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Journal of Island & Coastal Archaeology, 1:165–190, 2006Copyright © 2006 Taylor & Francis Group, LLCISSN: 1556-4894 print / 1556-1828 onlineDOI:10.1080/15564890600934129

Historical Ecology andBiogeography of NorthPacific Pinnipeds: Isotopesand Ancient DNA fromThree ArchaeologicalAssemblagesMadonna L. Moss,1 Dongya Y. Yang,2, 3 Seth D. Newsome,4

Camilla F. Speller,2, 3 Iain McKechnie,3 Alan D. McMillan,3

Robert J. Losey,5 and Paul L. Koch6

1Department of Anthropology, University of Oregon, Eugene, Oregon, USA2Ancient DNA Laboratory, Department of Archaeology, Simon Fraser

University, Burnaby, British Columbia, Canada3Department of Archaeology, Simon Fraser University, Burnaby,

British Columbia, Canada4Carnegie Geophysical Lab, Washington, DC, USA5Department of Anthropology, University of Alberta, Edmonton, Canada6Department of Earth Sciences, University of California, Santa Cruz,

California, USA

ABSTRACT

Zooarchaeology has the potential to make significant contri-butions to knowledge of pinniped biogeography of import toboth archaeologists and environmental scientists. We analyzednorthern fur seal remains found in three archaeological siteslocated along the outer coast of the Northeast Pacific Ocean:Cape Addington Rockshelter in southeast Alaska, Ts’ishaa on thewest coast of Vancouver Island, and the Netarts Sandspit site

Received 17 June 2006; accepted 31 July 2006.Address correspondence to Madonna L. Moss, Department of Anthropology, University of Oregon,Eugene, OR 97403, USA. E-mail: [email protected]

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Madonna L. Moss et al.

on the Oregon Coast. These three sites occur along an 850 kmstretch of coastline between 45◦ to 55◦ N. and 123◦ to 134◦ W.,far southeast of the primary breeding area for northern fur sealstoday, located on the Pribilof Islands at 57◦ N. 170◦ W. We useancient DNA (aDNA) and carbon (δ13C) and nitrogen (δ15N)isotopes to investigate whether northern fur seal remains fromthese archaeological sites originated with migratory PribilofIslands populations. For sites located in Oregon and points north,the isotope values are not distinct from those of the Pribilof furseals. Although aDNA was recovered from three pinniped species(northern fur seal, Steller sea lion, and Guadalupe fur seal), thepaucity of published genetic data from modern northern fur sealsprevents us from distinguishing the archaeological specimensfrom modern Pribilof seals.

Keywords zooarchaeology, marine mammals, carbon and nitrogen isotopes, ancient DNA,northern fur seals, historical ecology

INTRODUCTION

Worldwide, there currently exist 33pinniped species, not counting theCaribbean monk seal, which was drivento extinction in the twentieth century(Reeves et al. 1992:8–9). Maritime peo-ples have hunted pinnipeds for mil-lennia, but harvesting intensified in re-cent centuries with the emergence ofglobal markets for furs, oil, meat, andother products (Allen 1880; Busch 1985;Scammon 1968 [1874]). In the NorthPacific, hunting during the nineteenthand twentieth centuries radically alteredthe biogeography of pinnipeds, drivingseveral species to the brink of extinc-tion. As various pinniped species re-covered under programs of governmentprotection, some late twentieth -centurypopulations now are thought to repre-sent more “natural” conditions. Archae-ological data make it clear that humanpopulations have been interacting withpinniped populations for thousands ofyears, however, and the biogeographyand behavior of North Pacific pinnipedshave fluctuated through space and time.Archaeologists and marine ecologists are

using new analytical tools in their effortsto reconstruct the historical ecology ofpinnipeds, including studies of isotopesand DNA (both ancient and modern).

Over the last 30 years, zooarchae-ologists working on faunal assemblagesfrom across the North Pacific Coast haveidentified pinniped bones at numeroussites spanning much of the Holocene.The occurrence and relative abundanceof different species at various localitieshave helped document the importanceof marine mammals to coastal and islandcommunities over time. For specificsites, archaeologists have used marinemammal assemblages to infer: 1) humanuse of particular littoral, nearshore, andoffshore habitats; 2) hunting methods,butchering patterns, and storage tech-niques; 3) seasonality of site occupation;4) degree of economic reliance on ma-rine mammals vis a vis other animals;and 5) technological, demographic,and social changes over time (e.g.,Carlson 2003; Colten 2002; Erlandsonet al. 2004:57–61; Hildebrandt 1981,1984a; Huelsbeck 1988, 1994; Lefevreet al. 1997; Lyman 1991; Stewart andStewart 1996). Beyond specific sites,

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Historical Ecology of North Pacific Pinnipeds

investigators have compiled data for sub-regions of the North Pacific to arguefor geographically more extensive pat-terns of changing resource use throughtime. Such changes have been variouslyattributed to technological evolution,adaptations to changing environments,resource intensification by growing hu-man populations, over-exploitation ofsome species and subsequent prey-switching, or some combination of these(see Etnier 2002a; Hildebrandt and Jones1992, 2002, 2004; Jones and Hildebrandt1995; Lyman 1989, 1991, 1995, 2003;Niimi 1994; Porcasi et al. 2000; Walkeret al. 1999; Yamaura 1998; Yesner 1988).

Gretchen Lyon (1937) was probablythe first scientist working along thePacific Coast to recognize discrepanciesbetween the content of archaeologicalsamples and then contemporary pat-terns of marine mammal abundance.Years later, Gustafson (1968a, 1968b),Walker and Craig (1979), and Calvert(1980) noted the presence or relativeabundance of species in precontact as-semblages that were rare or absent inthe modern habitats surrounding thesites they studied. Hildebrandt (1984b),Clark (1986), and Lyman (1988) alsodetected zoogeographic discrepanciesbetween the past and present. Recently,investigators have focused more intentlyon demonstrating the utility of study-ing pinniped remains for understandinglong-term changes in the biogeographyof key species prior to the industrialera (Burton and Koch 1999; Burtonet al. 2001, 2002; Crockford et al. 2002;Gifford-Gonzalez et al. 2005).

