeography of the pygmy rabbit (Brachylagirs iduhoensis) in eastern Washington Jo~lrnalof Mammalogy 72 11amp17

Lyman R L 1994a The Olympic Mountain goat controversy a different perspective Conservation Biology 8 898-901

Lyman R L 1994b Vertebrute Taphonomy Cambridge Cambridge University Press

Lyman R L 1995a Determining when rare (zoo)archacological phenomena are truly absent Journal of Archueologicul Method unrl Theory 2 369-424

Lyman R L 199 Inaccurate data and the Olympic National Park mountain goat controversy North wect Science 69 234-8

McAllister K and Allen H 1993 Stutuc of the Pygmy Rabbit (Brachylagus idahoensis) in Wushington Olympia Washington Department of Wildlife Wildlife Management Division

McAllister K and Allen H 1994 Wushington Stute Draft Recovery Plun for the Pygmy Rabbit (Brachylagus idahocnsis) Olympia Washington Department of Wildlife Wildlife Management Division

MacArthur R H and Wilson E 0 1967 The Theory of Islunci Biogeography Princeton NJ Princeton University Press

MacAskie I 1987 Updated status of the sea otter Enhydru lutris in Canada Cunudiun Field- Nuturcrlist 101 279-83

McDonald J N 1981 North American Bison Their Clascicution and Evolution Berkeley University of California Press

Mcltzcr D J 1995 Clocking the first Americans Annual Review of Anthropology 24 71-45

124 R Lee E y n ~ u n

OBricn S J and Mays E 1991 Burcauc1atic mischief rccogni~ing enciangcrcd spccics and subspecies Scicncc 251 1 187--8

Ogden A 1933 Russian sea ottcr anal seal hunting on the Califorilia coast Caij(~rrrirrNisroricci1 Qmcrrrerly 12 29-5 1

Olscn S I and Olscn I W 1981 A conin-icnt on nomcnclaturc in faunal studies iui~cric~un Antiquity 46 192-4

Peck J M Miclucllc 14 C and Wright M 61987 Arc bison cuotic in the Wrangcll-St Elia National Park and Pre~cr~c Envivnnnlcrzrcrl iWrlrlcrgcineilr I I l(195 3

Quinn J Iand ifarrison S P 1988 Effects of habitat fragmentation anhi isolation on spci-is richness evidence from biogeographic patterns Oecologiii 75 132-40

Rasmusson E M 1985 El Nino and asiations in climate Atzcricr~z Scic3nritt 73 168-77

Raup D M 1984 Death of spccics I n Errirrcrions (etl M 14 Nitccllti) dlhicago University of Chicago Press pp 1-19

IcnsbcrgcrJ M and Barnosky A I) 1993 Short-term fluctuations in slniill mammals ot the lato Pleistocene from eastern Washington Wn iWorphologicrrl Chr~rigc irr C)iir~t~rricrr~ Mrrinnzcrls (J ~ i i~ t l~ Alzcrica (eds R A Martin and 4 Igt 131rnosky) Qambridgc Ciimbsidgc University Prcss pp 299-342

Richards M Smallcy KSykcs R and Iledges BP 1993 iir-ch21eology and genetics anaiysirig DNA from skeletal remains Worlrl~lrchtrcolog~5 18-28

Richards M B Sykcs B C and Hedges MF M 1995 Authenticating D N A extriicted Irorri ancient skeletal remains Iourr7crl of Arcl~aeologicril Scicncc 22 2W-9

Faohison ND 1978 Zooarchaeology its hiirto~y and dcvclopnirn~7r7rrcssrc Anthropologic~ul A~orirition Miscellancotrs Pripcr 2 1-2

Rocst A 1 1973 Subspecies of the sea ottcs Er7llytira Iiitris Io5 ii7gcles Coirnty Nnturcrl tiisrory Mltscirrn (ontributior~s in Scicncr 252

Rocst A 1 1979 A re-evaluation oi sea o t t c ~ taxonomy Paper prcscntcci at the Sea Ottcr Workshop Santa Barbara Cilifornia

Scheffer V B 1993 Tlie Olympic Mo~rntain goat conrrovcrs) a pcr5pectivc Coriscrvtrtior Biology 7 916-19

Sclicffcr V B and Wilkc F 1950 Validity of the subspccics E~llzydrii 1iltris r7crcic the southcrn s ix ottcr Joltvnill qf the Wcr~I7ir7gton Accrtlcuny 01 S~icr~cr402672

Simons I) R 1083 Interactions lsctween Calilornin corrdors and humans in prehistoric far wcstern North America In Vcrltcrrr Biology und ~tlunuget~ierlt (cds S KWilbur and J A Jackson) Berkeley University or Ciliforni~i Press pp 470-91

Steadman D xi 1995 Prehistoric cstinctions of Pacific isiarld birds biodiversity nlccts zooarchacology Scieilce 267 1 123-3 1

Trillrnich F and Limbcrger D 1985 Drastic effects of 871 Nirio on Ga1apigos piniiipcd Oecologiu 67 19-22

VanBlaricorn 6Rand Estcs 9 A (cds) 1988 l h p Bot~irnlrniij bcologj oj Seu O t r e r ~ (Berlin

Springer-Verliig

Wagner F 11 Forest RC i i l l R R Mcdullough 14 R Pcltoil M MPorTcr W F and Salwasscr 13 1995 WildlifePolicies in the U S Nirrionul Purks Washington DC Island Prcsi

Weisler M I and Gargett R H 1993 Pacific island avian cstinctions the taphonorny or human predation Arclzucoloyj in Occuniu 28 85-03

Wendell F E flardy R A An-rc5 1 land Burgi K T 1986 Tcrnporalanci spatial patterns in

The relevance of faunal analjgtsis to wildlife management 119

in the past surely will help in locating future preserves And as Graham and Graham (1994) indicate detailed knowledge of the historical and ecological biogeography of a taxon could prove critical to reconstructing appropriate habitats for reintroduced populations of the taxon Such knowledge can often come only from the fossil record

Extinction -of populations of species of lineages - is a natural fact of life (eg Raup 1984) But ignorance resulted in the loss of the passenger pigeon (Ectopistes migratorius) from North America the Great Auk (Pingninns impennis) from the North Atlantic and the dodo (Raphus cuci~llatus) from Mauritius in the Indian Ocean Our often stumbling attempts at wildlife management may due to ignorance ultimately lead to other such losses (eg Graham 1988 Hess 1993 Kay 1994) Ihe international move to maintain biological diversity - not just the taxonomic richness of biotic communities but the particular structure of those communities - seems to be in the correct direction if we are to maintain some quality of life and wilderness Understanding the loss of diversity often demands zooarchaeological and related data (eg Culotta 1995 Steadman 1995) and will help us avoid future inadvertent and unintentional losses Diamonds (1984) Rosetta Stone of using knowledge of modern extinctions to understand prehistoric extinctions could and perhaps should be turned on its head

Discussion

Nearly eighty years ago archaeologist W J Wintemberg (1919) noted how zooarchaeo- logical research could contribute to zoological knowledge and indirectly wildlife management His comments were repeated and elaborated thirty years later by biologist R M Gilmore (1949) Their remarks have largely gone unheeded perhaps because like Legge (1978 129 130) many of us see our job as aiming to contribute to the sum of the archaeological knowledge and that bone data are archaeological data I agree with Legge but also with Wintemberg and Gilmore we should be more than just consumers of wildlife data (cf Will 1982)

The potential topics under the broad umbrella of wildlife management to which zooarchaeological research might contribute are numerous I have touched on only a fevv of the more readily identifiable ones above There are to be sure others of some significance For example the modern fisheries industry of the eastern Pacific Ocean has been disrupted several times in the recent past by major changes in atmospheric circulation patterns causing alteration of ocean currents These events - variously referred to as El Nifio or Southern Oscillation events -appear to have some periodicity (Rasmusson 1985) but the historic record is of limited utility in plotting the precise nature of that periodicity These events clearly influence much of the oceanic biota (eg Arnold and Tissot 1993 Trillmich and Limberger 1985) Given sufficient temporal resolution the zooarchaeologi- cal record thus may eventually provide the time depth necessary to clarify the periodicity and magnitude of El Nifio events and thus allow more accurate predictions of future events

Zooarchaeological research has also shown that humans have long had a hand in modifying ecosystems Based on his extensive research on the prehistoric extinctions of Pacific Island bird populations and species Steadman (1995 1130) concludes that humans

tilay find Yt difficult to manage scarce resources in a sustainable way That humans contributed to these extinctions seems clear (cg Weisler and Cargett 1993) and thus Steadmans conclusion has the proper ominous tone But zooarchaeological research cair also perhaps help us out of such situations The wapiti (Cervns elaphus) of Yellowstone National Park in the United States are an example Herc nearly 60000 wapiti are managed and to a significant degree protected from human predation However historic and zooarchaeological research indicates that this population is a function of modern wildlife management historic records virtually vithout exception fail to mention wapiti i l l

the area and zooarchaeological data indicate this large ungulate was very riirc there during the last several thousand years (Kay 1994) Clearly if the Yellowstone ecosSsten1 is to be managed and maintained as a pristine wilderness in perpetuity (a laudable but in reality unattainable goal see Wager et al 1995) things must change before the ecosystem is irreparably altered A virtually identical situation exists in Rocky Mountain National Park sotile 550 km southwest of Yellowstone (Hcss 1993) where again zooarchaeologicai and other data from prehistoric times might help rcsolve debates

