Paleozoology in the Service of Conservation Biology

Evolutionary Anthropology 15:11–19 (2006)

ARTICLES

R. LEE LYMAN

Paleozoological data reveal past conditions created by anthropogenic and nat- grapple with the fact that ecosystems ural processes. These conditions can serve as benchmarks of ecological proper- and landscapes are not static for natu- ties and processes desired by conservation biologists. Paleozoological data pro- ral and anthropogenic reasons.16 Their vide empirical evidence analogous to experimental results of anthropogenic and desire to manage a minimally anthro- environmental causes. They can be used to determine whether a taxon is native or pogenically influenced ecosystem intro- exotic to an area, distinguish invasive from recolonizing taxa, choose a manage- duces the difficulty of identifying the ment action likely to produce a desired result, test benchmarks based on historic boundary between natural and unnatu- data, reveal unanticipated effects of conservation efforts, and identify causes of ral.17–19 But nonanthropogenically in- ecological conditions. It is time to use paleoecological knowledge in the service of fluenced ecosystems are not always modern conservation biology. desired. For example, some anthropo- genically introduced exotic taxa such as game birds in the western United States Conservation biologists, restoration compositions of biological communi- are economically beneficial and ecolog- ecologists, and wildlife managers of- ties occupying tens to hundreds of ically benign. ten select an ecological benchmark,1,2 hectares, as well as to ecosystems con- The paleozoological record pro- ecological baseline,3 or historical sisting of organisms, geology, fire re- vides unprecedented data that reflect landscape4,5 that they seek to recreate gimes, and so on, as well as ecological the long-term operation of many eco- or maintain in an area. A benchmark and evolutionary processes.6–8 Typi- logical and anthropogenic processes is a goal toward which conservation cally, a benchmark is established by and may provide guidance to distin- activities are aimed; it is an ecological reference to the early historic period guishing effects of the two.20–23 My condition or process that is desired. because written records are available specific goal here is to show that pa- Benchmarks vary in scale from a par- and also because anthropogenic, par- leozoological data are a significant ticular gene pool or range of pheno- ticularly industrial-era influences, are source of information on bench- types to the presence or absence of a usually undesirable. Conservation bi- marks. I focus on mammals, but any species in an area of a few hectares to ologists realize that any chosen taxon of plant or animal can provide benchmark is a moving target given data concerning a benchmark. My the vagaries of both particularistic general goal is to encourage paleo- contingencies and evolutionary histo- ecologists to consider how their re- ries.9 They worry about long-term cli- search might be of value to conserva- R. Lee Lyman became interested in the matic change and anthropogenic vari- tionists and to publish their research utility of paleozoological data to conservation biology in the middle 1980s, when ables and their influence on plant and in journals such as Biological Conser- he was studying the morphometry of the animal taxa and ecosystems.10,11 vation, Conservation Biology, Bio- prehistoric sea otter (Enhydra lutris)inthe Conservation biologists find multidis- Science, Ecological Restoration, and eastern Pacific Ocean. That interest ex- panded with his examination, in the late ciplinary research necessary to contend Environmental Management to inform 1980s, of the controversy regarding the with ecological, biological, and land- conservation biologists of the value of exotic or native status of mountain goats 12–14 (Oreamnos americanus) in Olympic Na- scape degradation. The Long Term paleoecological data. tional Park. Together with Ken Cannon, Ecological Research (LTER) Network Paleoecologists publish in these Lyman recently edited the volume Zooar- established by the United States Na- venues, but they seldom identify the chaeology and Conservation Biology. tional Science Foundation monitors, exact management implications of over long periods, how and why ecosys- their observations.24–30 Perhaps this is Key words: ecosystems, conservation biology, tems and ecological variables and pro- because they believe it would be “dan- paleoecology, paleozoology cesses interact and operate.15 LTER gerous” to offer suggestions outside recognizes that research must exceed a their field of expertise.31 I believe, © 2006 Wiley-Liss, Inc. season or two, a year or two, or a even however, that we must make explicit DOI 10.1002/evan.20083 Published online in Wiley InterScience decade or two if we are to understand suggestions because conservationists (www.interscience.wiley.com). ecosystems. Conservation biologists do not always perceive the value of 12 Lyman ARTICLES

an ecological benchmark is a moving target.