Jackson et al. (2001:629) pointed outthat most ecological research is basedon local field studies of short duration,conducted some time after the 1950s.Yet some pinniped populations were thefocus of massive commercial hunting inthe nineteenth and twentieth centuries,primarily for their oil and hides (e.g.,

Allen 1880; Sims 1906; Starks 1922).Even after pinniped numbers sharplydeclined, some species were subjectto continuing hunting pressure becausethey were perceived to compete withhumans for declining fisheries. For ex-ample, all the Pacific Coast states andBritish Columbia had predator controlor eradication programs for harbor seals,some that continued into the 1960s.Such programs included open seasons,government-paid hunters, and bountysystems (Mate 1981:440). It was not untilthe 1994 amendments to the 1972 Ma-rine Mammal Protection Act that inten-tional lethal take of any marine mammalwas made illegal in U.S. waters, exceptfor subsistence hunting by Alaska Na-tives or where necessary to protect hu-man life. Illegal shootings, entanglementin commercial fishing gear and marinedebris, and collisions with boats con-tinue to cause pinniped mortality, as evi-denced in stranding records (e.g., NOAA2001:15, 2003b:5). In British Columbia,harbor seals (Phoca vitulina), Californiasea lions (Zalophus californianus), andSteller sea lions (Eumetopias jubatus)can still be legally shot if they take fishheld or reared in sea cages (Fisheries andOcean Canada 2001). Despite this, somepinniped populations are reboundingand even extending their ranges, includ-ing northern elephant seals (Miroungaangustirostris), California sea lions, andthe eastern stock of Steller sea lions(e.g., Hodder et al. 1998; LeBoeuf 1996;NOAA 2003a; Pitcher et al. in press). Incontrast, other pinniped populations arein serious decline, including the PribilofIslands stock of northern fur seals andthe western stock of Steller sea lions(Gentry 1998; O’Harra 2005; Trites andLarkin 1996). Since modern patterns ofpinniped abundance, distribution, andecology reflect the distinct histories ofeach population or stock (e.g., Cooperand Stewart 1983; Howorth 1993;

JOURNAL OF ISLAND & COASTAL ARCHAEOLOGY 167


Stewart et al. 1993), they cannot beprojected onto the more ancient past.Gifford-Gonzalez et al. (2005) haveshown how studies of the historical ecol-ogy of human-pinniped interactions cancontribute not just anthropological in-sights, but biological understandings ofprecontact marine environments. Zooar-chaeological records therefore have thepotential to contribute to improvedwildlife conservation and resource man-agement (e.g., Etnier 2004; Lyman 1996,2006; Lyman and Cannon 2004).

In this paper, we use isotopic andgenetic data from three archaeologicalsites on the Northwest Coast of NorthAmerica to assess the biogeography ofone species, the northern fur seal, Cal-lorhinus ursinus. Today, northern furseals range from Japan to the Bering Seato southern California (Gentry 1998).Their breeding sites in the United Statesinclude the Pribilof Islands in the BeringSea, Bogoslof Island in the eastern Aleu-tians (since 1982), and San Miguel Island

in southern California (since 1968; Peter-son et al. 1968; See Figure 1). The bulk(74%) of the world’s Callorhinus pop-ulation breeds on the Pribilof Islands, aseries of major colonies (NOAA 2005a).These islands were the primary target ofindustrial-scale commercial sealing untilan international treaty was reached in1911 (Gentry 1998:24; Loughlin et al.1994). Modern Pribilof fur seals givebirth from late June to late July, femalesnurse their pups until weaning, andthen migrate south in November (Gentry1998:22). Females and juveniles of bothsexes migrate offshore, following “thegeneral patterns of water flow of theNorth Pacific Current and the southernboundary current of the Alaska Gyre”(Ream et al. 2005:838–839). Some fe-males and juveniles travel as far south asBaja California, while others spend thewinter foraging offshore along the coastsof California, Oregon, and Washington.Adult males spend the winter in thehigh latitudes, foraging around the

Figure 1. The location of archaelogical sites discussed in the text in relation to Pribilof Islands andSan Miguel Island northern fur seal breeding areas (prepared by lain McKechnie).

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Aleutian Islands and western Gulf ofAlaska (Gentry 1998:36; Kajimura 1984).Northern fur seals occur across theouter coast of the North Pacific be-tween November and June, and canbe locally abundant (as described forthe outer coast of Washington byMcConnaughey and McConnaughey1994 [1985]:405).

If one were to rely exclusively ona limited, normative view of twentiethcentury northern fur seal biology, thepresence of these seals in archaeologicalsites on the islands and coastlines ofthe North Pacific would most likelyindicate the presence of northern furseals migrating to and from the PribilofIslands. Yet Burton et al. (2001) notedthe marked abundance of northern furseals in archaeological assemblages dat-ing from the Middle and Late Holocenein Oregon (citing Lyman 1988, 1989)and in California. Their study of theisotopic composition of archaeologicalfur seal bones led these investigatorsto infer that northern fur seals foragedas far offshore in the distant past asthey do today, but remained in thevicinity of California year-round. Burtonet al. (2001) proposed that northern furseals were the predominant pinnipedin California prehistorically, and thatthey were hunted at mainland colonies.This argument heavily relied on datafrom northern fur seal pup remainsfound at the Moss Landing site (CA-MNT-234) in central California. Burtonet al. (2001, 2002) clearly showed thatnorthern fur seals had different breedingand migration patterns in the ancientpast than they did during the firsthalf of the twentieth century. Workingon other assemblages, Etnier (2002a)inferred the presence of previouslyunidentified fur seal colonies in theAleutian Islands and off the coast ofthe Olympic Peninsula in Washingtonthrough the osteometric analysis ofnorthern fur seal pup bones in archae-

ological sites. Based on the presenceof pre-weaning age pups, Crockfordet al. (2002) also presented a case fora locally breeding, non-migratory pop-ulation of northern fur seals along thecentral Northwest Coast, between CapeFlattery, Washington, and Barkley Soundon the west coast of Vancouver Island.

The questions we address here relateto whether or not northern fur sealremains found at three recently inves-tigated archaeological sites on the NorthPacific Coast derive from the PribilofIslands population of northern fur seals.We attempted to use the variability inthe isotopic and genetic data to assesswhether more localized northern furseal populations existed and whetherthis provides indirect evidence of pre-contact breeding sites in the vicinity ofthese archaeological sites prior to com-mercial sealing. The archaeological sitesare the Cape Addington Rockshelter (49-CRG-188) located on Noyes Island insoutheast Alaska, Ts’ishaa (DfSi-16) lo-cated in Barkley Sound on Vancouver Is-land, and the Netarts Sandspit site (35-TI-1) located on the Oregon Coast mainland(Figure 2). These three sites occur alongan 850-mile stretch of the outer coastof the Northeast Pacific. While the pre-vious archaeological studies challengingprevailing biological tenets relied onisotopic studies of pinniped bones, ouranalyses include both isotopic and ge-netic data. The sites represent widelyseparated points on both geographicand cultural spectra, providing anextensive framework within which toassess the isotopic and genetic results.

THE ARCHAEOLOGICAL SITES

Cape Addington Rockshelter (49-CRG-188) underwent limited excavations (9.5m3) in 1996 and 1997 (Moss 2004).The site is located near the southernend of Tlingit territory, not far fromthe Haida frontier. It is situated on



Noyes Island, one of the outer islands ofthe Prince of Wales Archipelago, about130 km west of Ketchikan, Alaska. Theoccupations represented at the CapeAddington Rockshelter are dated to calAD 50–1680, apparently pre-dating theHaida incursion into what is now Alaska.Of 8,918 vertebrate remains recoveredfrom the site, 74% were fish and 9%were marine mammals. The latter in-clude 292 pinniped bones, with harborseal, Steller sea lion, and northern furseal represented, in order of abundance(Table 1). Among the 20 northern furseal specimens, 10 elements were frompups (Moss 2004:183).