The few examples of how zooarchaeological research can be applied to t i ioder~~ problems I have outlined are rather specific to a place andlor to a taxon I do not perccive that to bc a weakness of an applied zooarchaeology but rather as an intellectual challenge Broadening the scope of zooarchaeology in the manner in which 1 have discussed will not be easy but i t will be beneficial to our Suture not only from the perspective of contributing to ensuring the preservation of biological diversity for future generations but also as 1conclude below from the perspective of our discipline which tilight otherwise become increasingly perceived as the pursuit of esoteric knotvledge of little practical use

One might argue that the discussion here could be expanded by suggesting that the detection of animal residues such as blood and hair (eg Loy 1993) could also providr information on the taxonomic identity of species native to the locale where the tools were collected (eg Hyland et al 1990) I doubt however that such a study would convince wildlife managers or all zooarchaeologists that a species was native given that stone tools were often transported significant distances by their tiiakers (eg Meltzer 1995) 1 beiievc multiple instances of such residues from multiple sites would be necessary to build 21

strong argument A partial skeleton or selected isolated bones and teeth of the species in que5tion recovered from a s~ngle ~ t e uould probably be muclr more conblnclng (Lyman 1994b)

I have attempted here to describe the structure of an applied zooarchaeology This is not a plea to legitimize what some of us do for fun andlor a living Rather i t is an attempt to broaden the scope of zooarchaeological research 1 am all for research for the sake of research if there is a problem in need of solving or a question in need of an answer But B am also concerned with the future of zooarchaeology Too many of our new doctoral str~dcnts are finding that the job market is closed and too marly of us arc finding it increasingly difficult to secure funding for research Perhaps a developed applied zooarchaeology wil

The relevance o f faunal analysis t o wildlife rnanagement 121

provide a new job market and new sources of funding It seems self-evident to me that if a national park for example has a staff archaeologist who deals with the archaeological resources on park lands and has a staff biologist who deals with biological resources on park lands then a marriage of the two would benefit both However given the increasing specialization of both archaeology and biology a viable marriage may not be possible Ihc obvious alternative is a park staff zooarchaeologist who has a deep knowledge ancl understanding of both disciplines and has been trained to integrate the two

New sources of funding may also become available if we can convince the wildlife management community that because its decisions and policies are typically aimed at tht future knowing something of the past can result in wise decisions That knowledge will of course come with a price tag The cost we may encounter is that members of the wildlife management community -while they may well have a solid knowledge of the time depth of evolutionary processes- often tend to view biotas as static ever enduring entities That is biotas have no past or at least their history is insignificant because they are being managed for the future While we certainly cannot predict exactly when the next ice age or glacial period will begin we can I think argue convincingly that climates and environments will change and knowing this we can use the prehistoric record to test our predictions about how certain kinds of changes may affect biotas of the future And confirmed - or even rejected or falsified -predictions would be the strong selling point Federal land-managing agencies whose charge includes wildlife management might well pay a zooarchaeologist to help them make wise management decisions The other selling point is that a less desirable kind of price -ecosystem destruction and loss of biodiversity -might otherwise have to be paid

Acknowledgements

I thank Donald K Grayson Adrian Lister Michael J OBrien and Ken Ihornas for comments on earlier versions of the manuscript

Departtnent o f Anthropology University o f Missouri-Columbia

References

Anonymous 1992 Draft guidclincs for rcintroductions hZws1etttr of the ILJCN feintvorirlctiorr Specinlist Group 4 2-3

Anunscn C S and Anunsen R 1993 Rcsponsc to Sclleffer Conservation Biology 7 054-7

Arnold J E andTissot B N 1993 Measurcmcnt of significant rnarine palcotcmperature variation using black abalone shells from prehistoric middens Quaternurj~ Iesearch 39 3901

Brewer D J 1992 Zooarchacology method tlleory and goals In Archaeological Metl~od and Theory Vol 4 (ed M B Schiffer) Tucson University of Arizona Press pp 195-244

Brown J H 1971 Mammals on mountaintops nonequilibrium insular biogcograplly American Vr~turulist105 467-78

Brown J PI 1978 The theory of insular biogeography and the distribution of boreal mammals and birds In Intermountain Biogeogrur~hy A Synzposium (eds KT Harper and J L Rcvcal) Great Basin Natural Memoir 2 pp 209-28

Brown J H 1986 Two decades of interaction between the MacArthiu-Wilson model and tile complexities of mammalian distributions Biological Journal d t h c Lirlnearl Society 28 23151

Cookc H B S 1976 Suidac from Plio-Pleistocene strata of the Rudolf Basin In Earlicst Mair a ~ r d Environmcrlts in the Lakc Rlrdolf Basin Stratigraphy Paleoecology and Evolutioir (cds Y Coppcns F C Flowcll G LI Isaac and RE F Lcakey) Chicago Univcrsity of Cllicago Press pp 25 1-63

Culotta E 1995 Many suspects to blame in Madagascar cxtinctions Science 268 1568-9

Davis J and Lidicker W Z Jr 1975 Tlle taxonomic status of tllc soutl~ern sea otter P r o c ~ ~ d i i z g oJtlze Cr~lijorrliri Acriderny o f Sciences 40(14) 429-37

Diamond J M 1984 Historic extinctions a Rosetta Stone for understanding prcllistoric extinctions in Quuterrlury Extir1ctions A Prehistoric Revolution (eds P S Martin and R 6Klcin) Tucson University of Arizona Prcss pp 824-62

Emslic S D 1987 Age and diet of fossil California condors in Grand Canyon Arizona Science 237 768-70

Fisller J W Jr 1995 Bone surface modifications in 7ooarchaeology Journal ofArchacologica1 Method rind Theory 2 7-68

Gifford-Gonzalez D 1991 Bones arc not enough analogues knowledge and intcrprctivc strategies in zooarchaeology Journal ofAntlzropologica1 Archrreologj 10 215-54

Gilmorc R M 1949 Tllc identification and value of mammal bones from archeological excavations Journal of Mummulogy 30 163-9

Graham RW 1988 The role of climatic change in the design of biological reserves tho palcoccological perspccti~rc for conser~ration biology Corltervution Biologjl 2 391-4

Graham RW and Graham M A 1994 Late Quaternary distribution of Marter in North America In Mcrrten~ Sublcs urtd Fislrcrs Biology and Conservation (eds S W Ruskirk 14 S Harestad M 6Raphael and RA Powell) Itllaca NY Corncll University Prcss pp 2 6 5 8

Grayson D M 1991 Tllc biogeographic history of small mammals in the Great Basin observations on thc last 20000 years Journal of Mammalogy 68 359-75

Green J S and Flinders J T 1980a Brachylrig~s idahoerisis Manznzalian Specics 125 1--4

Green J S and Flindcrs J T 1980b Habitat and dietary relationships of the pygmy rabbit dournnl Kringe Marlrigernent 33 13amp42

I3css K J r 1993 Rocky Times in Rocky Mourltairl hcrtiorzril Park Niwot Colo University Prcss of Colorado

Flestcr F E 1991 The U S National Park Scrvicc experience with exotic species Nriturril Areri ~ Jourr~ril1 1 127-8

Hoerr W 1993 The concept of naturalness in environmental tliscoursc Natural Areas Journal 1329-32

Ilouston D B and Schrciner E 6 1995 Alien specie in national parks drawing lines in space and time Conservation Biology 9 204-9

IHouston D B Schrciner E 6and Moorhcad R R 1994 Mourltairl Goats in Olympic Nutionul Park Biologjl and Marlugernerlt of an Introduced Species U S National Park Scrvicc Scientific Monograph NPSINROLYMINRSM-9325

I-Iudson J (cd) 1993 From Bones to Bc~lzavior Etlznotrrchaeologicul and E~plcrimcntill Coiltri- butiorls ro rhe Intcrprc~tation of Fblrr~al liernairls Carbondalc Southern Illinois University Center for Archaeological Investigations Occasional Papcr No 21

T h e relevance of faunal analysis t o wildlife management 123

Hyland D C Tersak J M Adovasio J M and Sicgal M I 1990 Identification of thc speciesof origin of residual blood on lithic material Americntl Antiquity 55 104-12

Jameson R J Kenyon K W Johnson A M and Wight H M 1982 History and status of translocated sea otter populations in North America Wildlife Society Bulletin 10 100-7

Jameson R J Kenyon K W Jeffries S and VanBlaricom G R 1986 Status of a I ranslocatcd sea otter population and its habitat in Washington The Murrelet 67 84-7

Kay C E 1994 Aboriginal overkill the role of Native Americans in structur~ng western ecosystems Hurnnt~ Nrrrure 5 359-98

Kenyon K W 1969 The sea otter In the eastern Pacific Ocean Worth Amer~crrtz Firuna 68

Korn H 1994 Genetic demographic spatial environmental and catastrophic effects on the survival probability of small populations of mammals In Mitlimum Animrrl Popu1rrrior1s (ed H Remmert) Berlin Springer-Verlag pp 3)-49

Lcggc A J 1978 Archacozoology - or zooarchacology In Research Prohlems in Zoourchueology (eds D R Brothwell K D Thomas and J Clutton-Brock) Univcrsity of London Institute of Archaeology Occasional Publications No 3 pp 129-32

Leopold A S Cain S A Cottam C IM Gabrielson I N and Kimhall T L 1963 Wildlife management in the national parks Trunsuctions of the Twenty-Eighth North American Wildlife und Nutural Resoirrce~ Cd70nferencepp 2945