EXOTIC TAXA AND NATIVE TAXA One conservation activity is to translocate animals from one population to another to regulate the size of the donor population to enhance or reestablish another population, or both. Translocation requires that we know the benchmark of local indigenous taxa.42 Paleozoological data as- Figure 1. A model of how the environmental tolerances of a species influence its abun- sist with the determination of whether dance on the landscape. Any environmental variable such as number of frost-free days, annual precipitation, or temperature range can be plotted on the x axis. a taxon is native or exotic to a loca- tion. Definitions of exotic taxa vary.43 For purposes of this paper, exotic, alien, nonindigenous taxa are those paleoecological data. As more papers such terms as natural, native, pristine, that did not previously exist in an with such suggestions are published, and the like be explicitly defined. In area.44 Efforts in many national parks members of the conservation profes- the United States, the terms usually are devoted to eradicating established sions will realize that the long-term are defined as “pre-Columbian or im- exotic taxa and ensuring that no new 37,38 data provided by paleoecology is wor- mediately post-Columbian.” It is exotic taxa become established.45–47 thy of serious consideration. Then re- often implied that “Columbian era” Plans to reintroduce the North search as well as job opportunities for signifies nonanthropogenic, which ig- American wapiti (Cervus elaphus)to paleoecologists may increase and a nores the 12,000-plus years that the state of Missouri have been dis- new vault of funding may open. American Indians have been in North cussed for two decades. The historic Conservation biologists often mud- America and presumes that Native record indicates that this large ungu- dle along with imperfect knowledge, Americans had minimal influence on late was present in the state in the not knowing the exact consequences pre-Columbian ecosystems. Paleoeco- nineteenth century, but does not indi- 32 of their activities. To not act because logical data indicate that people cate all locations where wapiti were of imperfect knowledge represents the throughout the world had all sorts of found and not found. The paleozoo- 33 “paralysis of analysis.” Not only do ecological influences.39 Terms that logical record indicates that wapiti paleoecological data represent knowl- imply nonanthropogenically influ- were only present in the state when edge, they underscore the moving-tar- enced ecosystems and landscapes climates were cooler than at present get nature of benchmarks, and high- should be avoided.34 and that they occupied the topograph- light the fact that ecological stasis is The second point is that every taxon ically rugged and timbered Ozark Pla- unattainable. They also may suggest has been and continues to be shaped teau of the south-central portion of which of several benchmarks is the by natural selection to live within a the state.48 The state Fish and Wildlife most feasible to attain or maintain. A certain range of temperature and pre- Department plans to release wapiti in chosen benchmark will depend on so- cipitation, vegetation, geology and to- areas with open forest and prairie cial, political, economic, and ecologi- pography, predation, and other envi- habitats and little topographic relief. 34 cal variables. To convince anyone of ronmental variables (Fig. 1). If one of The transplant effort has been de- the “applied” value of paleoecological those variables changes, the taxon layed, but if the state plan is followed data, examples must be identified. has three options: to become locally will the wapiti survive where there is Here I describe several examples in extirpated, migrate to an area where no evidence that they are native? which paleozoological data have been the environmental variable has not Banff National Park straddles the brought to bear on particular conser- changed, or adapt to the new environ- crest of Canada’s Rocky Mountains. vation, management, and restoration ment.40 The third alternative can take Biologists are contemplating releas- issues. Before doing that, however, I at least two forms that are not neces- ing bison (Bison bison) into this rug- must address two background issues. sarily exclusive. The organism can ged wilderness area. Zooarcheological adapt by decreasing in abundance, al- data indicate that bison were present tering its morphometry, or a combi- there during the last 10,000 years and BACKGROUND nation of the two.41 A species may dis- that individuals were adult males; no Many federal agencies such as the appear from an area and reappear at a cow-calf herds are represented.49 United States National Park Service later date; its abundance may increase There is no evidence of differential are charged with managing “natural” or decrease only to decrease or in- preservation of male remains, so or “pristine” landscapes and bio- crease at a later date; or individuals should bison in the park be managed tas.35,36 This charge demands that may shrink or grow larger. In short, as a sink population into which excess ARTICLES Paleozoology and Conservation Biology 13 male bison from surrounding areas were it to be observed today off the between Oregon State and Washing- immigrate or should park bison be coast of Washington. Prehistoric ton State up to Celilo Falls, 324 km managed as a source population with northern fur seal (Callorhinus ursi- upstream of the river’s mouth in the a high reproduction rate and out-mi- nus) remains from several areas along Pacific Ocean.63 Those seals made gration?50 The archetypical bison is the Washington, Oregon, and north- that journey throughout the Holo- the Plains bison (B. b. bison); the ern California coasts57–59 indicate that cene, never getting farther inland due wood bison (B. b. athabascae), now late twentieth-century records of this to the impassability of the falls. Con- extinct, occurred in the northern taxon along the California coast60 rep- struction of Bonneville Dam at river plains.51,52 Remains of both have been resent recolonization rather than in- km 250 in 1938 ended the access of reported in Banff National Park,49 so vasion. Stable isotopes in prehistoric pinnipeds to upstream areas. Zooar- the release of Plains bison there seems remains of this taxon recovered from cheological remains indicate that har- acceptable. sites in central California also indicate bor seals were pursuing salmon (On- Whether a taxon is native or exotic recolonization.61 corhynchus spp.) that were making is but one side of the coin, the side Grayson and Delpech26 argue that their annual spawning run upstream. concerning determination of which long-term climatic change repre- Were Bonneville Dam to be removed, 64 taxa to allow in an area. Some exotic sented by the shift from a glacial pe- a remote possibility, it is likely that taxa are beneficial or ecologically be- riod to an interglacial period resulted harbor seals would once again be ob- nign. The other side of the coin con- in the loss of reindeer (Rangifer taran- served at Celilo Falls. Commercial cerns determination of which taxa salmon fishermen are likely to per- should be denied access to an