Extensive excavations at Ts’ishaa(DfSi-16) took place in 1999, 2000, and2001 (McMillan and St. Claire 2005). Thesite is located within the territory ofthe Tseshaht, one of the Nuu-chah-nulthFirst Nations in British Columbia. Thesite is a large ethnographically knownvillage situated on Benson Island inthe Broken Group islands in BarkleySound on the west side of VancouverIsland. The main village area dates tocal AD 80–1700 (McMillan and St. Claire2005:42–45), which is contemporane-ous with the occupation at Cape Adding-ton. From the main village area, 163 m3

were excavated from several spatiallydispersed, but contemporary parts ofthe site. The back terrace area of thesite is substantially older (5320–2950 calBP) and dates to a period when rela-tive sea level was approximately 3–4 mhigher than today in this region of theNorthwest Coast (Friele and Hutchinson1993). From the back terrace, 44.7 m3

were excavated (McMillan and St. Claire2005:72–74). Of the 48,962 vertebrate

Figure 2. Maps of archaeological sites dis-cussed in the text (drafted by IainMcKechnie).

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Table 1. Pinniped bones (NISP) from Cape Addington Rockshelter, Ts’ishaa, and Netarts.

49-CRG-188 DfSi-16 35-TI-1

Arctocephalus townsendii Guadalupe fur seal 0 0 3

Callorhinus ursinus Northern fur seal 20 2501 31

Eumetopias jubatus Steller sea lion 52 19 166

Mirounga angustirostris Northern elephant seal 0 1 0

Phoca vitulina Harbor seal 97 43 124

Zalophus californianus California sea lion 0 1 2

Otariid 31 16 175

Pinniped 92 159 141

Total 292 489 642

1Only 31 northern fur seal bones were identified from the back terrace, or older part of thesite, and all others were from the main village, dating to within the past 2000 years (Frederick andCrockford 2005:180).

Note: Three specimens identified as northern fur seal in Moss (2004) are Steller sea lion; onespecimen identified as northern fur seal in Losey (2002) is Steller sea lion. One specimen identifiedas harbor seal in Moss (2004) is Steller sea lion, but was labeled as northern fur seal whensubmitted to Newsome and later Yang. One specimen identified as northern fur seal in Losey(2002) is Guadalupe fur seal.

remains examined, fish constitute 93%and marine mammals 3%. The latterinclude 489 pinniped bones; the major-ity of those identified to species werenorthern fur seal (n = 250). The rel-ative frequency of northern fur sealsincreased through time (Frederick andCrockford 2005:180), and fur seals ofboth sexes and all age groups (includingseven elements from pups younger thanfour months) provide evidence of afur seal breeding ground in the sitevicinity according to Crockford et al.(2002) and Frederick and Crockford(2005:180–181). Fur seals dominate themammalian assemblages at almost all ex-cavated sites along western VancouverIsland and around Cape Flattery (Calvert1980; Crockford et al. 2002; Gustafson1968a; McMillan 1999:140). Harbor seal,Steller sea lion, California sea lion, andnorthern elephant seal are also repre-sented in the Ts’ishaa assemblage.

The Netarts Sandspit site (35-TI-1) islocated within Tillamook territory on thenorthern Oregon Coast, approximately

100 km west of Portland. The site wasfirst dug during the 1940s, althoughwork by professional archaeologists didnot occur until 1952 (Cressman 1952).Extensive excavations (estimated atnearly 500 m2) continued in 1956, 1957,and 1958 (Losey 2002:193; Newman1959). Losey recovered additional mate-rial in his small-scale excavations usingintensive recovery methods in 1999,2000, and 2001, and dated the siteoccupations to cal AD 1300–1800 (Losey2002:246–259). He analyzed 5108 verte-brate specimens excavated in the 1950s(curated at the University of OregonMuseum of Natural and Cultural His-tory) and 62,156 specimens from his1999–2001 investigations. Of the total67,264 vertebrate remains, over 90%were fish and 2.5% were marine mam-mals. Of the marine mammals, 642 spec-imens were pinniped bones. In orderof abundance, Steller sea lion, harborseal, northern fur seal, California sealion, and Guadalupe fur seal are rep-resented. Of the 31 northern fur seal



specimens, seven elements are frompups.

Despite substantial differences inexcavation volume and faunal recovery,the analyzed samples of pinniped bonesfrom the three sites are within thesame order of magnitude (Table 1).Occupations at Cape Addington Rock-shelter, the main village at Ts’ishaa, andat Netarts, are all dated to within thelast 2000 years (i.e., they are roughlycontemporaneous).1

CONTEMPORARY PATTERNS OFPINNIPED ABUNDANCES

Northern fur seals spend much of theirlives far offshore and are rarely seennearshore today, except for the modernbreeding sites on the Pribilof Islands inthe Bering Sea and the Channel Islandsoff southern California. In the vicinityof Cape Addington, or almost anywhereelse in southeast Alaska, northern furseals are rare. Alaska Department ofFish and Game biologist Ken Pitcher(1998, pers. comm. to Moss) reportedthat several northern fur seals occasion-ally haul out on the Forrester Islandsduring the summer. The Forresters arean isolated group of small islands located68 km southwest of Cape Addington, asubstantial distance by canoe. Currently,the Forrester Islands (Lowrie Island inparticular) support the largest Steller sealion breeding site in the world at approx-imately 7,000 animals (Isleib 1992:2;Pitcher et al. In press). The Hazy Is-lands, located 66 km northwest of CapeAddington, serve as another substantialSteller sea lion breeding site with about3,000 animals counted annually (NMML2004). Near the western tip of CapeAddington itself, only 8 km (by water)from the archaeological site, an averageof 820 Steller sea lions haul out annually.Harbor seals are the most abundant

nearshore pinniped in southeast Alaska,but their absolute abundance in thevicinity of Cape Addington is not doc-umented. Harbor seals do not migratelong distances annually. Cape Addingtonlies considerably north of the modernrange of Guadalupe fur seals whereasthe occurence of California sea lionshas increased recently (Maniscalco et al.2004). Male northern elephant seals feedin the Aleutian Islands, but females andyoung generally occur south of 45◦N.(NOAA 2002:1). The primary breedingsites for northern elephant seals are inCalifornia and Baja California.

With respect to the site of Ts’ishaa,northern fur seals migrate throughBritish Columbia waters on their waynorth to the Pribilofs in April and ontheir way south in November (Kajimura1984; MacAskie 1979). The modern pop-ulations do not come inshore, but tendto forage along the edge of the conti-nental shelf, about 70 km from shore atthis latitude (Ream et al. 2005). There areno isolated offshore islands near Ts’ishaaequivalent to the Farallon (Pyle et al.2001) or Forrester Islands. The mostaccessible northern fur seals may havebeen those associated with a breedingsite (or sites) in the vicinity of CapeFlattery, inferred by Etnier (2002a) andCrockford et al. (2002). Cape Flattery isapproximately 75 km from Ts’ishaa.