Loy T H 1993 The artifact as site an example of the biomolccular analysis of organic residues on prehistoric tools World Archaeology 25 44-63

Lyman R L 1988a Significance for wildlife management of the late Quaternary biogeography of mountain goats (Oreurnnos urnericunus) in the Pacific Northwest USA Arctic and Alpine Reseurch 20 13-23

Lyman R L 1988b Zoogeography of Oregon coast marine mammals the last 3000 years Murine Murnrnul Science 4 247-64

Lyman R L 1991 Late Quaternary biogeography of the pygmy rabbit (Brachylagirs iduhoensis) in eastern Washington Jo~lrnalof Mammalogy 72 11amp17

Lyman R L 1994a The Olympic Mountain goat controversy a different perspective Conservation Biology 8 898-901

Lyman R L 1994b Vertebrute Taphonomy Cambridge Cambridge University Press

Lyman R L 1995a Determining when rare (zoo)archacological phenomena are truly absent Journal of Archueologicul Method unrl Theory 2 369-424

Lyman R L 199 Inaccurate data and the Olympic National Park mountain goat controversy North wect Science 69 234-8

McAllister K and Allen H 1993 Stutuc of the Pygmy Rabbit (Brachylagus idahoensis) in Wushington Olympia Washington Department of Wildlife Wildlife Management Division

McAllister K and Allen H 1994 Wushington Stute Draft Recovery Plun for the Pygmy Rabbit (Brachylagus idahocnsis) Olympia Washington Department of Wildlife Wildlife Management Division

MacArthur R H and Wilson E 0 1967 The Theory of Islunci Biogeography Princeton NJ Princeton University Press

MacAskie I 1987 Updated status of the sea otter Enhydru lutris in Canada Cunudiun Field- Nuturcrlist 101 279-83

McDonald J N 1981 North American Bison Their Clascicution and Evolution Berkeley University of California Press

Mcltzcr D J 1995 Clocking the first Americans Annual Review of Anthropology 24 71-45

124 R Lee E y n ~ u n

OBricn S J and Mays E 1991 Burcauc1atic mischief rccogni~ing enciangcrcd spccics and subspecies Scicncc 251 1 187--8

Ogden A 1933 Russian sea ottcr anal seal hunting on the Califorilia coast Caij(~rrrirrNisroricci1 Qmcrrrerly 12 29-5 1

Olscn S I and Olscn I W 1981 A conin-icnt on nomcnclaturc in faunal studies iui~cric~un Antiquity 46 192-4

Peck J M Miclucllc 14 C and Wright M 61987 Arc bison cuotic in the Wrangcll-St Elia National Park and Pre~cr~c Envivnnnlcrzrcrl iWrlrlcrgcineilr I I l(195 3

Quinn J Iand ifarrison S P 1988 Effects of habitat fragmentation anhi isolation on spci-is richness evidence from biogeographic patterns Oecologiii 75 132-40

Rasmusson E M 1985 El Nino and asiations in climate Atzcricr~z Scic3nritt 73 168-77

Raup D M 1984 Death of spccics I n Errirrcrions (etl M 14 Nitccllti) dlhicago University of Chicago Press pp 1-19

IcnsbcrgcrJ M and Barnosky A I) 1993 Short-term fluctuations in slniill mammals ot the lato Pleistocene from eastern Washington Wn iWorphologicrrl Chr~rigc irr C)iir~t~rricrr~ Mrrinnzcrls (J ~ i i~ t l~ Alzcrica (eds R A Martin and 4 Igt 131rnosky) Qambridgc Ciimbsidgc University Prcss pp 299-342

Richards M Smallcy KSykcs R and Iledges BP 1993 iir-ch21eology and genetics anaiysirig DNA from skeletal remains Worlrl~lrchtrcolog~5 18-28

Richards M B Sykcs B C and Hedges MF M 1995 Authenticating D N A extriicted Irorri ancient skeletal remains Iourr7crl of Arcl~aeologicril Scicncc 22 2W-9

Faohison ND 1978 Zooarchaeology its hiirto~y and dcvclopnirn~7r7rrcssrc Anthropologic~ul A~orirition Miscellancotrs Pripcr 2 1-2

Rocst A 1 1973 Subspecies of the sea ottcs Er7llytira Iiitris Io5 ii7gcles Coirnty Nnturcrl tiisrory Mltscirrn (ontributior~s in Scicncr 252

Rocst A 1 1979 A re-evaluation oi sea o t t c ~ taxonomy Paper prcscntcci at the Sea Ottcr Workshop Santa Barbara Cilifornia

Scheffer V B 1993 Tlie Olympic Mo~rntain goat conrrovcrs) a pcr5pectivc Coriscrvtrtior Biology 7 916-19

Sclicffcr V B and Wilkc F 1950 Validity of the subspccics E~llzydrii 1iltris r7crcic the southcrn s ix ottcr Joltvnill qf the Wcr~I7ir7gton Accrtlcuny 01 S~icr~cr402672

Simons I) R 1083 Interactions lsctween Calilornin corrdors and humans in prehistoric far wcstern North America In Vcrltcrrr Biology und ~tlunuget~ierlt (cds S KWilbur and J A Jackson) Berkeley University or Ciliforni~i Press pp 470-91

Steadman D xi 1995 Prehistoric cstinctions of Pacific isiarld birds biodiversity nlccts zooarchacology Scieilce 267 1 123-3 1

Trillrnich F and Limbcrger D 1985 Drastic effects of 871 Nirio on Ga1apigos piniiipcd Oecologiu 67 19-22

VanBlaricorn 6Rand Estcs 9 A (cds) 1988 l h p Bot~irnlrniij bcologj oj Seu O t r e r ~ (Berlin

Springer-Verliig

Wagner F 11 Forest RC i i l l R R Mcdullough 14 R Pcltoil M MPorTcr W F and Salwasscr 13 1995 WildlifePolicies in the U S Nirrionul Purks Washington DC Island Prcsi

Weisler M I and Gargett R H 1993 Pacific island avian cstinctions the taphonorny or human predation Arclzucoloyj in Occuniu 28 85-03

Wendell F E flardy R A An-rc5 1 land Burgi K T 1986 Tcrnporalanci spatial patterns in

tilay find Yt difficult to manage scarce resources in a sustainable way That humans contributed to these extinctions seems clear (cg Weisler and Cargett 1993) and thus Steadmans conclusion has the proper ominous tone But zooarchaeological research cair also perhaps help us out of such situations The wapiti (Cervns elaphus) of Yellowstone National Park in the United States are an example Herc nearly 60000 wapiti are managed and to a significant degree protected from human predation However historic and zooarchaeological research indicates that this population is a function of modern wildlife management historic records virtually vithout exception fail to mention wapiti i l l

the area and zooarchaeological data indicate this large ungulate was very riirc there during the last several thousand years (Kay 1994) Clearly if the Yellowstone ecosSsten1 is to be managed and maintained as a pristine wilderness in perpetuity (a laudable but in reality unattainable goal see Wager et al 1995) things must change before the ecosystem is irreparably altered A virtually identical situation exists in Rocky Mountain National Park sotile 550 km southwest of Yellowstone (Hcss 1993) where again zooarchaeologicai and other data from prehistoric times might help rcsolve debates

The few examples of how zooarchaeological research can be applied to t i ioder~~ problems I have outlined are rather specific to a place andlor to a taxon I do not perccive that to bc a weakness of an applied zooarchaeology but rather as an intellectual challenge Broadening the scope of zooarchaeology in the manner in which 1 have discussed will not be easy but i t will be beneficial to our Suture not only from the perspective of contributing to ensuring the preservation of biological diversity for future generations but also as 1conclude below from the perspective of our discipline which tilight otherwise become increasingly perceived as the pursuit of esoteric knotvledge of little practical use

One might argue that the discussion here could be expanded by suggesting that the detection of animal residues such as blood and hair (eg Loy 1993) could also providr information on the taxonomic identity of species native to the locale where the tools were collected (eg Hyland et al 1990) I doubt however that such a study would convince wildlife managers or all zooarchaeologists that a species was native given that stone tools were often transported significant distances by their tiiakers (eg Meltzer 1995) 1 beiievc multiple instances of such residues from multiple sites would be necessary to build 21

strong argument A partial skeleton or selected isolated bones and teeth of the species in que5tion recovered from a s~ngle ~ t e uould probably be muclr more conblnclng (Lyman 1994b)

I have attempted here to describe the structure of an applied zooarchaeology This is not a plea to legitimize what some of us do for fun andlor a living Rather i t is an attempt to broaden the scope of zooarchaeological research 1 am all for research for the sake of research if there is a problem in need of solving or a question in need of an answer But B am also concerned with the future of zooarchaeology Too many of our new doctoral str~dcnts are finding that the job market is closed and too marly of us arc finding it increasingly difficult to secure funding for research Perhaps a developed applied zooarchaeology wil

The relevance o f faunal analysis t o wildlife rnanagement 121

provide a new job market and new sources of funding It seems self-evident to me that if a national park for example has a staff archaeologist who deals with the archaeological resources on park lands and has a staff biologist who deals with biological resources on park lands then a marriage of the two would benefit both However given the increasing specialization of both archaeology and biology a viable marriage may not be possible Ihc obvious alternative is a park staff zooarchaeologist who has a deep knowledge ancl understanding of both disciplines and has been trained to integrate the two