area. ceive the seals as invasive. This involves “invasive” taxa, the ex- Whether a taxon is Paleozoological data may reveal otic taxa that cause ecological or eco- prehistoric patterns that contradict nomic damage.44 native or exotic is but historical data or that suggest anthro- one side of the coin, the pogenic causes of historically documented patterns. Paleozoological data DISTINGUISHING INVASIVE side concerning indicate that ibex (Capra ibex) occu- TAXA FROM RECOLONIZING determination of which pied more topographic positions TAXA across the Italian landscape than they taxa to allow in an area. do today. The mountain habitat at the Invasion biology is presently a ma- Some exotic taxa are 2,000 to 3,000 m elevation used by jor concern.53,54 An apparent invasion ibex today may be the result of an- may, however, reflect recolonization beneficial or thropogenic transplanting efforts.31 of pre-Columbian ranges rather than ecologically benign. The Stable isotope analysis of prehistoric colonization of new areas.43 Paleozoo- other side of the coin bighorn sheep (Ovis canadensis) re- logical data may distinguish the two. mains in Wyoming State indicates Darwent and Darwent55 found that concerns determination that this taxon’s prehistoric seasonal paleozoological data indicate that the of which taxa should be migration has been altered. Bighorn range and abundance of muskox sheep formerly wintered in lowlands (Ovisbos moschatus) in the high Arc- denied access to an now occupied by humans. Today big- tic of eastern Canada and Greenland area. horn are (re)utilizing what was for- fluctuated with climatic change over merly their seasonal range, prompting the last 5,000 years. Greenland mus- questions about suggested manage- kox are not in jeopardy of extirpation, ment actions.65 contrary to the belief of local conservation biologists, and one translo- dus) from southwestern Europe. Their cated group was released where it is paleozoological data indicate that rein- CHOOSING MANAGEMENT unlikely to survive. Muskox may re- deer will recolonize southwestern Eu- OPTIONS colonize some areas if climate rope if climate shifts to a period of changes appropriately and humans do Knowledge of the environmental tol- cooler summers and glaciation. Other not interfere. erances of endangered taxa, along with paleozoological data indicate that nu- Etnier56 identified the remains of knowledge of their prehistoric biogeo- juvenile male and female Guadalupe merous taxa will recolonize or invade graphic histories, may assist in identi- fur seal (Arctocephalus townsendi)in the presently hyperarid desert of the fying why those taxa are in jeopardy 500-year-old deposits on the northern Eastern Sahara should precipitation in- today and how we might or might not coast of Washington State. The far- crease there.62 Whether reindeer or protect them. The pygmy cottontail rab- thest north this species had been his- Saharan taxa are considered invaders bit (Brachylagus idahoensis) is found to- torically reported was just north of will likely depend on human land use day in the physiographic Great Basin of San Francisco Bay. The remains indi- at the time. western North America (eastern Cali- cate that this taxon would not be in- Zooarcheological remains of harbor fornia, Nevada, western Utah, southern vading new territory but instead recol- seals (Phoca vitulina) show that this Idaho, and southeastern Oregon) and in onizing previously occupied range species ascended the Columbia River a small isolated area of eastern Wash- 14 Lyman ARTICLES ington State. The modern distribution tions.72–74 Within the limits of resolu- populations are having serious im- of the pygmy cottontail implies that a tion of radiocarbon dating and pacts on what are supposed to be pro- corridor between southeastern Oregon stratigraphy, causes of varied rates of tected ecosystems.78–80 Before the his- and eastern Washington served as an extinction can be measured.75 Extinc- torical period, wapiti populations immigration route for individuals orig- tion rates and causes are critical infor- were, Kay79 believes, much smaller inating in the former population to mation for conservation biologists, as is than today, largely as a result of pre- have established the latter. Late Pleisto- information about the opposite process historic human predation. Kay bases cene remains of this taxon within this of persistence.76 In the case of pygmy his argument on two measures. One is hypothetical corridor indicate that it rabbits, the concern is with managing the modern abundance of wapiti in was the route.66,67 Were a conservation and protecting a depleted population. the Yellowstone ecosystem of north- biologist to want to replenish the Wash- In other cases, the concern may be with western Wyoming. The other com- ington population, the genetic source a population that is too large. prises a paleozoological benchmark seems clear. However, modern individ- made up of the summed pre-Colum- uals from Washington are genetically TESTING POPULATION SIZE bian zooarcheological record from distinct from nearby populations.68 Pa- seven western states (Wyoming, Mon- leozoological data indicate that the In North America, large mammals tana, Utah, Idaho, Nevada, Oregon, range and the population of pygmy rab- are today subjects of management and Washington) and spanning the bit 8,000 to 4,000 years ago were larger plans because they provide prey for last 10,000 years. Because the mea- than in those in the twentieth centu- sure representing the modern Yellow- ry.66,67,69 The earlier period was a time stone abundance of wapiti is consid- when big sagebrush (Artemisia triden- A taxon’s history will erably higher than the abundance of tata) was widespread and abundant; reveal benchmarks. wapiti indicated by the lumped paleo- pygmy rabbits depend on this plant for zoological data, Kay argues that wa- cover and food. The range and abun- Each benchmark will piti should be hunted down to pre- dance of sagebrush in eastern Washing- have unique spatio- Columbian levels to avoid further ton decreased as the climate cooled and degradation of western ecosystems. It effective moisture increased 4,000 years temporal coordinates is unlikely that his benchmark will be ago. So, too, did the pygmy rabbit and environmental adopted by conservation biologists for range, and likely its abundance, de- contexts. Thus we must two reasons. First, no management crease.67 But there was still a lot of sage- application will be uniformly applied brush after 4,000 B.P., and it was wide- avoid a “one size fits all” to the seven-state area from which his spread. Why, then, were there so few solution, which is sample derived. Second, wapiti abun- pygmy rabbits in the middle of the dances varied across both the area twentieth century and why were there anathema to biological and the last 10,000 years.81 fewer than 100 individuals remaining and ecological diversity. Paleozoological taxonomic abun- in eastern Washington by the year dances are, at best, ordinal scale and 2000?70 are best considered relative to the Cattle ranching was initiated in late abundances of other taxa.82–84 Data nineteenth century.71 Open-range graz- from eastern Washington State indi- ing destroyed sagebrush. The pygmy