Steller sea lions currently breed atthree sites in British Columbia, but theclosest to Ts’ishaa is the Scott Islandcomplex (Pitcher et al. In press), lo-cated approximately 350 km north ofBarkley Sound. Outside of the breedingseason, Steller sea lions are widely dis-persed. Today, small groups haul outnear Ts’ishaa on the northeast side ofBenson Island (less than a kilometerfrom the site, Frederick and Crockford2005:198) and on several exposed reefsin the outer Broken Group islands inBarkley Sound (Bigg 1985). Although

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California sea lions do not breed thisfar north (they breed mainly on theChannel Islands and off Baja California),some adult and subadult males over-winter in southern British Columbia(McTaggart Cowan and Guiguet 1965).Harbor seals are common year-roundin Vancouver Island waters. Northernelephant seals are “reported rarely butsporadically” from the west coast ofVancouver Island (Banfield 1974; Fred-erick and Crockford 2005:199). BarkleySound is considerably north of Mon-terey Bay, California, which is consid-ered the northern limit of the rangeof Guadalupe fur seals prior to theirnineteenth-century extirpation (NOAA2000:1).

Since Netarts Bay is positioned alongthe northern Oregon Coast, pinnipedabundances in both Washington andOregon should be considered. Today,northern fur seals are infrequently seenalong the coast except when they arestranded onshore. These tend to befemales and juveniles that migrate southfrom the Pribilofs, so they occur in Ore-gon and Washington waters during thewinter. In recent years, small numbersof northern fur seals have hauled outin the Shell Island—Simpson Reef areaoff Cape Arago (Oregon Coastal Atlas2005), but this site is 245 km south ofNetarts. Steller sea lions have no breed-ing sites on the coast of Washingtontoday, but they maintain three coloniesin southern Oregon (Rogue Reef,Orford Reef, and St. George Reef on theCalifornia border). Branded Steller sealions from the Forrester Islands breedingarea in Alaska (on Lowrie Island) andthe Rogue Reef colony in Oregon havebeen observed along the WashingtonCoast (Pitcher et al. In press). Stellersea lion pups are born at Three ArchRocks, located only 6 km north of theNetarts site, but biologists consider thenumbers of pups too few for this site to

be classified as a breeding site (RobinBrown 2005, pers. comm. to Moss).As described above, California sea lionsbreed in California, and there is noevidence that they breed in Oregontoday (McClenachan 2002), althoughLyman (1988) suggested that they didin the past. Some adult and subadultmales move north to winter in Ore-gon, Washington, and British Columbia.Those found in Oregon include animalsmoving to and from more northerlylocalities, as well as those that winterin Oregon. California sea lions also haulout at Three Arch Rocks and at thetip of Cape Lookout about 8 km southof Netarts. Harbor seals are year-roundresidents and reproduce everywherethey occur (Pearson 1968). Northernelephant seals have produced pups onShell Island in recent years, but thisappears to be the northernmost extentof their pupping range (Hodder et al.1998). Even this site is considerablynorth of the contemporary range ofGuadalupe fur seals in Monterey Bay.

To summarize, we list specific ques-tions related to northern fur seals whenarcheological findings (as briefly pre-sented in the previous section) arecompared to contemporary patterns ofabundance:

1. A few northern fur seal pup remainsfrom Cape Addington are foundin the absence of a contemporarybreeding site for this species inthe vicinity of the archaeologicalsite. Are these remains from animalsmigrating to or from the PribilofIslands, or was there a northern furseal breeding area somewhere insoutheast Alaska in the past?

2. Northern fur seals are the most abun-dant pinniped at Ts’ishaa, and furseals of both sexes and all age groupsare represented. Are these remainsfrom animals migrating to or from



the Pribilof Islands, or was therea northern fur seal breeding areasomewhere along the west coast ofVancouver Island (or the adjacentcoast of Washington) in the past?

3. The northern fur seals from Netartsinclude males, females, and pups.Are these remains from animalsmigrating to or from the PribilofIslands, animals associated with aVancouver Island or Washingtonbreeding area, or was there a breed-ing site somewhere along the Ore-gon Coast in the past?

MATERIALS AND METHODS

Moss and Losey identified the vertebrateremains from Cape Addington usingcomparative specimens in the Depart-ment of Anthropology at the Universityof Oregon. Pinniped bones from the sitewere also examined in reference to com-parative collections in the Departmentof Anthropology at Oregon State Univer-sity and the National Marine MammalLaboratory in Sand Point, Washington.Losey used collections at these same in-stitutions in his analysis of remains fromthe Netarts Sandspit site. From Ts’ishaa,Gay Frederick (Malaspina University-College, Nanaimo, British Columbia) andSusan Crockford (Pacific Identifications,Inc., Victoria, British Columbia) identi-fied the vertebrate remains using thecomparative collections of the Depart-ment of Anthropology, University ofVictoria. It is worth noting that theDNA analyses reported below revisesome of the identifications Moss (2004)and Losey (2002) reported on the CapeAddington and Netarts projects (see alsoMoss and Losey 2003).

Newsome measured the carbon andnitrogen isotope compositions of bonecollagen extracted from archaeologi-cal specimens from Cape Addington,

Ts’ishaa, and Netarts at the Stable Iso-tope Laboratory in the Departments ofEarth and Ocean Sciences, University ofCalifornia, Santa Cruz. To gauge the feed-ing behavior of the northern fur seals, healso measured the isotopic compositionfrom harbor seal bones from these samethree sites for comparison. Harbor sealsare well-documented nearshore residentforagers, whereas northern fur seals for-age offshore. The isotopic compositionfrom the remains of Steller sea lion andGuadalupe fur seal was also measured,as later identified by Yang and Spellerthrough their analyses of ancient DNA(see results). The techniques used toclean and treat the samples have beendescribed by Burton et al. (2001:110,2002:8). The isotopic results are ex-pressed as delta (δ) values where δ13Cor δ15N = ( [(Rsample(Rstandard) − 1) ×1000, and Rsample and Rstandard are the13C/12C or 15N/14N ratios of the sam-ple and the standard, respectively. Thestandards are Vienna Pee Dee Belemnitelimestone for carbon and atmosphericN2 for nitrogen, and the units are partsper thousand or per mil (�). Repeatedmeasurements of a gelatin standard (n =30) yielded a standard deviation of <0.20for both δ13C and δ15N values. Duplicateisotopic measurements were performedon ∼20% of all unknown samples andyielded an absolute difference of 0.20for both δ13C and δ15N values.