New sources of funding may also become available if we can convince the wildlife management community that because its decisions and policies are typically aimed at tht future knowing something of the past can result in wise decisions That knowledge will of course come with a price tag The cost we may encounter is that members of the wildlife management community -while they may well have a solid knowledge of the time depth of evolutionary processes- often tend to view biotas as static ever enduring entities That is biotas have no past or at least their history is insignificant because they are being managed for the future While we certainly cannot predict exactly when the next ice age or glacial period will begin we can I think argue convincingly that climates and environments will change and knowing this we can use the prehistoric record to test our predictions about how certain kinds of changes may affect biotas of the future And confirmed - or even rejected or falsified -predictions would be the strong selling point Federal land-managing agencies whose charge includes wildlife management might well pay a zooarchaeologist to help them make wise management decisions The other selling point is that a less desirable kind of price -ecosystem destruction and loss of biodiversity -might otherwise have to be paid

Acknowledgements

I thank Donald K Grayson Adrian Lister Michael J OBrien and Ken Ihornas for comments on earlier versions of the manuscript

Departtnent o f Anthropology University o f Missouri-Columbia

References

Anonymous 1992 Draft guidclincs for rcintroductions hZws1etttr of the ILJCN feintvorirlctiorr Specinlist Group 4 2-3

Anunscn C S and Anunsen R 1993 Rcsponsc to Sclleffer Conservation Biology 7 054-7

Arnold J E andTissot B N 1993 Measurcmcnt of significant rnarine palcotcmperature variation using black abalone shells from prehistoric middens Quaternurj~ Iesearch 39 3901

Brewer D J 1992 Zooarchacology method tlleory and goals In Archaeological Metl~od and Theory Vol 4 (ed M B Schiffer) Tucson University of Arizona Press pp 195-244

Brown J H 1971 Mammals on mountaintops nonequilibrium insular biogcograplly American Vr~turulist105 467-78

Brown J PI 1978 The theory of insular biogeography and the distribution of boreal mammals and birds In Intermountain Biogeogrur~hy A Synzposium (eds KT Harper and J L Rcvcal) Great Basin Natural Memoir 2 pp 209-28

Brown J H 1986 Two decades of interaction between the MacArthiu-Wilson model and tile complexities of mammalian distributions Biological Journal d t h c Lirlnearl Society 28 23151

Cookc H B S 1976 Suidac from Plio-Pleistocene strata of the Rudolf Basin In Earlicst Mair a ~ r d Environmcrlts in the Lakc Rlrdolf Basin Stratigraphy Paleoecology and Evolutioir (cds Y Coppcns F C Flowcll G LI Isaac and RE F Lcakey) Chicago Univcrsity of Cllicago Press pp 25 1-63

Culotta E 1995 Many suspects to blame in Madagascar cxtinctions Science 268 1568-9

Davis J and Lidicker W Z Jr 1975 Tlle taxonomic status of tllc soutl~ern sea otter P r o c ~ ~ d i i z g oJtlze Cr~lijorrliri Acriderny o f Sciences 40(14) 429-37

Diamond J M 1984 Historic extinctions a Rosetta Stone for understanding prcllistoric extinctions in Quuterrlury Extir1ctions A Prehistoric Revolution (eds P S Martin and R 6Klcin) Tucson University of Arizona Prcss pp 824-62

Emslic S D 1987 Age and diet of fossil California condors in Grand Canyon Arizona Science 237 768-70

Fisller J W Jr 1995 Bone surface modifications in 7ooarchaeology Journal ofArchacologica1 Method rind Theory 2 7-68

Gifford-Gonzalez D 1991 Bones arc not enough analogues knowledge and intcrprctivc strategies in zooarchaeology Journal ofAntlzropologica1 Archrreologj 10 215-54

Gilmorc R M 1949 Tllc identification and value of mammal bones from archeological excavations Journal of Mummulogy 30 163-9

Graham RW 1988 The role of climatic change in the design of biological reserves tho palcoccological perspccti~rc for conser~ration biology Corltervution Biologjl 2 391-4

Graham RW and Graham M A 1994 Late Quaternary distribution of Marter in North America In Mcrrten~ Sublcs urtd Fislrcrs Biology and Conservation (eds S W Ruskirk 14 S Harestad M 6Raphael and RA Powell) Itllaca NY Corncll University Prcss pp 2 6 5 8

Grayson D M 1991 Tllc biogeographic history of small mammals in the Great Basin observations on thc last 20000 years Journal of Mammalogy 68 359-75

Green J S and Flinders J T 1980a Brachylrig~s idahoerisis Manznzalian Specics 125 1--4

Green J S and Flindcrs J T 1980b Habitat and dietary relationships of the pygmy rabbit dournnl Kringe Marlrigernent 33 13amp42

I3css K J r 1993 Rocky Times in Rocky Mourltairl hcrtiorzril Park Niwot Colo University Prcss of Colorado

Flestcr F E 1991 The U S National Park Scrvicc experience with exotic species Nriturril Areri ~ Jourr~ril1 1 127-8

Hoerr W 1993 The concept of naturalness in environmental tliscoursc Natural Areas Journal 1329-32

Ilouston D B and Schrciner E 6 1995 Alien specie in national parks drawing lines in space and time Conservation Biology 9 204-9

IHouston D B Schrciner E 6and Moorhcad R R 1994 Mourltairl Goats in Olympic Nutionul Park Biologjl and Marlugernerlt of an Introduced Species U S National Park Scrvicc Scientific Monograph NPSINROLYMINRSM-9325

I-Iudson J (cd) 1993 From Bones to Bc~lzavior Etlznotrrchaeologicul and E~plcrimcntill Coiltri- butiorls ro rhe Intcrprc~tation of Fblrr~al liernairls Carbondalc Southern Illinois University Center for Archaeological Investigations Occasional Papcr No 21

T h e relevance of faunal analysis t o wildlife management 123

Hyland D C Tersak J M Adovasio J M and Sicgal M I 1990 Identification of thc speciesof origin of residual blood on lithic material Americntl Antiquity 55 104-12

Jameson R J Kenyon K W Johnson A M and Wight H M 1982 History and status of translocated sea otter populations in North America Wildlife Society Bulletin 10 100-7

Jameson R J Kenyon K W Jeffries S and VanBlaricom G R 1986 Status of a I ranslocatcd sea otter population and its habitat in Washington The Murrelet 67 84-7

Kay C E 1994 Aboriginal overkill the role of Native Americans in structur~ng western ecosystems Hurnnt~ Nrrrure 5 359-98

Kenyon K W 1969 The sea otter In the eastern Pacific Ocean Worth Amer~crrtz Firuna 68

Korn H 1994 Genetic demographic spatial environmental and catastrophic effects on the survival probability of small populations of mammals In Mitlimum Animrrl Popu1rrrior1s (ed H Remmert) Berlin Springer-Verlag pp 3)-49

Lcggc A J 1978 Archacozoology - or zooarchacology In Research Prohlems in Zoourchueology (eds D R Brothwell K D Thomas and J Clutton-Brock) Univcrsity of London Institute of Archaeology Occasional Publications No 3 pp 129-32

Leopold A S Cain S A Cottam C IM Gabrielson I N and Kimhall T L 1963 Wildlife management in the national parks Trunsuctions of the Twenty-Eighth North American Wildlife und Nutural Resoirrce~ Cd70nferencepp 2945

Loy T H 1993 The artifact as site an example of the biomolccular analysis of organic residues on prehistoric tools World Archaeology 25 44-63

Lyman R L 1988a Significance for wildlife management of the late Quaternary biogeography of mountain goats (Oreurnnos urnericunus) in the Pacific Northwest USA Arctic and Alpine Reseurch 20 13-23

Lyman R L 1988b Zoogeography of Oregon coast marine mammals the last 3000 years Murine Murnrnul Science 4 247-64

Lyman R L 1991 Late Quaternary biogeography of the pygmy rabbit (Brachylagirs iduhoensis) in eastern Washington Jo~lrnalof Mammalogy 72 11amp17

Lyman R L 1994a The Olympic Mountain goat controversy a different perspective Conservation Biology 8 898-901

Lyman R L 1994b Vertebrute Taphonomy Cambridge Cambridge University Press

Lyman R L 1995a Determining when rare (zoo)archacological phenomena are truly absent Journal of Archueologicul Method unrl Theory 2 369-424

Lyman R L 199 Inaccurate data and the Olympic National Park mountain goat controversy North wect Science 69 234-8

McAllister K and Allen H 1993 Stutuc of the Pygmy Rabbit (Brachylagus idahoensis) in Wushington Olympia Washington Department of Wildlife Wildlife Management Division

McAllister K and Allen H 1994 Wushington Stute Draft Recovery Plun for the Pygmy Rabbit (Brachylagus idahocnsis) Olympia Washington Department of Wildlife Wildlife Management Division

MacArthur R H and Wilson E 0 1967 The Theory of Islunci Biogeography Princeton NJ Princeton University Press

MacAskie I 1987 Updated status of the sea otter Enhydru lutris in Canada Cunudiun Field- Nuturcrlist 101 279-83

McDonald J N 1981 North American Bison Their Clascicution and Evolution Berkeley University of California Press

Mcltzcr D J 1995 Clocking the first Americans Annual Review of Anthropology 24 71-45

124 R Lee E y n ~ u n

OBricn S J and Mays E 1991 Burcauc1atic mischief rccogni~ing enciangcrcd spccics and subspecies Scicncc 251 1 187--8