hunters or wildlife-viewing opportu- cate that Kay’s benchmark of wapiti rabbit is a burrower and the cattle com- nities for tourists. The long-term his- population size is unfounded there. If pacted local soil, making burrowing dif- tory of how these taxa responded to humans depressed local wapiti popu- ficult.70 Sagebrush loss was exacer- varying levels of predation, environ- lations during the first thousand years bated by tillage agriculture in the early mental change, and efforts to supple- that people were present, this time pe- twentieth century. If the remaining ment their gene pools may be revealed riod is not represented in the samples population of pygmy rabbits in eastern by paleozoological data. A taxon’s his- of faunal remains that Kay used.81 Washington is to be protected, the last tory will reveal benchmarks. Each Ungulate remains in 86 samples span- native stands of big sagebrush must not benchmark will have unique spatio- ning the last 10,000 years indicate that be destroyed.70 In the case of eastern temporal coordinates and environ- there was no decrease in wapiti abun- Washington’s pygmy rabbits, migration mental contexts. Thus we must avoid dance relative to other ungulates over corridors are implausible. However, de- a “one size fits all” solution, which is this time span (Fig. 2). There is no creased destruction of stands of big anathema to biological and ecological evidence of intensified processing or sagebrush may be plausible. That such diversity. alteration in the demography of the destruction depleted the preferred hab- Political ecologist Charles Kay77 ar- kill, both of which are expected to ap- itat of this small rabbit is suggested by gued that wapiti (Cervus elaphus) pop- pear if, given an increasingly sophisti- historic data; it is confirmed with the ulations in the western United States cated technology and increasing hu- independent data of the paleozoological are too high today as a result of mod- man population, hunters took similar record. ern hunting regulations and reduced abundances of wapiti from a succes- A related issue concerns rates of ex- natural predation. This is especially sively more depressed population over tinction of small isolated popula- the case in national parks where large time. There is no evidence here that ARTICLES Paleozoology and Conservation Biology 15 human hunting influenced the wapiti metapopulation. This does not mean that humans did not influence that metapopulation, only that such an influence is undetectable in the data available. There are no early historic zooarcheological collections from eastern Washington with which to compare modern wapiti abundance; such collections would reveal early impacts, if any, of Euroamerican predation regimes and potentially would corroborate Kay’s benchmark. We can monitor temporal and spa- tial trends with paleozoological data; trends present a series of benchmarks, each of slightly different magnitude or value in one respect or another. That benchmarks are a time slice of a long- term trend of shifting values under- scores the fact that ecosystem man- Figure 2. Abundance of elk remains relative to all ungulate remains in 86 assemblages from agement decisions are choices and so eastern Washington. The horizontal line is the simple best-fit regression line; its slope are benchmarks. Importantly, paleo- (ϭ0.000001, from youngest to oldest) is insignificant. zoological data may reveal unanticipated effects of conservation efforts. last 180 years but did not affect Oregon and on the Columbia River UNANTICIPATED EFFECTS deer? I hypothesize that the 1920s ad- floodplain in the Portland Basin of Two subspecies of wapiti occurred dition of Rocky Mountain wapiti to northwestern Oregon and south- in Washington State in the nine- the gene pool in the Cascade Moun- western Washington. It is one of the teenth and twentieth centuries. The tains, through a process of hybridiza- original 78 taxa in the United States eastern quarter of the state is occu- tion with indigenous wapiti and intro- listed in 1968 as federally endan- pied by the Rocky Mountain subspe- gression, produced the smaller wapiti gered. Hunting of this subspecies is cies (Cervus elaphus nelsoni); the present today in southwestern Wash- now restricted, and many of the western quarter is occupied by the ington and northwestern Oregon. This Portland Basin population’s mem- on-average larger Roosevelt wapiti is an unexpected outcome. The hy- bers are in the Columbian White- (C. e. roosevelti). The identity of the pothesis needs to be tested by genetic Tailed Deer National Wildlife Ref- subspecies that occupied the Cas- analysis of prehistoric remains. One uge, established in 1972 to protect cade Mountain range of the central test implication is already well estab- the subspecies. The foothills around portion of the state is debated.85 Lo- lished. What might be considered the the floodplain are occupied by the cal wapiti populations were depleted “Yellowstone genetic signature” has Columbian black-tailed deer (O. by the beginning of the twentieth been tracked across various modern hemionus columbianus), a subspe- 87 century as a result of unregulated populations. That genetic signature cies conspecific with the mule deer hunting. To supplement one rem- should not be detected in late prehis- (O. h. hemionus) of the Rocky Moun- nant population, early in the twenti- toric and early historic wapiti remains tains and eastern Washington. More eth century Rocky Mountain wapiti from Washington if the hypothesis is or less sympatric with mule deer in were captured in the Yellowstone correct. the eastern part of the state is the ecosystem and transplanted to the Any evidence that allows anticipa- northwest white-tailed deer (O. v. southern Cascade Mountains of cen- tion or retrodictive recognition of the ochrourus). Increase in the abun- tral Washington and many other lo- consequences of management activi- dance of both subspecies of white- cations across the continent.86 ties is of high value. This point arises tailed deer since 1935 is attributed to Late prehistoric (A.D. 1500 to 1792) again in the context of choosing one modern management practices. Re- and early historic (A.D. 1792 to 1835) management option over another ceived wisdom holds that white- wapiti remains from near modern Port- based on a suspected cause of a con- tailed deer were abundant in Wash- land, Oregon, are larger than both mod- servation problem. ington and Oregon until the second ern Roosevelt wapiti and Rocky Moun- half of the nineteenth century, when tain wapiti (Fig. 3). Deer (Odocoileus IDENTIFYING CAUSES habitat modification and human virginianus and O. hemionus) remains predation depleted the metapopula- from the same collection are the same The Columbian white-tailed deer tion. Paleozoological data indicate size as modern deer (Fig. 4). What (Odocoileus virginianus leucurus)is that this species’ prehistoric range caused wapiti diminution during the found today only in southwestern was larger than its modern range.88 16 Lyman ARTICLES

Figure 3. Bivariate scatterplot of distal width (ADb) and lateral length (ALl) of elk astragali from archeological sites in the Portland Basin relative to astragali of two subspecies of modern elk.