Yang and Speller extracted ancientDNA from the archaeological speci-mens from the three sites in the ded-icated Ancient DNA Laboratory in theDepartment of Archaeology at SimonFraser University. The protocols usedfor bone preparation, bone decontam-ination, and DNA extraction can befound in Yang et al. (1998, 2004,2005) and Speller et al. (2005). Boththe conservative cytochrome b and thehyper-variable D-loop control region ofmitochondrial DNA were analyzed to

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generate genetic information for thisstudy. The D-loop fragment of 199bpwas amplified by forward primer NFS-F99-DL (5-CTCCCCCTATGTACTTCGTGCA-3) and reverse primer NFS-R301-DL (5-GTACACTTTTCACAAGGGTTGCTG-3); a slighter shorter cytochromeb gene fragment of 180bp was tar-geted using forward primer NFS-F5-CtB (5-CCAACATTCGAAAAGTTCATCC-3) and reverse primer NFS-R185-CtB (5- GCTGTGGTGGTGTCTGAGGT-3). All PCR amplifications were con-ducted in a Mastercycler Personal (Ep-pendorf, Hamburg, Germany) in a 30µL reaction volume containing 50 mMKCl and 10 mM Tris-HCl, 2.5 mM MgCl2,0.2 mM dNTP, 1.5 mg/mL BSA, 0.3µM each primer, 3 µL DNA sample,and 2.25 U AmpliTaq GoldTM. PCR wasrun for 60 cycles at 94◦C for 30 sec-onds, 55◦C for 30 seconds, and 72◦Cextension for 40 seconds, with an initialdenaturing at 95◦C for 12 minutes. FiveµL of PCR product were separated byelectrophoresis on 2% agarose gel andvisualized using SYBR-Green on a DarkReader Box. PCR products were purifiedusing Qiagen’s MinEluteTM. For the ma-jority of samples, both directions weresequenced with relevant PCR primersfrom the same PCR products or differentPCR products of the same DNA sample.

RESULTS

Carbon and nitrogen isotope composi-tions of bone collagen were measuredfrom 19 archaeological specimens fromCape Addington and 29 from Netarts(identified as northern fur seals and har-bor seals). Of these same specimens, the10 identified as northern fur seal fromCape Addington and the 16 identifiedas northern fur seal from Netarts under-went genetic analysis. From Ts’ishaa, 10specimens underwent genetic analysis.

Isotopic Measurements

A host of biological and physicalvariables interact to produce the iso-topic composition of pinniped bones.Stable isotope values of plants and ani-mals in marine ecosystems vary along anumber of gradients, including latitude,nearshore versus offshore, proximity toupwelling, etc. (Aurioles et al. 2006;Kelly 2000; West et al. 2006). These iso-topic differences cascade up food websmodified by fractionation at each suc-cessive trophic level (Kelly 2000; Kurle2002; Newsome et al. 2006; Wu et al.1999). Burton and Koch (1999) and Bur-ton et al. (2001, 2002) have pioneeredinterpretation of isotopic values of pin-niped bones. They have demonstratedhow comparison between species andacross different localities can be usedto detect changes in patterns of forag-ing and migration over the millennia,specifically of northern fur seals. Ageand sex are also key variables; adult maleand adult female northern fur seals havedifferent foraging patterns, and nursingpups feed on their mother’s tissuesat a trophic level higher than nursingfemales, exhibiting enriched nitrogenisotope values.

One complication that arises whencomparing data from modern animalsto archaeological specimens is that theisotopic composition at the base ofmarine food webs may have shifteddue to anthropogenic perturbations.The best known of these is the Suesseffect. The combustion of fossil fuels,which are relatively rich in 12C, hascaused the δ13C value of many Earthsurface carbon reservoirs, including theatmosphere and surface ocean in someregions, to drop by ∼1� relative tovalues in the Late Holocene (e.g., Inder-muhle et al. 1999; Quay et al. 1992).There is debate about the extent towhich this effect would impact vertically



Figure 3. Carbon and nitrogen isotope ratios from Cape Addington, Netarts, and Monterey Bayarchaeological sites compared to values for modern specimens.

well-mixed surface waters in high-latitude regions, such as the BeringSea (Schell 2001). Here, however, weassume the effect is uniform throughoutthe North Pacific, and add 1� to δ13Cvalues from modern pinnipeds.

Table 2 shows the isotopic valuesfor individual pinniped specimens fromCape Addington and Netarts. Table 3shows mean isotopic values from pin-niped remains from these sites alongwith those from the Moss Landing sitein Monterey Bay (CA-MNT-234, Burton etal. 2001) and two modern samples (onefrom the Pribilof Islands, one from SanMiguel Island; see also Figure 3). First wecompare the harbor seal values to thoseof adult female northern fur seals. Burtonand Koch (1999) have shown that thebone collagen δ13C values in modernharbor seals as nearshore feeders are∼2� higher than those of offshore feed-ers such as northern fur seals. At MossLanding, the difference is 1.7� and atNetarts the difference is 1.3�, roughly

consistent with expectations. No adultfemale northern fur seals from CapeAddington are available for comparisonwith harbor seals.

The δ15N values of the Moss Landingharbor seals are not significantly differ-ent from that of the northern fur sealsfrom that site. In contrast, the CapeAddington harbor seal mean δ15N valuesare 1.1� lower than that of northern furseals. In this case, the high values for theCape Addington fur seals are probablydue to the age of the animals whoseremains were tested; young fur sealsare represented, but no adults. Previousinvestigators have shown that becausenursing pups feed at a higher trophiclevel than adults, their δ15N values willbe enriched (Newsome et al. 2006). Atthe Netarts site, while the harbor sealδ15N values are 0.4� lower than thatof the adult male northern fur sealsfrom the site, they are 2.4� higherthan the female northern fur seals. Thereare several possibilities to explain this

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Table 2. Carbon and nitrogen isotope values from Netarts and Cape Addington.