Ogden A 1933 Russian sea ottcr anal seal hunting on the Califorilia coast Caij(~rrrirrNisroricci1 Qmcrrrerly 12 29-5 1

Olscn S I and Olscn I W 1981 A conin-icnt on nomcnclaturc in faunal studies iui~cric~un Antiquity 46 192-4

Peck J M Miclucllc 14 C and Wright M 61987 Arc bison cuotic in the Wrangcll-St Elia National Park and Pre~cr~c Envivnnnlcrzrcrl iWrlrlcrgcineilr I I l(195 3

Quinn J Iand ifarrison S P 1988 Effects of habitat fragmentation anhi isolation on spci-is richness evidence from biogeographic patterns Oecologiii 75 132-40

Rasmusson E M 1985 El Nino and asiations in climate Atzcricr~z Scic3nritt 73 168-77

Raup D M 1984 Death of spccics I n Errirrcrions (etl M 14 Nitccllti) dlhicago University of Chicago Press pp 1-19

IcnsbcrgcrJ M and Barnosky A I) 1993 Short-term fluctuations in slniill mammals ot the lato Pleistocene from eastern Washington Wn iWorphologicrrl Chr~rigc irr C)iir~t~rricrr~ Mrrinnzcrls (J ~ i i~ t l~ Alzcrica (eds R A Martin and 4 Igt 131rnosky) Qambridgc Ciimbsidgc University Prcss pp 299-342

Richards M Smallcy KSykcs R and Iledges BP 1993 iir-ch21eology and genetics anaiysirig DNA from skeletal remains Worlrl~lrchtrcolog~5 18-28

Richards M B Sykcs B C and Hedges MF M 1995 Authenticating D N A extriicted Irorri ancient skeletal remains Iourr7crl of Arcl~aeologicril Scicncc 22 2W-9

Faohison ND 1978 Zooarchaeology its hiirto~y and dcvclopnirn~7r7rrcssrc Anthropologic~ul A~orirition Miscellancotrs Pripcr 2 1-2

Rocst A 1 1973 Subspecies of the sea ottcs Er7llytira Iiitris Io5 ii7gcles Coirnty Nnturcrl tiisrory Mltscirrn (ontributior~s in Scicncr 252

Rocst A 1 1979 A re-evaluation oi sea o t t c ~ taxonomy Paper prcscntcci at the Sea Ottcr Workshop Santa Barbara Cilifornia

Scheffer V B 1993 Tlie Olympic Mo~rntain goat conrrovcrs) a pcr5pectivc Coriscrvtrtior Biology 7 916-19

Sclicffcr V B and Wilkc F 1950 Validity of the subspccics E~llzydrii 1iltris r7crcic the southcrn s ix ottcr Joltvnill qf the Wcr~I7ir7gton Accrtlcuny 01 S~icr~cr402672

Simons I) R 1083 Interactions lsctween Calilornin corrdors and humans in prehistoric far wcstern North America In Vcrltcrrr Biology und ~tlunuget~ierlt (cds S KWilbur and J A Jackson) Berkeley University or Ciliforni~i Press pp 470-91

Steadman D xi 1995 Prehistoric cstinctions of Pacific isiarld birds biodiversity nlccts zooarchacology Scieilce 267 1 123-3 1

Trillrnich F and Limbcrger D 1985 Drastic effects of 871 Nirio on Ga1apigos piniiipcd Oecologiu 67 19-22

VanBlaricorn 6Rand Estcs 9 A (cds) 1988 l h p Bot~irnlrniij bcologj oj Seu O t r e r ~ (Berlin

Springer-Verliig

Wagner F 11 Forest RC i i l l R R Mcdullough 14 R Pcltoil M MPorTcr W F and Salwasscr 13 1995 WildlifePolicies in the U S Nirrionul Purks Washington DC Island Prcsi

Weisler M I and Gargett R H 1993 Pacific island avian cstinctions the taphonorny or human predation Arclzucoloyj in Occuniu 28 85-03

Wendell F E flardy R A An-rc5 1 land Burgi K T 1986 Tcrnporalanci spatial patterns in

The relevance o f faunal analysis t o wildlife rnanagement 121

provide a new job market and new sources of funding It seems self-evident to me that if a national park for example has a staff archaeologist who deals with the archaeological resources on park lands and has a staff biologist who deals with biological resources on park lands then a marriage of the two would benefit both However given the increasing specialization of both archaeology and biology a viable marriage may not be possible Ihc obvious alternative is a park staff zooarchaeologist who has a deep knowledge ancl understanding of both disciplines and has been trained to integrate the two

New sources of funding may also become available if we can convince the wildlife management community that because its decisions and policies are typically aimed at tht future knowing something of the past can result in wise decisions That knowledge will of course come with a price tag The cost we may encounter is that members of the wildlife management community -while they may well have a solid knowledge of the time depth of evolutionary processes- often tend to view biotas as static ever enduring entities That is biotas have no past or at least their history is insignificant because they are being managed for the future While we certainly cannot predict exactly when the next ice age or glacial period will begin we can I think argue convincingly that climates and environments will change and knowing this we can use the prehistoric record to test our predictions about how certain kinds of changes may affect biotas of the future And confirmed - or even rejected or falsified -predictions would be the strong selling point Federal land-managing agencies whose charge includes wildlife management might well pay a zooarchaeologist to help them make wise management decisions The other selling point is that a less desirable kind of price -ecosystem destruction and loss of biodiversity -might otherwise have to be paid

Acknowledgements

I thank Donald K Grayson Adrian Lister Michael J OBrien and Ken Ihornas for comments on earlier versions of the manuscript

Departtnent o f Anthropology University o f Missouri-Columbia

References

Anonymous 1992 Draft guidclincs for rcintroductions hZws1etttr of the ILJCN feintvorirlctiorr Specinlist Group 4 2-3

Anunscn C S and Anunsen R 1993 Rcsponsc to Sclleffer Conservation Biology 7 054-7

Arnold J E andTissot B N 1993 Measurcmcnt of significant rnarine palcotcmperature variation using black abalone shells from prehistoric middens Quaternurj~ Iesearch 39 3901

Brewer D J 1992 Zooarchacology method tlleory and goals In Archaeological Metl~od and Theory Vol 4 (ed M B Schiffer) Tucson University of Arizona Press pp 195-244

Brown J H 1971 Mammals on mountaintops nonequilibrium insular biogcograplly American Vr~turulist105 467-78

Brown J PI 1978 The theory of insular biogeography and the distribution of boreal mammals and birds In Intermountain Biogeogrur~hy A Synzposium (eds KT Harper and J L Rcvcal) Great Basin Natural Memoir 2 pp 209-28

Brown J H 1986 Two decades of interaction between the MacArthiu-Wilson model and tile complexities of mammalian distributions Biological Journal d t h c Lirlnearl Society 28 23151

Cookc H B S 1976 Suidac from Plio-Pleistocene strata of the Rudolf Basin In Earlicst Mair a ~ r d Environmcrlts in the Lakc Rlrdolf Basin Stratigraphy Paleoecology and Evolutioir (cds Y Coppcns F C Flowcll G LI Isaac and RE F Lcakey) Chicago Univcrsity of Cllicago Press pp 25 1-63

Culotta E 1995 Many suspects to blame in Madagascar cxtinctions Science 268 1568-9

Davis J and Lidicker W Z Jr 1975 Tlle taxonomic status of tllc soutl~ern sea otter P r o c ~ ~ d i i z g oJtlze Cr~lijorrliri Acriderny o f Sciences 40(14) 429-37

Diamond J M 1984 Historic extinctions a Rosetta Stone for understanding prcllistoric extinctions in Quuterrlury Extir1ctions A Prehistoric Revolution (eds P S Martin and R 6Klcin) Tucson University of Arizona Prcss pp 824-62

Emslic S D 1987 Age and diet of fossil California condors in Grand Canyon Arizona Science 237 768-70

Fisller J W Jr 1995 Bone surface modifications in 7ooarchaeology Journal ofArchacologica1 Method rind Theory 2 7-68

Gifford-Gonzalez D 1991 Bones arc not enough analogues knowledge and intcrprctivc strategies in zooarchaeology Journal ofAntlzropologica1 Archrreologj 10 215-54

Gilmorc R M 1949 Tllc identification and value of mammal bones from archeological excavations Journal of Mummulogy 30 163-9

Graham RW 1988 The role of climatic change in the design of biological reserves tho palcoccological perspccti~rc for conser~ration biology Corltervution Biologjl 2 391-4

Graham RW and Graham M A 1994 Late Quaternary distribution of Marter in North America In Mcrrten~ Sublcs urtd Fislrcrs Biology and Conservation (eds S W Ruskirk 14 S Harestad M 6Raphael and RA Powell) Itllaca NY Corncll University Prcss pp 2 6 5 8

Grayson D M 1991 Tllc biogeographic history of small mammals in the Great Basin observations on thc last 20000 years Journal of Mammalogy 68 359-75

Green J S and Flinders J T 1980a Brachylrig~s idahoerisis Manznzalian Specics 125 1--4

Green J S and Flindcrs J T 1980b Habitat and dietary relationships of the pygmy rabbit dournnl Kringe Marlrigernent 33 13amp42

I3css K J r 1993 Rocky Times in Rocky Mourltairl hcrtiorzril Park Niwot Colo University Prcss of Colorado