Remains of both Columbian white- tion from rebounding as a conse- mortality was greater.96 A manage- tailed deer and Columbian black- quence of decreased American Indian ment implication of these observa- tailed deer have been identified in the predation. The Columbian white- tions is that when climates are harsh, Portland Basin.89 There was no statis- tailed deer population was subse- if cub survival (recruitment) is a con- tically significant change in the rela- quently depressed by increasing fire- cern, then the harvest of prime-age tive abundance of either subspecies of arm hunting by Euroamericans in male bears should increase because deer from the pre-Euroamerican con- conjunction with industrial habitat adult males kill youngsters. Here, pa- tact (A.D. 1450 to 1792) period to the modification such as logging. If this is leozoological data provide insights post-Euroamerican contact period correct, and if more Columbian white- into how a chosen condition might be (A.D. 1792 to 1835). This is surprising, tail deer are desired, then conserva- maintained. given the loss of half to two-thirds of tionists should restrict hunting and the local American Indian population initiate traditional use of anthropo- during the late eighteenth and earliest genic fire. The latter has been recom- CONCLUSION nineteenth centuries as a result of in- mended in nearby contexts,93 but Anthropologists have long known troduced European diseases.90 Forag- requires public education if it is to that humans influence ecosystems.97 ing theory suggests that release from be successful.94 Paleozoological data Historical ecologists have demon- predation pressure will result in more here reveal a cause for population de- strated that modern ecosystems are deer.91 There is zooarcheological evi- pletion and what it might take to re- historical phenomena.98 Using pa- dence in the area that there were verse that depletion.95 leozoological data to assess bench- changes in the availability of some an- In Europe, correspondence be- marks integrates these observations. imal taxa.92 tween cave bear (Ursus spelaeus) de- A benchmark’s validity must be sub- Local precontact and post-contact mography and climate suggests how ject to testing, its causes must be American Indians set fire to floodplain modern ursids might be managed. identified, and it must be determined and foothill vegetation to enhance pri- During periods of interstadial cli- if it can be created and maintained. mary productivity and to create open mates, cave bears were less sexually Paleozoological data can contribute spaces in which to hunt.93 Euroameri- segregated during hibernation, more to all of those points. can fire suppression in the late nine- bears grew to older ages, and cub Many issues in conservation biology teenth century closed the canopy and mortality was lower relative to that are increasingly important to the depressed primary productivity.94 during stadial periods. During the lat- long-term health of humanity as we This likely resulted in dietary stress ter, more males died in hibernation enter the third millennium and as the that prevented the local deer popula- loci, fewer bears grew old, and cub ever more resource-hungry human ARTICLES Paleozoology and Conservation Biology 17

8 Smith TB, Bruford MW, Wayne RK. 1993. The preservation of process: the missing element of conservation programs. Biodiversity Lett 1:164– 167. 9 Russell E. 2003. Evolutionary history: prospec- tus for a new field. Env Hist 8:204–228. 10 Glick P, Inkley D, Tufts C. 2001. Climate change and wildlife: integrating global climate policy implementation with local conservation action. Trans N Am Wildl Nat Res Conf 66:380– 391. 11 Joyce LA, Hansen A. 2001. Climate change: ecosystem restructuring, natural disturbances and land use. Trans N Am Wildl Nat Res Conf 66:327–343. 12 Dayton PK. 2003. The importance of the natural sciences to conservation. Am Nat 162:1–13. 13 Ludwig D, Mangel M, Haddad B. 2001. Ecol- ogy, conservation, and public policy. Ann Rev Ecol Syst 32:481–517. 14 Noss R. 1999. Is there a special conservation biology? Ecography 22:113–122. 15 Hobbie JE. 2003. Scientific accomplishments of the Long Term Ecological Research Program: an introduction. BioSci 53:17–20. 16 Suding KN, Gross KL, Houseman GR. 2004. Alternative states and positive feedbacks in restoration ecology. Trends Ecol Evol 19:46–53. Figure 4. Bivariate scatterplot of distal width (ADb) and lateral length (ALl) of deer astragali 17 Angermeier PL. 2000. The natural imperative for biological conservation. Cons Biol 14:373– from archeological sites in the Portland Basin relative to astragali of two species (three 381. subspecies) of modern deer. 18 Higgs ES. 1997. What is good ecological restoration? Cons Biol 11:338–348. 