Provenience Species Element Sex/Age δ13C δ15NSFU#NF-

NETARTS 35-TI-1

House 12 brown

sand and shell

layer

Arctocephalus L tibia Juvenile −15.9 21.4 11

House 13 rim Callorhinus R mandible Adult male −13.7 18.6 2

House 10 fill Callorhinus R calcaneous Adult male? −14.2 19.3 14

House 13, upper

occupation

Callorhinus ilium Adult male −14.3 18.5 16

House 13 fill Callorhinus L femur Adult male −14.3 18.4 4

House 12 fill Callorhinus L femur Adult male −14.8 17.5 15

House 1 carbon

stratum

Callorhinus R femur Adult male −16.2 20.4 6

House 13, sand lens

below outside

Callorhinus L mandible Subadult

male

−14.1 17.7 9

House 13 fill Callorhinus L femur Adult female −14.8 17.0 10

House 13 shell layer

gray sand

Callorhinus R femur adult female −14.2 18.1 5

House 12 fill Callorhinus R femur Adult female −14.8 15.6 3

House 13 fill Callorhinus L femur Adult female −14.6 16.6 12

House 2 Callorhinus L femur Adult female −15.6 16.1 7

House 12 fill Callorhinus L ulna No estimate −14.6 16.3 1

House 12 fill Callorhinus L ulna Adult female −14.4 16.3 8

House 10 fill Eumetopias R astragalus Adult female −13.6 18.7 13

CAPE ADDINGTON

49-CRG-188

Unit 2/Stratum VI Callorhinus Ulna Juvenile −15.0 20.1 34

Unit 7, 23–35 cm Callorhinus Premaxilla Pup −14.6 18.9 32

Unit 7, 25–35 cm Callorhinus Maxilla Pup −14.5 18.8 37

Unit 7, 15–25 cm Callorhinus Maxilla w/canine

& 1 post-canine

Pup −14.4 18.8 30

Unit 2/Stratum IVB Callorhinus Pubis, ischium,

part of

acetabulum

Pup −14.6 18.3 35

Unit 7/ SE bulk

sample

Callorhinus Calcaneous Juvenile −15.7 19.1 33

Unit 7, 65–75 cm Eumetopias Scapula Adult −13.4 21.2 36

Unit 6, 15–25 cm Eumetopias Scapula Adult −14.8 20.3 31

Unit 6–7 slough Eumetopias Femur Juvenile −14.6 19.6 29

Unit 7, 40 cm Eumetopias Auditory bulla

and zygomatic

Adult −14.3 20.0 28



Table 3. Mean carbon and nitrogen isotope values from three archaeological sites and twomodern populations. The modern δ13C data have been corrected for the Suess effectby adding 1.0 per mil to the mean δ13C values.

δ13C δ15N

Site/Locality Mean SD Mean SD Species N Source

Pribilof (modern) −12.7 1.0 17.4 1.8 Harbor seal 37 Burton et al. 2001

−13.9 0.7 16.3 1.0 Northern fur seal

(female)

9 Burton et al. 2001


(male)

9 Burton et al. 2001

−13.3 — 17.9 — Northern fur seal 19 Hirons et al. 2001

Cape Addington −12.0 0.7 17.9 1.1 Harbor seal 9 This study


(pup-juvenile)

6 This study

−14.3 0.6 20.3 0.7 Steller sea lion 4 This study

Netarts −13.3 0.9 19.1 0.8 Harbor seal 13 This study


(female)

8 This study


(male)

6 This study

−13.6 — 18.7 — Steller sea lion 1 This study

−15.9 — 21.4 — Guadalupe fur seal 1 This study

Moss Landing −11.5 0.3 18.1 0.3 Harbor seal 13 Burton et al. 2001

−12.8 0.4 18.5 1.0 California sea lion 17 Burton et al. 2001

−13.2 0.5 18.5 1.4 Northern fur seal 61 Burton et al. 2001

San Miguel (modern) −13.5 0.4 18.4 0.4 Northern fur seal 6 Burton et al. 2001

pattern. The Netarts harbor seals appearto be somewhat 15N-enriched relative toconspecifics at other sites in Californiaand the Pacific Northwest. Harbor sealvalues of ∼18� are common in thisregion, but values up to 19� are anoma-lous, suggesting greater access to ahigher trophic level food resource (pos-sibly salmon).2 The values for femalenorthern fur seals at Netarts, in contrast,are similar to values for females from thePribilofs and other sites along the PacificCoast. Finally if the male northern furseals from Netarts fed offshore, as wesuspect from their δ13C values, then theyeither fed at a trophic level higher thanfemales, or they fed further south, offthe California Coast, in waters similar

to those used by animals from MossLanding.

At Cape Addington, the δ15N valuesof Steller sea lion are 1.3� and 2.4�greater than that of northern fur sealand harbor seal respectively. For theδ13C values, the Cape Addington Stellersea lions are 2.3� lower than the har-bor seals, but 0.5 � higher than thenorthern fur seals. This suggests that theSteller sea lion foraging pattern moreclosely resembles that of the northernfur seals at the site than it does thatof the harbor seals (i.e., more offshorethan nearshore). From Netarts, isotopicvalues are available from only one Stellersea lion. For this specimen, the δ13Cvalues are closer to that of the harbor

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seals (−0.3 �) than to the northernfur seals (+1.0�). The δ15N values ofSteller sea lion fall within the wide rangebetween the male and female northernfur seals (2.0� more than females, 0.8�less than males). The small sample sizeprecludes further discussion.

To summarize, the high δ15N valuesappear to confirm the presence of north-ern fur seals less than 1 year old at CapeAddington. We say “appear to confirm”because bone collagen takes as much as10 months to turnover, and young-of-the-year could still have high nitrogenisotope values for several months asthe weaning signal is diluted out of thesystem (Newsome et al. 2006). At bothCape Addington and Netarts, northernfur seals were feeding offshore, as indi-cated in their lower δ13C values whencompared to harbor seals, conformingto predictions (Burton and Koch 1999;Burton et al. 2001). Yet the isotopevalues do not allow us to distinguish thenorthern fur seals of Cape Addingtonor Netarts from those of the modernPribilof Islands.

Ancient DNA

Yang and Speller analyzed 37 bonesamples: 16 from Netarts (sample num-bers NF1-NF16), 10 from Ts’ishaa (NF17-NF26), and 11 from Cape Addington(NF27-NF37). The specimens from Ne-tarts and Cape Addington used for iso-topic analysis were also used for ge-netic analysis. The results show that theanalyzed samples contain good qualityDNA. Of 37 samples, 35 yielded positivePCR amplifications for the cytochromeb fragment and only one sample failedfor the D-loop amplification. When theamplified DNA sequences are comparedwith those from GenBank (Wynen et al.2001) through phylogenetic analysis,the obtained trees reveal that the cy-tochrome b fragment clearly affiliates

the ancient DNA samples with availablemodern reference DNA sequences, al-lowing for confident species identifica-tions (Figure 4). The D-loop fragment,on the other hand, reveals a significantamount of variation within the ancientsequences, although the same speciesidentifications can also be determined(Figure 5).

Several bone samples previouslyidentified morphologically as northernfur seal were determined to be fromother species through DNA analysis.These include four specimens fromCape Addington and one specimen fromNetarts identified through genetic anal-ysis as Steller sea lion.3 This discrep-ancy may reflect intrinsic difficultiesassociated with morphological identifi-cation of some fragmentary and juvenileskeletal remains. Although Steller sealion remains were identified in bothassemblages, without comparative spec-imens representing the full range of mor-phological variation among pinnipedspecies, identification errors were made.An additional specimen from Netartswas identified genetically as Guadalupefur seal (Arctocephalus townsendi).This specimen can be added to twoGuadalupe fur seal bones previouslyidentified by Losey (2002) from the site.This result is consistent with Etnier’s(2002b) study identifying Guadalupe furseal at Ozette, showing that this speciesranged further north at the time thesesites were occupied than it does to-day. Ideally, the species identificationsreported here should be confirmed withmore reference DNA sequences.

Analyses of both the D-loop andcytochrome b fragments indicate thatsample NF19 from Ts’ishaa cannot easilybe assigned to either the Guadalupe furseal (A. townsendi) or the southern furseal (A. philippii) based on its positionin the trees and its low bootstrap val-ues (below 50%). Genetic analyses of



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modern southern fur seal populationshave revealed a very close genetic re-lationship between A. philippii and A.townsendi (Wynen et al. 2001), and A.townsendi has also experienced a lossof genetic diversity (Weber et al. 2004).This makes it difficult to distinguish thetwo species based on limited fragmentsof mtDNA.