Flestcr F E 1991 The U S National Park Scrvicc experience with exotic species Nriturril Areri ~ Jourr~ril1 1 127-8

Hoerr W 1993 The concept of naturalness in environmental tliscoursc Natural Areas Journal 1329-32

Ilouston D B and Schrciner E 6 1995 Alien specie in national parks drawing lines in space and time Conservation Biology 9 204-9

IHouston D B Schrciner E 6and Moorhcad R R 1994 Mourltairl Goats in Olympic Nutionul Park Biologjl and Marlugernerlt of an Introduced Species U S National Park Scrvicc Scientific Monograph NPSINROLYMINRSM-9325

I-Iudson J (cd) 1993 From Bones to Bc~lzavior Etlznotrrchaeologicul and E~plcrimcntill Coiltri- butiorls ro rhe Intcrprc~tation of Fblrr~al liernairls Carbondalc Southern Illinois University Center for Archaeological Investigations Occasional Papcr No 21

T h e relevance of faunal analysis t o wildlife management 123

Hyland D C Tersak J M Adovasio J M and Sicgal M I 1990 Identification of thc speciesof origin of residual blood on lithic material Americntl Antiquity 55 104-12

Jameson R J Kenyon K W Johnson A M and Wight H M 1982 History and status of translocated sea otter populations in North America Wildlife Society Bulletin 10 100-7

Jameson R J Kenyon K W Jeffries S and VanBlaricom G R 1986 Status of a I ranslocatcd sea otter population and its habitat in Washington The Murrelet 67 84-7

Kay C E 1994 Aboriginal overkill the role of Native Americans in structur~ng western ecosystems Hurnnt~ Nrrrure 5 359-98

Kenyon K W 1969 The sea otter In the eastern Pacific Ocean Worth Amer~crrtz Firuna 68

Korn H 1994 Genetic demographic spatial environmental and catastrophic effects on the survival probability of small populations of mammals In Mitlimum Animrrl Popu1rrrior1s (ed H Remmert) Berlin Springer-Verlag pp 3)-49

Lcggc A J 1978 Archacozoology - or zooarchacology In Research Prohlems in Zoourchueology (eds D R Brothwell K D Thomas and J Clutton-Brock) Univcrsity of London Institute of Archaeology Occasional Publications No 3 pp 129-32

Leopold A S Cain S A Cottam C IM Gabrielson I N and Kimhall T L 1963 Wildlife management in the national parks Trunsuctions of the Twenty-Eighth North American Wildlife und Nutural Resoirrce~ Cd70nferencepp 2945

Loy T H 1993 The artifact as site an example of the biomolccular analysis of organic residues on prehistoric tools World Archaeology 25 44-63

Lyman R L 1988a Significance for wildlife management of the late Quaternary biogeography of mountain goats (Oreurnnos urnericunus) in the Pacific Northwest USA Arctic and Alpine Reseurch 20 13-23

Lyman R L 1988b Zoogeography of Oregon coast marine mammals the last 3000 years Murine Murnrnul Science 4 247-64

Lyman R L 1991 Late Quaternary biogeography of the pygmy rabbit (Brachylagirs iduhoensis) in eastern Washington Jo~lrnalof Mammalogy 72 11amp17

Lyman R L 1994a The Olympic Mountain goat controversy a different perspective Conservation Biology 8 898-901

Lyman R L 1994b Vertebrute Taphonomy Cambridge Cambridge University Press

Lyman R L 1995a Determining when rare (zoo)archacological phenomena are truly absent Journal of Archueologicul Method unrl Theory 2 369-424

Lyman R L 199 Inaccurate data and the Olympic National Park mountain goat controversy North wect Science 69 234-8

McAllister K and Allen H 1993 Stutuc of the Pygmy Rabbit (Brachylagus idahoensis) in Wushington Olympia Washington Department of Wildlife Wildlife Management Division

McAllister K and Allen H 1994 Wushington Stute Draft Recovery Plun for the Pygmy Rabbit (Brachylagus idahocnsis) Olympia Washington Department of Wildlife Wildlife Management Division

MacArthur R H and Wilson E 0 1967 The Theory of Islunci Biogeography Princeton NJ Princeton University Press

MacAskie I 1987 Updated status of the sea otter Enhydru lutris in Canada Cunudiun Field- Nuturcrlist 101 279-83

McDonald J N 1981 North American Bison Their Clascicution and Evolution Berkeley University of California Press

Mcltzcr D J 1995 Clocking the first Americans Annual Review of Anthropology 24 71-45

124 R Lee E y n ~ u n

OBricn S J and Mays E 1991 Burcauc1atic mischief rccogni~ing enciangcrcd spccics and subspecies Scicncc 251 1 187--8

Ogden A 1933 Russian sea ottcr anal seal hunting on the Califorilia coast Caij(~rrrirrNisroricci1 Qmcrrrerly 12 29-5 1

Olscn S I and Olscn I W 1981 A conin-icnt on nomcnclaturc in faunal studies iui~cric~un Antiquity 46 192-4

Peck J M Miclucllc 14 C and Wright M 61987 Arc bison cuotic in the Wrangcll-St Elia National Park and Pre~cr~c Envivnnnlcrzrcrl iWrlrlcrgcineilr I I l(195 3

Quinn J Iand ifarrison S P 1988 Effects of habitat fragmentation anhi isolation on spci-is richness evidence from biogeographic patterns Oecologiii 75 132-40

Rasmusson E M 1985 El Nino and asiations in climate Atzcricr~z Scic3nritt 73 168-77

Raup D M 1984 Death of spccics I n Errirrcrions (etl M 14 Nitccllti) dlhicago University of Chicago Press pp 1-19

IcnsbcrgcrJ M and Barnosky A I) 1993 Short-term fluctuations in slniill mammals ot the lato Pleistocene from eastern Washington Wn iWorphologicrrl Chr~rigc irr C)iir~t~rricrr~ Mrrinnzcrls (J ~ i i~ t l~ Alzcrica (eds R A Martin and 4 Igt 131rnosky) Qambridgc Ciimbsidgc University Prcss pp 299-342

Richards M Smallcy KSykcs R and Iledges BP 1993 iir-ch21eology and genetics anaiysirig DNA from skeletal remains Worlrl~lrchtrcolog~5 18-28

Richards M B Sykcs B C and Hedges MF M 1995 Authenticating D N A extriicted Irorri ancient skeletal remains Iourr7crl of Arcl~aeologicril Scicncc 22 2W-9

Faohison ND 1978 Zooarchaeology its hiirto~y and dcvclopnirn~7r7rrcssrc Anthropologic~ul A~orirition Miscellancotrs Pripcr 2 1-2

Rocst A 1 1973 Subspecies of the sea ottcs Er7llytira Iiitris Io5 ii7gcles Coirnty Nnturcrl tiisrory Mltscirrn (ontributior~s in Scicncr 252

Rocst A 1 1979 A re-evaluation oi sea o t t c ~ taxonomy Paper prcscntcci at the Sea Ottcr Workshop Santa Barbara Cilifornia

Scheffer V B 1993 Tlie Olympic Mo~rntain goat conrrovcrs) a pcr5pectivc Coriscrvtrtior Biology 7 916-19

Sclicffcr V B and Wilkc F 1950 Validity of the subspccics E~llzydrii 1iltris r7crcic the southcrn s ix ottcr Joltvnill qf the Wcr~I7ir7gton Accrtlcuny 01 S~icr~cr402672

Simons I) R 1083 Interactions lsctween Calilornin corrdors and humans in prehistoric far wcstern North America In Vcrltcrrr Biology und ~tlunuget~ierlt (cds S KWilbur and J A Jackson) Berkeley University or Ciliforni~i Press pp 470-91

Steadman D xi 1995 Prehistoric cstinctions of Pacific isiarld birds biodiversity nlccts zooarchacology Scieilce 267 1 123-3 1

Trillrnich F and Limbcrger D 1985 Drastic effects of 871 Nirio on Ga1apigos piniiipcd Oecologiu 67 19-22

VanBlaricorn 6Rand Estcs 9 A (cds) 1988 l h p Bot~irnlrniij bcologj oj Seu O t r e r ~ (Berlin

Springer-Verliig

Wagner F 11 Forest RC i i l l R R Mcdullough 14 R Pcltoil M MPorTcr W F and Salwasscr 13 1995 WildlifePolicies in the U S Nirrionul Purks Washington DC Island Prcsi

Weisler M I and Gargett R H 1993 Pacific island avian cstinctions the taphonorny or human predation Arclzucoloyj in Occuniu 28 85-03

Wendell F E flardy R A An-rc5 1 land Burgi K T 1986 Tcrnporalanci spatial patterns in

Brown J PI 1978 The theory of insular biogeography and the distribution of boreal mammals and birds In Intermountain Biogeogrur~hy A Synzposium (eds KT Harper and J L Rcvcal) Great Basin Natural Memoir 2 pp 209-28

Brown J H 1986 Two decades of interaction between the MacArthiu-Wilson model and tile complexities of mammalian distributions Biological Journal d t h c Lirlnearl Society 28 23151

Cookc H B S 1976 Suidac from Plio-Pleistocene strata of the Rudolf Basin In Earlicst Mair a ~ r d Environmcrlts in the Lakc Rlrdolf Basin Stratigraphy Paleoecology and Evolutioir (cds Y Coppcns F C Flowcll G LI Isaac and RE F Lcakey) Chicago Univcrsity of Cllicago Press pp 25 1-63