19 Huff DE. 1997. Defining ecosystem health in national parks. Trans N Am Wildl Nat Res Conf 99 population continues to grow. Nu- ecosystems. It is time that we speak to 62:448–453. merous conservation issues might be those most in need of the data we can 20 Deevey ES. 1969. Coaxing history to conduct addressed with paleozoological data. provide and that we not only produce, experiments. BioSci 19:40–43. For example, based on comparisons but also use those data in the service 21 Jackson JBC. 2001. What was natural in the coastal oceans? Proc Natl Acad Sci (USA) 98: between remains of prehistoric wild of conservation biology. 5411–5418. animals and remains of conspecific 22 Lyman RL. 1996. Applied zooarchaeology: the zoo-raised animals, O’Regan and relevance of faunal analysis to wildlife manage- ACKNOWLEDGMENTS ment. World Archaeol 28:110–125. Turner100 show that the latter may be 23 Stahl PW. 1996. Holocene biodiversity: an ar- phenotypically ill equipped to survive I thank Yin Lam and two anony- chaeological perspective from the Americas. Ann if released in the wild. Sanders and mous reviewers for thoughtful com- Rev Anthropol 25:105–126. Miller101 demonstrate that a histori- ments on an earlier draft. This 24 Barsh R. 2003. Shell middens yield rich cul- manuscript began as a presentation tural deposits for fine-scale modeling of prein- cally documented seasonal migration dustrial ecosystems. Ecol Restoration 21:319– route used by a herd of pronghorn solicited by Steve Wolverton of the 320. antelope (Antilocapra americana) has Department of Geography, Univer- 25 Bowen T. 2004. Archaeology, biology and sity of North Texas. The audience conservation on islands in the Gulf of California a time depth of 5,000 to 6,000 years. Env Cons 31:199–206. provided several useful ideas. Late twentieth-century construction 26 Grayson DK, Delpech F. 2005. Pleistocene re- may disrupt that behavioral pattern indeer and global warming. Cons Biol 19:557– and jeopardize the herd. Badenhorst REFERENCES 562. 102 27 Hadly EA. 2003. The interface of paleontology and Plug highlight the use of paleo- and mammalogy: past, present, and future. J zoological training to solve forensic 1 Hunter M Jr. 1996. Benchmarks for managing Mamm 83:347–353. ecosystems: are human activities natural? Cons problems in wildlife management. Is a Biol 10:695–697. 28 Peacock E, Haag WR, Warren ML Jr. 2005. Prehistoric decline in freshwater mussels coinci- hide or skull from a poached animal? 2 Lawton JH. 1997. The science and non-science dent with the advent of maize agriculture. Cons In order to deepen conservation bi- of conservation biology. Oikos 79:3–5. Biol 19:547–551. ology’s appreciation of what paleozo- 3 Young TP. 2000. Restoration ecology and con- 29 Swetnam TW, Allen CD, Betancourt JL. 1999. servation biology. Biol Cons 92:73–83. ology can offer, we must advertise our Applied historical ecology: using the past to man- 4 Jordan WR III. 1999. Nature and culture. Ecol age for the future. Ecol Applications 9:1189– 102 skills. We should publish case stud- Restoration 17:187–188. 1206. ies in journals read by conservation 5 Povilitis T. 2002. What is a natural area? Nat 30 van der Leeuw S, Redman CL. 2002. Placing biologists, restoration ecologists, and Areas J 22:70–74. archaeology at the center of socio-natural stud- 6 Crandall KA, Bininda-Emonds ORP, Mace GM, ies. Am Antiquity 67:597–605. the like. I have focused here on mam- Wayne RK. 2000. Considering evolutionary pro- 31 Phoca-Cosmetatou N. 2004. A zooarchaeo- mals, but any sort of paleobiological cesses in conservation biology. Trends Ecol Evol logical reassessment of the habitat and ecology of or paleoecological data can be of 15:290–295. the ibex. In: Lauwerier RCGM, Plug I, editors. 7 Landres PB. 1992. Temporal scale perspectives The future from the past: archaeozoology in value to those charged with recreating in managing biological diversity. Trans N Am wildlife conservation and heritage management. and maintaining particular kinds of Wildl Nat Res Conf 57:292–307. Oxford: Oxbow Books. p 64–78. 18 Lyman ARTICLES

32 Bailey JA. 1982. Implications of “muddling ary impact of invasive species. Proc Natl Acad Sci 73 Pimm SL, Jones HL, Diamond J. 1988. On the through” for wildlife management. Wildl Soc (USA) 98:5446–5451. risk of extinction. Am Nat 132:757–785. Bull 10:363–369. 54 Vermeij GJ. 1996. An agenda for invasion bi- 74 Steadman DW. 1995. Prehistoric extinctions 33 Hutchins M. 1995. Olympic mountain goat ology. Biol Cons 78:3–9. of Pacific Island birds: biodiversity meets zooar- controversy continues. Cons Biol 9:1324–1326. 55 Darwent CM, Darwent J. 2004. Where the chaeology. Science 267:1123–1131. 34 Lyman RL, Cannon KP. 2004. Applied zooar- muskox roamed: biogeography of tundra muskox 75 Steadman DW, Pregill GK, Burley DV. 2002. chaeology, because it matters. In: Lyman RL, (Ovisbos moschatus) in the eastern Arctic. In: Rapid prehistoric extinction of iguanas and birds Cannon KP, editors. Zooarchaeology and conser- Lyman RL, Cannon KP, editors. Zooarchaeology in Polynesia. Proc Natl Acad Sci (USA) 99:3673– vation biology. Salt Lake City: University of Utah and conservation biology. Salt Lake City: Univer- 3677. Press. p 1–24. sity of Utah Press. p 61–87. 