The phylogenetic analysis currentlyshows 23 haplotypes of the D-looppresent within the 29 samples identifiedas northern fur seal, representing a highdiversity of haplotypes. Because thesame haplotype is found among boththe Ts’ishaa and Netarts samples (NF21and NF14), the minimum number ofindividual northern fur seals representedat the three sites (based on aDNA) is24, with a MNI of four individuals fromCape Addington, nine from Ts’ishaa, and11 from Netarts. At this stage of theanalysis, we cannot use the aDNA dataalone to distinguish between the sameindividual, the same maternal family, ora closely related maternal lineage. Basedon morphology and provenience, how-ever, samples NF37, NF32, and NF30from Cape Addington could be the sameindividual. This would mean that at leasttwo pups and two juvenile northern furseals are represented among the samplestested from Cape Addington (Table 2).From Ts’ishaa, all nine specimens havedifferent haplotypes (i.e., they represent

nine individual animals), with a mixof adults, possibly two subadults andtwo juveniles represented (McKechnie,unpublished data). From Netarts, twosets of samples are the same haplotype(NF2, NF16, NF4 and NF9, NF5). Con-sidering the morphological and prove-nience data, NF2, NF16, and NF4 couldbe from the same individual, but NF9and NF5 cannot, resulting in an MNIvalue of 12 northern fur seals fromNetarts.

The large number of haplotypes(23 from the 29 samples) suggests amuch higher genetic variability in theprecontact populations, although therelatively small number of modern DNAhaplotypes used in this study preventsus from drawing firm conclusions re-garding the extent of recent geneticbottlenecks. The phylogenetic trees donot reveal distinctive clades within thearchaeological DNA sequences. Thesesequences are scattered across the fiveavailable modern northern fur seal DNAsequences from GenBank. The archaeo-logical samples do not cluster into sitegroupings (i.e., there is no correlationbetween the DNA haplotype and geo-graphic location).

The lack of strong geographic as-sociation fails to confirm the presenceof local northern fur seal breeding sitesalong the Pacific Coast of Oregon, BritishColumbia, and southeast Alaska. Yet the

←−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−Figure 4. Results of phylogenetic analysis of northern fur seals from Cape Addington (circles),

Ts’ishaa (squares), and Netarts (triangles) archaeological sites inferred from partialcytochrome b sequences. The phylogenetic tree displays the cytochrome b gene haplotypediversity of the 35 archaeological samples (NF27 and NF29 failed to yield DNA) based onthe amplified cytochrome b gene fragment. The tree (NJ with Kimura 2-parameter) wascomposed using Mega3 software (Kumar et al. 2004) with harbor seal (Phoca vitulina) asthe outgroup. The numbers at the nodes indicate those bootstrap values above 50% after2000 replications. As expected, less haplotype diversity was observed when compared tothat of the D-loop fragments (Figure 5) and the tree is consistent with those built by longercytochrome b gene fragments revealing the evolutionary histories of Otariidae (Wynenet al. 2001). For some reference sequences, the code after the species name in the sequencelabel is the haplotype code listed in GenBank.



182 VOLUME 1 • ISSUE 2 • 2006


DNA results cannot be used to rejectthe hypothesis that such breeding sitesexisted at some time in the past. Ifthe northern fur seal had at least twogroups according to their behaviors (oneor more “residential” or “non-migratory”at mid-latitudes and the other migra-tory from the Pribilofs), we should seegenetic difference between the groupsonly if: 1) they were reproducibly iso-lated from each other for a significantamount of time; and 2) there was noemigration/immigration between onegroup to the other. Unless populationsare geographically circumscribed (likesome terrestrial species), opportunisticswitchover cannot be ruled out. Ourlimited aDNA data are consistent with amigratory mode of behavior for northernfur seals.

Archaeologists have used the pur-ported distinction between pinnipedsthat are resident or migratory breedersto debate the nature of technologicaland cultural evolutionary change (e.g.,Hildebrandt and Jones 1992; Jones andHildebrandt 1995; Lyman 1995). YetLyman (1995:50–51) pointed out thatthe categories of resident breeder andmigratory breeder “are founded on thehistoric behaviors of these species,”and that these categories might notadequately characterize the past. Eventhough northern fur seals are generallylabeled as “migratory,” the San Miguelnorthern fur seals are now considered

a separate stock that is essentially non-migratory (NOAA 2003b). This contem-porary behavior, as well as evidenceaccumulating from archaeological stud-ies (e.g., Crockford et al. 2002; Gifford-Gonzalez et al. 2005), show that itis a mistake to consider long-distancemigrations as characteristic of all north-ern fur seals throughout the Holocene,when even today this species exhibitssignificant behavioral flexibility.

The DNA variation observed fromour phylogenetic analyses does not con-firm the hypothesis of long-term and/orisolated breeding sites used by non-migratory populations. Alternately, theDNA data do not allow us to reject thehypothesis that such breeding areas ex-isted, even if not in total isolation. At thispoint in time, due to the limited lengthof the analyzed ancient DNA fragmentsand the lack of an adequate number ofmodern DNA samples against which tocompare, we cannot use this analysis toconfirm the presence of colonies at loca-tions near the three archaeological sitesunder consideration. With the additionof more northern fur seal data pointsto GenBank, our results will be worthre-visiting in the future.

DISCUSSION AND CONCLUSIONS

Study of ancient DNA along with mor-phological analyses have allowed theidentification of a minimum of four

←−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−Figure 5. Results of phylogenetic analysis of northern fur seals from Cape Addington (circles),

Ts’ishaa (squares), and Netarts (triangles) archaeological sites inferred from partialD-loop sequences. The phylogenetic tree displays the D-loop haplotype diversity of the36 archaeological samples (NF27 failed to yield DNA) based on the amplified mtDNAcontrol region fragment. The tree (NJ with Kimura 2-parameter) was composed usingMega3 software (Kumar et al. 2004) with harbor seal (Phoca vitulina) as the outgroup. Thenumbers at the nodes indicate those bootstrap values above 50% after 2000 replications.Although the DNA fragment analyzed may not be sufficiently informative to reflectOtariidae species evolutionary histories (Wynen et al. 2001), the fragment is useful forassessing haplotype relationships within species. For some reference sequences, the codeafter the species name in the sequence label is the haplotype code listed in GenBank.



individual northern fur seals from CapeAddington, nine from Ts’ishaa, and12 from Netarts among our samples.From Cape Addington, the presence oftwo young-of-the-year and two juvenilenorthern fur seals was confirmed. TheNetarts sample we tested contains north-ern fur seal adults of both sexes andone subadult. Unfortunately, none of theseven northern fur seal pup bones foundat the site (Losey 2002) were included inthe sample that underwent isotopic oraDNA analyses. The Ts’ishaa sample wetested includes a mix of adults, possiblytwo subadults, and two juveniles. Onlyone of the bones we tested is includedamong the seven northern fur sealbones (representing seven individuals)Crockford et al. (2002:162) identified as“rookery age animals.”