Culotta E 1995 Many suspects to blame in Madagascar cxtinctions Science 268 1568-9

Davis J and Lidicker W Z Jr 1975 Tlle taxonomic status of tllc soutl~ern sea otter P r o c ~ ~ d i i z g oJtlze Cr~lijorrliri Acriderny o f Sciences 40(14) 429-37

Diamond J M 1984 Historic extinctions a Rosetta Stone for understanding prcllistoric extinctions in Quuterrlury Extir1ctions A Prehistoric Revolution (eds P S Martin and R 6Klcin) Tucson University of Arizona Prcss pp 824-62

Emslic S D 1987 Age and diet of fossil California condors in Grand Canyon Arizona Science 237 768-70

Fisller J W Jr 1995 Bone surface modifications in 7ooarchaeology Journal ofArchacologica1 Method rind Theory 2 7-68

Gifford-Gonzalez D 1991 Bones arc not enough analogues knowledge and intcrprctivc strategies in zooarchaeology Journal ofAntlzropologica1 Archrreologj 10 215-54

Gilmorc R M 1949 Tllc identification and value of mammal bones from archeological excavations Journal of Mummulogy 30 163-9

Graham RW 1988 The role of climatic change in the design of biological reserves tho palcoccological perspccti~rc for conser~ration biology Corltervution Biologjl 2 391-4

Graham RW and Graham M A 1994 Late Quaternary distribution of Marter in North America In Mcrrten~ Sublcs urtd Fislrcrs Biology and Conservation (eds S W Ruskirk 14 S Harestad M 6Raphael and RA Powell) Itllaca NY Corncll University Prcss pp 2 6 5 8

Grayson D M 1991 Tllc biogeographic history of small mammals in the Great Basin observations on thc last 20000 years Journal of Mammalogy 68 359-75

Green J S and Flinders J T 1980a Brachylrig~s idahoerisis Manznzalian Specics 125 1--4

Green J S and Flindcrs J T 1980b Habitat and dietary relationships of the pygmy rabbit dournnl Kringe Marlrigernent 33 13amp42

I3css K J r 1993 Rocky Times in Rocky Mourltairl hcrtiorzril Park Niwot Colo University Prcss of Colorado

Flestcr F E 1991 The U S National Park Scrvicc experience with exotic species Nriturril Areri ~ Jourr~ril1 1 127-8

Hoerr W 1993 The concept of naturalness in environmental tliscoursc Natural Areas Journal 1329-32

Ilouston D B and Schrciner E 6 1995 Alien specie in national parks drawing lines in space and time Conservation Biology 9 204-9

IHouston D B Schrciner E 6and Moorhcad R R 1994 Mourltairl Goats in Olympic Nutionul Park Biologjl and Marlugernerlt of an Introduced Species U S National Park Scrvicc Scientific Monograph NPSINROLYMINRSM-9325

I-Iudson J (cd) 1993 From Bones to Bc~lzavior Etlznotrrchaeologicul and E~plcrimcntill Coiltri- butiorls ro rhe Intcrprc~tation of Fblrr~al liernairls Carbondalc Southern Illinois University Center for Archaeological Investigations Occasional Papcr No 21

T h e relevance of faunal analysis t o wildlife management 123

Hyland D C Tersak J M Adovasio J M and Sicgal M I 1990 Identification of thc speciesof origin of residual blood on lithic material Americntl Antiquity 55 104-12

Jameson R J Kenyon K W Johnson A M and Wight H M 1982 History and status of translocated sea otter populations in North America Wildlife Society Bulletin 10 100-7

Jameson R J Kenyon K W Jeffries S and VanBlaricom G R 1986 Status of a I ranslocatcd sea otter population and its habitat in Washington The Murrelet 67 84-7

Kay C E 1994 Aboriginal overkill the role of Native Americans in structur~ng western ecosystems Hurnnt~ Nrrrure 5 359-98

Kenyon K W 1969 The sea otter In the eastern Pacific Ocean Worth Amer~crrtz Firuna 68

Korn H 1994 Genetic demographic spatial environmental and catastrophic effects on the survival probability of small populations of mammals In Mitlimum Animrrl Popu1rrrior1s (ed H Remmert) Berlin Springer-Verlag pp 3)-49

Lcggc A J 1978 Archacozoology - or zooarchacology In Research Prohlems in Zoourchueology (eds D R Brothwell K D Thomas and J Clutton-Brock) Univcrsity of London Institute of Archaeology Occasional Publications No 3 pp 129-32

Leopold A S Cain S A Cottam C IM Gabrielson I N and Kimhall T L 1963 Wildlife management in the national parks Trunsuctions of the Twenty-Eighth North American Wildlife und Nutural Resoirrce~ Cd70nferencepp 2945

Loy T H 1993 The artifact as site an example of the biomolccular analysis of organic residues on prehistoric tools World Archaeology 25 44-63

Lyman R L 1988a Significance for wildlife management of the late Quaternary biogeography of mountain goats (Oreurnnos urnericunus) in the Pacific Northwest USA Arctic and Alpine Reseurch 20 13-23

Lyman R L 1988b Zoogeography of Oregon coast marine mammals the last 3000 years Murine Murnrnul Science 4 247-64

Lyman R L 1991 Late Quaternary biogeography of the pygmy rabbit (Brachylagirs iduhoensis) in eastern Washington Jo~lrnalof Mammalogy 72 11amp17

Lyman R L 1994a The Olympic Mountain goat controversy a different perspective Conservation Biology 8 898-901

Lyman R L 1994b Vertebrute Taphonomy Cambridge Cambridge University Press

Lyman R L 1995a Determining when rare (zoo)archacological phenomena are truly absent Journal of Archueologicul Method unrl Theory 2 369-424

Lyman R L 199 Inaccurate data and the Olympic National Park mountain goat controversy North wect Science 69 234-8

McAllister K and Allen H 1993 Stutuc of the Pygmy Rabbit (Brachylagus idahoensis) in Wushington Olympia Washington Department of Wildlife Wildlife Management Division

McAllister K and Allen H 1994 Wushington Stute Draft Recovery Plun for the Pygmy Rabbit (Brachylagus idahocnsis) Olympia Washington Department of Wildlife Wildlife Management Division

MacArthur R H and Wilson E 0 1967 The Theory of Islunci Biogeography Princeton NJ Princeton University Press

MacAskie I 1987 Updated status of the sea otter Enhydru lutris in Canada Cunudiun Field- Nuturcrlist 101 279-83

McDonald J N 1981 North American Bison Their Clascicution and Evolution Berkeley University of California Press

Mcltzcr D J 1995 Clocking the first Americans Annual Review of Anthropology 24 71-45

124 R Lee E y n ~ u n

OBricn S J and Mays E 1991 Burcauc1atic mischief rccogni~ing enciangcrcd spccics and subspecies Scicncc 251 1 187--8

Ogden A 1933 Russian sea ottcr anal seal hunting on the Califorilia coast Caij(~rrrirrNisroricci1 Qmcrrrerly 12 29-5 1

Olscn S I and Olscn I W 1981 A conin-icnt on nomcnclaturc in faunal studies iui~cric~un Antiquity 46 192-4

Peck J M Miclucllc 14 C and Wright M 61987 Arc bison cuotic in the Wrangcll-St Elia National Park and Pre~cr~c Envivnnnlcrzrcrl iWrlrlcrgcineilr I I l(195 3

Quinn J Iand ifarrison S P 1988 Effects of habitat fragmentation anhi isolation on spci-is richness evidence from biogeographic patterns Oecologiii 75 132-40

Rasmusson E M 1985 El Nino and asiations in climate Atzcricr~z Scic3nritt 73 168-77

Raup D M 1984 Death of spccics I n Errirrcrions (etl M 14 Nitccllti) dlhicago University of Chicago Press pp 1-19

IcnsbcrgcrJ M and Barnosky A I) 1993 Short-term fluctuations in slniill mammals ot the lato Pleistocene from eastern Washington Wn iWorphologicrrl Chr~rigc irr C)iir~t~rricrr~ Mrrinnzcrls (J ~ i i~ t l~ Alzcrica (eds R A Martin and 4 Igt 131rnosky) Qambridgc Ciimbsidgc University Prcss pp 299-342

Richards M Smallcy KSykcs R and Iledges BP 1993 iir-ch21eology and genetics anaiysirig DNA from skeletal remains Worlrl~lrchtrcolog~5 18-28

Richards M B Sykcs B C and Hedges MF M 1995 Authenticating D N A extriicted Irorri ancient skeletal remains Iourr7crl of Arcl~aeologicril Scicncc 22 2W-9

Faohison ND 1978 Zooarchaeology its hiirto~y and dcvclopnirn~7r7rrcssrc Anthropologic~ul A~orirition Miscellancotrs Pripcr 2 1-2

Rocst A 1 1973 Subspecies of the sea ottcs Er7llytira Iiitris Io5 ii7gcles Coirnty Nnturcrl tiisrory Mltscirrn (ontributior~s in Scicncr 252

Rocst A 1 1979 A re-evaluation oi sea o t t c ~ taxonomy Paper prcscntcci at the Sea Ottcr Workshop Santa Barbara Cilifornia

Scheffer V B 1993 Tlie Olympic Mo~rntain goat conrrovcrs) a pcr5pectivc Coriscrvtrtior Biology 7 916-19

Sclicffcr V B and Wilkc F 1950 Validity of the subspccics E~llzydrii 1iltris r7crcic the southcrn s ix ottcr Joltvnill qf the Wcr~I7ir7gton Accrtlcuny 01 S~icr~cr402672