76 Mangel M, Tier C. 1994. Four facts every con- 35 Bonnicksen TM, Stone EC. 1985. Restoring 56 Etnier MA. 2002. Occurrences of Guadalupe servation biologist should know about persis- naturalness to national parks. Env Management fur seals (Arctocephalus townsendi) on the Wash- tence. Ecol 75:607–614. 9:479–486. ington coast over the past 500 years. Marine 77 Kay CE. 1994. Aboriginal overkill: the role of 36 Bowden MJ. 1992. The invention of American Mamm Sci 18:551–557. Native Americans in structuring western ecosys- tradition. J Hist Geogr 18:3–26. 57 Crockford SJ, Frederick SG, Wigen RJ. 2002. tems. Hum Nat 5:359–398. 37 Graber DM. 2003. Ecological restoration in The Cape Flattery fur seal: an extinct species of 78 Hess K Jr. 1993. Rocky times in Rocky Moun- wilderness: natural versus wild in National Park Callorhinus in the eastern north Pacific? Can J tain National Park. Niwot: University Press of Service wilderness. George Wright Forum 20:34– Archaeol 26:152–174. Colorado. 41. 58 Lyman RL. 1988. Zoogeography of Oregon 79 Kay CE. 1997. Ungulate herbivory, willows, 38 Houston DB, Schreiner EG. 1995. Alien spe- coast marine mammals: the last 3000 years. Ma- and political ecology in Yellowstone. J Range cies in national parks: drawing lines in space and rine Mamm Sci 4:247–264. Management 50:139–145. time. Cons Biol 9:204–209. 59 Hildebrandt WR. 1984. Archaeological pres- 80 Porter WF, Underwood HB. 1999. Of ele- 39 Redman CL. 1999. Human impact on ancient ence of the northern fur seal (Callorhinus ursi- phants and blind men: deer management in the environments. Tucson: University of Arizona nus) along the coast of northern California. Mur- U.S. National Parks. Ecol Applications 9:3–9. Press. relet 65:28–29. 81 Lyman RL. 2004. Aboriginal overkill in the 40 Gauthreaux SA Jr. 1980. The influences of 60 Peterson RS, Le Boeuf BJ, Delong RL. 1968. long-term and short-term climatic changes on intermountain west of North America: zooar- Fur seals from the Bering Sea breeding in Cali- chaeological tests and implications. Hum Nat 15: the dispersal and migration of organisms. In: fornia. Nature 219:899–901. Gauthreaux SA, editor. Animal migration, orien- 169–208. 61 Burton RK, Snodgrass JJ, Gifford-Gonzalez tation, and navigation. New York: Academic 82 Gilinsky NL, Bennington JB. 1994. Estimating Press. p 103–174. D, Guilderson T, Brown T, Koch PL. 2001. Holo- numbers of whole individuals from collections of cene changes in the ecology of northern fur seals: 41 Lyman RL, O’Brien MJ. 2005. Within-taxon body parts: a taphonomic limitation of the paleon- insights from stable isotopes and archaeofauna. tological record. Paleobiology 20:245–258. morphological diversity as a paleoenvironmental Oecologia 128:107–115. indicator: Late-Quaternary Neotoma in the Bon- 83 Grayson DK. 1984. Quantitative zooarchaeol- neville Basin, northwestern Utah. Quaternary 62 Peter J, Po¨llath N. 2004. Holocene faunas of ogy. Orlando, FL: Academic Press. Res 63:274–282. the Eastern Sahara: zoogeographical and palaeoecological aspects. In: Lauwerier RCGM, 84 Lyman RL, Power E, Lyman RJ. 2003. Quan- 42 Seddon PJ, Soorae PS. 1999. Guidelines for Plug I, editors. The future from the past: ar- tification and sampling of faunal remains in owl subspecific substitutions in wildlife restoration chaeozoology in wildlife conservation and her- pellets. J Taphonomy 1:3–14. projects. Cons Biol 13:177–184. itage management. Oxford: Oxbow Books. p 85 Bryant LD, Maser C. 1982. Classification and 43 Shrader-Frechette K. 2001. Non-indigenous 34–50. distribution. In: Thomas JW, Toweill DE, editors. species and ecological explanation. Biol Philos 63 Lyman RL, Harpole JL, Darwent C, Church R. Elk of North America: ecology and management. 16:507–519. 2002. Prehistoric occurrence of pinnipeds in the Harrisburg, PA: Stackpole Books. p 1–59. 44 Lodge DM, Shrader-Frechette K. 2003. Non- lower Columbia River. Northwest Nat 83:1–6. 86 Robbins RL, Redfearn DE, Stone CP. 1982. indigenous species: ecological explanation, envi- Refuges and elk management. In: Thomas JW, ronmental ethics, and public policy. Cons Biol 64 Levine PS, Schiewe MH. 2001. Preserving salmon biodiversity. Am Sci 89:220–227. Toweill DE, editors. Elk of North America: ecol- 17:31–37. ogy and management. Harrisburg, PA: Stackpole 45 Goigel M, Bratton S. 1983. Exotics in the 65 Hughes SS. 2004. Postcontact changes in Books. p 479–507. parks. National Parks 57:24–29. the behavior and distribution of Rocky Moun- tain bighorn sheep (Ovis canadensis) in north- 87 Polziehn RO, Hamr J, Mallory FF, Strobeck C. 46 Hester FE. 1991. The U. S. National Park Ser- western Wyoming. In: Lyman RL, Cannon KP, 1998. Phylogenetic status of North American wa- vice experience with exotic species. Nat Areas J editors. Zooarchaeology and conservation biol- piti (Cervus elaphus) subspecies. Can J Zool 76: 11:127–128. ogy. Salt Lake City: University of Utah Press. p 998–1010. 