Until we better understand the vari-ables involved in producing the isotopicsignatures in bone, and until more DNAdata on modern northern fur seals havebeen published, the best evidence fora breeding area may be morphologicalage estimates that show that more thana few pups are present. Of the three sitesreviewed in this paper, the sample fromTs’ishaa presents the strongest case for abreeding area somewhere in the vicinityof this site. Crockford et al. (2002:162)identified the remains of seven indi-vidual “rookery-age” northern fur seals,arguing that it is not possible that theseyoung animals migrated from the PribilofIslands. Even from Tsi’ishaa, however,the number of pups is relatively small.The numbers of northern fur seal pupsfrom Cape Addington and Netarts areeven smaller. Greater numbers of allthe age and sex groups present willbe necessary to generate robust harvestprofiles. The isotopic values of young furseals from Cape Addington are sugges-tive of “rookery age,” although becausethe weaning signal persists for so long,and because of the small sample size, we

cannot rule out the possibility that theseare stragglers or stranded animals.

For archaeological sites locatedalong the Pacific Coast from Oregonnorth to Alaska, it appears that avail-able isotopic data are hard to use asdirect evidence to support a case for oragainst the presence of local northernfur seal breeding areas. Isotopic valuesfrom northern fur seals in California aredistinct from those in the Pribilofs, butonce animals are foraging north of theNorth Pacific Current (which flows westto east between 40◦ N. to 50◦ N.), theyappear to have values similar to the mod-ern Pribilof population. One goal of theancient DNA analysis was to provide datacomplementary to the isotopic resultsto better understand the relationshipbetween prehistoric and modern north-ern fur seals. At this time, our geneticdata have not allowed us to answer thequestion of local breeding areas, partlybecause of the limited modern northernfur seal DNA data available throughGenBank. As Mulligan (2006:368) stated,a “sufficiently comprehensive and repre-sentative dataset on modern individuals”must exist to answer research questionssuch as those posed in this study.

Nevertheless, this is one of thefirst archaeological studies to reportsuccessful extraction and amplificationof ancient DNA from three pinnipedspecies: northern fur seal, Steller sealion, and Guadalupe fur seal. As moregenetic data become available, we maybe able to answer questions related tohow humans interacted with northernfur seals in the past. If ancient breedingareas of the northern fur seal can beidentified (or inferred), we might betterunderstand archaeological site season-ality and human settlement patternswithin subregions of the outer North Pa-cific Coast. Alternatively, if northern furseals harvested at these sites exhibitedthe same migratory behavior as that of

184 VOLUME 1 • ISSUE 2 • 2006


the Pribilof population today, it wouldappear that people were venturing con-siderable distances offshore in ocean-going watercraft to take northern furseals.

With additional genetic work, theconsequences of bottleneck events thatoccurred in the past might be identi-fied. Understanding the genetic diver-sity of northern fur seals is importantbecause this species has been listedas a depleted stock under the MarineMammal Protection Act of 1988 (NOAA2005b:iv). Although the size of theEastern Pacific stock of northern furseals appears healthy at an estimated888,120, pup production in recent yearshas dropped below the 1921 level onSt. Paul Island and below the 1916 levelon St. George Island (NOAA 2005b:16).From their 2004 field survey, Towellet al. (2006:489) estimated that pupproduction on the Pribilof Islands is lessthan half of what it was in 1968. Archae-ological data suggest, moreover, thatnorthern fur seals were more widely dis-tributed in the past, when they may havebeen less vulnerable to catastrophiccollapse. A loss of genetic diversity sincethe eighteenth and nineteenth centurieshas been documented for other marinemammals (Larson et al. 2002; Weberet al. 2004), and the same may be truefor northern fur seals.

To a great extent, the distributionand abundance of many of the 33 pin-niped species extant today has beenshaped by human activity over thepast 250 years. Eventually, we hopethat archaeological research can helpus establish a baseline from which toevaluate the rapid changes that haveoccurred during the era of industrialand commercial northern fur seal ex-ploitation. Clearly, more study of largersamples of northern fur seal remainsfrom other Pacific Coast archaeologicalsites is warranted.

ACKNOWLEDGEMENTS

The Cape Addington work was sup-ported by National Science Founda-tion Grant SBR-9705014 and the CraigRanger District of the Tongass NationalForest. Excavations at Ts’ishaa werejointly funded by Parks Canada andthe Tseshaht First Nation. The Ne-tarts investigations were supported bydissertation improvement grant NSF-9907019 and Sigma Xi. We are gratefulto Diane Gifford-Gonzalez and DanCosta for their pioneering interdisci-plinary work on northern fur seals,and for co-organizing the Interna-tional Workshop on Northern Fur SealEcology, Biogeography, and Manage-ment in Historic Perspective (with PaulKoch), held at the University of Cali-fornia, Santa Cruz in April, 2004. Wethank Mike Etnier for his importantresearch and for always sharing hisperspectives and insights. Moss extendsspecial thanks to Ken Pitcher (AlaskaDepartment of Fish and Game) andRobin Brown (Oregon Department ofFish and Wildlife) for sharing dataand ideas. Pam Endzweig (Universityof Oregon Museum of Natural andCultural History) facilitated loan ofthe Netarts specimens. Thanks to SusanCrockford and Gay Frederick (PacificIdentifications, Inc., Victoria, B.C.) foranalyzing the Ts’ishaa fauna and facil-itating access to specimens. Moss andLosey are grateful to Bob DeLong ofthe National Marine Fisheries Servicefor his assistance. The ancient DNAanalyses was supported in part by So-cial Sciences and Humanities ResearchCouncil of Canada (SSHRC) grants toD. Yang. We also thank Marcel VanTuinen for sharing some PCR primersequences. Sarah Padilla and KellyKim of the Simon Fraser Univer-sity Ancient DNA Laboratory providedvaluable technical assistance. The



isotopic analysis was supported by Na-tional Science Foundation Grants EAR-0000895 and OCE-0345943 awardedto Paul Koch and Diane Gifford-Gonzalez. We thank journal editors,Scott M. Fitzpatrick and Jon M. Erland-son, for their patience and helpful com-ments. We also thank reviewers R. LeeLyman, for pushing us to improve theclarity of the text, and Terry Jones, whoalso contributed to the development ofthis paper.

END NOTES

1. Of the 250 northern fur seal bones identifiedfrom Ts’ishaa, 219 are from the main village,and 31 are from the older back terrace.

2. Ideally one should control for sex amongthe harbor seal values also. Although adultmale harbor seals are somewhat larger thanadult females, faunal analysts generally do notdistinguish them osteologically.

3. The isotopic composition of only one of thefour Steller sea lion specimens from CapeAddington fell outside the range exhibited bythe northern fur seal specimens at this site.

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