Simons I) R 1083 Interactions lsctween Calilornin corrdors and humans in prehistoric far wcstern North America In Vcrltcrrr Biology und ~tlunuget~ierlt (cds S KWilbur and J A Jackson) Berkeley University or Ciliforni~i Press pp 470-91

Steadman D xi 1995 Prehistoric cstinctions of Pacific isiarld birds biodiversity nlccts zooarchacology Scieilce 267 1 123-3 1

Trillrnich F and Limbcrger D 1985 Drastic effects of 871 Nirio on Ga1apigos piniiipcd Oecologiu 67 19-22

VanBlaricorn 6Rand Estcs 9 A (cds) 1988 l h p Bot~irnlrniij bcologj oj Seu O t r e r ~ (Berlin

Springer-Verliig

Wagner F 11 Forest RC i i l l R R Mcdullough 14 R Pcltoil M MPorTcr W F and Salwasscr 13 1995 WildlifePolicies in the U S Nirrionul Purks Washington DC Island Prcsi

Weisler M I and Gargett R H 1993 Pacific island avian cstinctions the taphonorny or human predation Arclzucoloyj in Occuniu 28 85-03

Wendell F E flardy R A An-rc5 1 land Burgi K T 1986 Tcrnporalanci spatial patterns in

T h e relevance of faunal analysis t o wildlife management 123

Hyland D C Tersak J M Adovasio J M and Sicgal M I 1990 Identification of thc speciesof origin of residual blood on lithic material Americntl Antiquity 55 104-12

Jameson R J Kenyon K W Johnson A M and Wight H M 1982 History and status of translocated sea otter populations in North America Wildlife Society Bulletin 10 100-7

Jameson R J Kenyon K W Jeffries S and VanBlaricom G R 1986 Status of a I ranslocatcd sea otter population and its habitat in Washington The Murrelet 67 84-7

Kay C E 1994 Aboriginal overkill the role of Native Americans in structur~ng western ecosystems Hurnnt~ Nrrrure 5 359-98

Kenyon K W 1969 The sea otter In the eastern Pacific Ocean Worth Amer~crrtz Firuna 68

Korn H 1994 Genetic demographic spatial environmental and catastrophic effects on the survival probability of small populations of mammals In Mitlimum Animrrl Popu1rrrior1s (ed H Remmert) Berlin Springer-Verlag pp 3)-49

Lcggc A J 1978 Archacozoology - or zooarchacology In Research Prohlems in Zoourchueology (eds D R Brothwell K D Thomas and J Clutton-Brock) Univcrsity of London Institute of Archaeology Occasional Publications No 3 pp 129-32

Leopold A S Cain S A Cottam C IM Gabrielson I N and Kimhall T L 1963 Wildlife management in the national parks Trunsuctions of the Twenty-Eighth North American Wildlife und Nutural Resoirrce~ Cd70nferencepp 2945

Loy T H 1993 The artifact as site an example of the biomolccular analysis of organic residues on prehistoric tools World Archaeology 25 44-63

Lyman R L 1988a Significance for wildlife management of the late Quaternary biogeography of mountain goats (Oreurnnos urnericunus) in the Pacific Northwest USA Arctic and Alpine Reseurch 20 13-23

Lyman R L 1988b Zoogeography of Oregon coast marine mammals the last 3000 years Murine Murnrnul Science 4 247-64

Lyman R L 1991 Late Quaternary biogeography of the pygmy rabbit (Brachylagirs iduhoensis) in eastern Washington Jo~lrnalof Mammalogy 72 11amp17

Lyman R L 1994a The Olympic Mountain goat controversy a different perspective Conservation Biology 8 898-901

Lyman R L 1994b Vertebrute Taphonomy Cambridge Cambridge University Press

Lyman R L 1995a Determining when rare (zoo)archacological phenomena are truly absent Journal of Archueologicul Method unrl Theory 2 369-424

Lyman R L 199 Inaccurate data and the Olympic National Park mountain goat controversy North wect Science 69 234-8

McAllister K and Allen H 1993 Stutuc of the Pygmy Rabbit (Brachylagus idahoensis) in Wushington Olympia Washington Department of Wildlife Wildlife Management Division

McAllister K and Allen H 1994 Wushington Stute Draft Recovery Plun for the Pygmy Rabbit (Brachylagus idahocnsis) Olympia Washington Department of Wildlife Wildlife Management Division

MacArthur R H and Wilson E 0 1967 The Theory of Islunci Biogeography Princeton NJ Princeton University Press

MacAskie I 1987 Updated status of the sea otter Enhydru lutris in Canada Cunudiun Field- Nuturcrlist 101 279-83

McDonald J N 1981 North American Bison Their Clascicution and Evolution Berkeley University of California Press

Mcltzcr D J 1995 Clocking the first Americans Annual Review of Anthropology 24 71-45

124 R Lee E y n ~ u n

OBricn S J and Mays E 1991 Burcauc1atic mischief rccogni~ing enciangcrcd spccics and subspecies Scicncc 251 1 187--8

Ogden A 1933 Russian sea ottcr anal seal hunting on the Califorilia coast Caij(~rrrirrNisroricci1 Qmcrrrerly 12 29-5 1

Olscn S I and Olscn I W 1981 A conin-icnt on nomcnclaturc in faunal studies iui~cric~un Antiquity 46 192-4

Peck J M Miclucllc 14 C and Wright M 61987 Arc bison cuotic in the Wrangcll-St Elia National Park and Pre~cr~c Envivnnnlcrzrcrl iWrlrlcrgcineilr I I l(195 3

Quinn J Iand ifarrison S P 1988 Effects of habitat fragmentation anhi isolation on spci-is richness evidence from biogeographic patterns Oecologiii 75 132-40

Rasmusson E M 1985 El Nino and asiations in climate Atzcricr~z Scic3nritt 73 168-77

Raup D M 1984 Death of spccics I n Errirrcrions (etl M 14 Nitccllti) dlhicago University of Chicago Press pp 1-19

IcnsbcrgcrJ M and Barnosky A I) 1993 Short-term fluctuations in slniill mammals ot the lato Pleistocene from eastern Washington Wn iWorphologicrrl Chr~rigc irr C)iir~t~rricrr~ Mrrinnzcrls (J ~ i i~ t l~ Alzcrica (eds R A Martin and 4 Igt 131rnosky) Qambridgc Ciimbsidgc University Prcss pp 299-342

Richards M Smallcy KSykcs R and Iledges BP 1993 iir-ch21eology and genetics anaiysirig DNA from skeletal remains Worlrl~lrchtrcolog~5 18-28

Richards M B Sykcs B C and Hedges MF M 1995 Authenticating D N A extriicted Irorri ancient skeletal remains Iourr7crl of Arcl~aeologicril Scicncc 22 2W-9

Faohison ND 1978 Zooarchaeology its hiirto~y and dcvclopnirn~7r7rrcssrc Anthropologic~ul A~orirition Miscellancotrs Pripcr 2 1-2

Rocst A 1 1973 Subspecies of the sea ottcs Er7llytira Iiitris Io5 ii7gcles Coirnty Nnturcrl tiisrory Mltscirrn (ontributior~s in Scicncr 252

Rocst A 1 1979 A re-evaluation oi sea o t t c ~ taxonomy Paper prcscntcci at the Sea Ottcr Workshop Santa Barbara Cilifornia

Scheffer V B 1993 Tlie Olympic Mo~rntain goat conrrovcrs) a pcr5pectivc Coriscrvtrtior Biology 7 916-19

Sclicffcr V B and Wilkc F 1950 Validity of the subspccics E~llzydrii 1iltris r7crcic the southcrn s ix ottcr Joltvnill qf the Wcr~I7ir7gton Accrtlcuny 01 S~icr~cr402672

Simons I) R 1083 Interactions lsctween Calilornin corrdors and humans in prehistoric far wcstern North America In Vcrltcrrr Biology und ~tlunuget~ierlt (cds S KWilbur and J A Jackson) Berkeley University or Ciliforni~i Press pp 470-91

Steadman D xi 1995 Prehistoric cstinctions of Pacific isiarld birds biodiversity nlccts zooarchacology Scieilce 267 1 123-3 1

Trillrnich F and Limbcrger D 1985 Drastic effects of 871 Nirio on Ga1apigos piniiipcd Oecologiu 67 19-22

VanBlaricorn 6Rand Estcs 9 A (cds) 1988 l h p Bot~irnlrniij bcologj oj Seu O t r e r ~ (Berlin

Springer-Verliig

Wagner F 11 Forest RC i i l l R R Mcdullough 14 R Pcltoil M MPorTcr W F and Salwasscr 13 1995 WildlifePolicies in the U S Nirrionul Purks Washington DC Island Prcsi

Weisler M I and Gargett R H 1993 Pacific island avian cstinctions the taphonorny or human predation Arclzucoloyj in Occuniu 28 85-03

Wendell F E flardy R A An-rc5 1 land Burgi K T 1986 Tcrnporalanci spatial patterns in

124 R Lee E y n ~ u n

OBricn S J and Mays E 1991 Burcauc1atic mischief rccogni~ing enciangcrcd spccics and subspecies Scicncc 251 1 187--8

Ogden A 1933 Russian sea ottcr anal seal hunting on the Califorilia coast Caij(~rrrirrNisroricci1 Qmcrrrerly 12 29-5 1

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