47 Lyman RL. 1998. White goats, white lies: the 116–135. 88 Livingston SD. 1987. Prehistoric biogeogra- abuse of science in Olympic National Park. Salt 66 Lyman RL. 1991. Late Quaternary biogeography of white-tailed deer in Washington and Ore- Lake City: University of Utah Press. phy of the pygmy rabbit (Brachylagus idahoensis) gon. J Wildl Management 51:649–654. 48 Harpole JL. 2004. Zooarchaeological implica- in eastern Washington. J Mamm 72:110–117. 89 Lyman RL. n.d. Late prehistoric and early tions for Missouri’s elk (Cervus elaphus) reintro- historic abundance of Columbian white-tailed duction effort. In: Lyman RL, Cannon KP, edi- 67 Lyman RL. 2004. Biogeographic and paleoenvironmental implications of late Quaternary deer, Portland Basin, Washington and Oregon, tors. Zooarchaeology and conservation biology. U.S.A. J Wildl Management 69. In press. Salt Lake City: University of Utah Press. p 103– pygmy rabbits (Brachylagus idahoensis) in east- 115. ern Washington. Western N Am Nat 64:1–6. 90 Boyd R. 1999. The coming of the spirit of 68 Warheit KI. 2001. Genetic diversity and pop- pestilence. Vancouver, BC: University of British 49 Langemann EG. 2004. Zooarchaeological re- Columbia Press. search in support of a reintroduction of bison to ulation differentiation of pygmy rabbits (Brachy- Banff National Park, Canada. In: Lauwerier lagus idahoensis). Olympia: Washington Depart- 91 Grayson DK. 2001. The archaeological record RCGM, Plug I, editors. The future from the past: ment of Fish and Wildlife. of human impacts on animal populations. J archaeozoology in wildlife conservation and her- 69 Grayson DK. 1987. The biogeographic history World Prehist 15:1–68. itage management. Oxford: Oxbow Books. p 79– of small mammals in the Great Basin: observa- 92 Butler VL. 2000. Resource depression on the 89. tions on the last 20,000 years. J Mamm 68:359– Northwest Coast of North America. Antiquity 74: 50 Pulliam HR. 1988. Sources, sinks, and popu- 375. 649–661. lation regulation. Am Nat 132:652–661. 70 Thines NJS, Shipley LA, Sayler RD. 2004. Ef- 93 Leopold EB, Boyd R. 1999. An ecological his- 51 Polziehn RO, Beech R, Sheraton J, Strobeck fects of cattle grazing on ecology and habitat of tory of old prairie areas in southwestern Wash- C. 1996. Genetic relationships among North Columbia Basin pygmy rabbits (Brachylagus ida- ington. In: Boyd R, editor. Indians, fire and the American bison populations. Can J Zool 74:734– hoensis). Biol Cons 119:525–534. land in the Pacific Northwest. Corvallis: Oregon 749. 71 Galbraith WA, Anderson EW. 1971. Grazing State University Press. p 139–163. 52 van Zyll de Jong CG, Gates C, Reynolds H, history of the Northwest. J Range Management 94 Dods RR. 2002. The death of Smokey Bear: Olson W. 1995. Phenotypic variation in remnant 24:6–12. the ecodisaster myth and forest management populations of North American bison. J Mamm 72 Foley P. 1994. Predicting extinction times practices in prehistoric North America. World 76:391–405. from environmental stochasticity and carrying Archaeol 33:475–487. 53 Mooney HA, Cleland EE. 2001. The evolution- capacity. Cons Biol 8:124–137. 95 Wray J, Anderson MK. 2003. Restoring Indi- ARTICLES Paleozoology and Conservation Biology 19

an-set fires to prairie ecosystems on the Olympic 99 Lyman RL, Cannon KP, editors. 2004. Zooar- cana) migration in the Upper Green River Basin Peninsula. Ecol Restoration 21:296–301. chaeology and conservation biology. Salt Lake of Wyoming: implications for wildlife manage- 96 Germonpre´ M. 2004. Influence of climate on City: University of Utah Press. ment. In: Lyman RL, Cannon KP, editors. Zoo- sexual segregation and cub mortality in penigla- 100 O’Regan HJ, Turner A. 2004. The interface archaeology and conservation biology. Salt Lake City: University of Utah Press. p 164–177. cial cave bear. In: Lauwerier RCGM, Plug I, edi- between conservation biology, palaeontology tors. The future from the past: archaeozoology in and archaeozoology: morphometrics and pop- 102 Badenhorst S, Plug I. 2004. Archaeozoology, wildlife conservation and heritage management. ulation viability analysis. In: Lauwerier RCGM, law enforcement and nature conservation in the Oxford: Oxbow Books. p 51–63. Plug I, editors. The future from the past: ar- Republic of South Africa: perspectives from the chaeozoology in wildlife conservation and her- Transvaal Museum, Pretoria. In: Lauwerier 97 Heizer RF. 1955. Primitive man as an ecologic itage management. Oxford: Oxbow Books. p RCGM, Plug I, editors. The future from the past: factor. Kroeber Anthropol Soc Papers 13:1–31. 90–96. archaeozoology in wildlife conservation and heritage 98 Baleé W, editor. 1998. Advances in historical 101 Sanders PH, Miller ME. 2004. Archaeologi- management. Oxford: Oxbow Books. p 117–121. ecology. New York: Columbia University Press. cal evidence of pronghorn (Antilocapra Ameri- © 2006 Wiley-Liss, Inc.