dogs, humans and island ecosystems: of domestic dogs ......dogs, humans and island ecosystems: the...

12
PROOF ONLY Kennett and Winterhalder, 2006). Developments in the domesti- cation of plants and animals significantly contributed to the accel- eration of human population growth and the growing impacts of people on Earth’s natural ecosystems. A major source of such impacts is related to the translocation (intentional or uninten- tional) of both wild and domestic animals to new regions of the world. Such biotic introductions often result(ed) in significant alterations in natural landscapes and ecological communities, including numerous extinctions (Kirch and Hunt, 1997; Grayson, Introduction The domestication of animals and plants was a major milestone in human history, when numerous species around the world were selectively bred to increase their value (food, labour, protection, etc.) for people (see Bellwood, 2004; Barker, 2006; Zeder, 2006; Dogs, humans and island ecosystems: the distribution, antiquity and ecology of domestic dogs (Canis familiaris) on California’s Channel Islands, USA Torben C. Rick, 1 * Phillip L. Walker, 2 Lauren M. Willis, 1 Anna C. Noah, 3 Jon M. Erlandson, 4,5 René L. Vellanoweth, 6 Todd J. Braje 5 and Douglas J. Kennett 5 ( 1 Department of Anthropology, Southern Methodist University, Dallas TX 75275-0336, USA; 2 Department of Anthropology, University of California, Santa Barbara CA 93106-3210, USA; 3 Cotsen Institute of Archaeology, University of California, Los Angeles CA 90095-1510, USA; 4 Museum of Natural and Cultural History, University of Oregon, Eugene OR 97403-1224, USA; 5 Department of Anthropology, University of Oregon, Eugene OR 97403-1218, USA; 6 Department of Anthropology, Humboldt State University, Arcata CA 95521-8299, USA) Received 23 October 2007; revised manuscript accepted 8 April 2008 Abstract: Archaeologists have made significant contributions to our understanding of ancient island environ- ments, including the timing and implications of the introduction of non-native animals (pigs, chickens, rats, etc.) by humans. Here, we focus on the historical ecology and biogeography of domestic dogs (Canis familiaris) on California’s Channel Islands during the Holocene. Dogs are the only animal known unequivocally to have been introduced by Native Americans to the islands, but relatively little is known about their distribution, antiq- uity or influence on native island fauna and flora. We identified a minimum of 96 dogs from 42 archaeological sites on six of the eight islands. Dogs were present for at least 6000 years and appear to have increased in abun- dance through time. Our analysis suggests that dogs, along with humans and island foxes (Urocyon littoralis), would have had an impact on native animals and ecosystems, especially breeding birds and marine mammals. Dogs and island foxes likely competed with one another for food, however, and the impacts of dogs on island ecosystems may have been reduced by the presence of island foxes and the symbiotic relationship between dogs and humans. Dogs have been removed from all but one of the islands today, eliminating one of the few terres- trial carnivores present for most of the Holocene. Key words: Dogs, palaeoecology, human environmental impacts, zooarchaeology, Canis familiaris, Channel Islands, Holocene. The Holocene 18,7 (2008) pp. 01–11 © 2008 SAGE Publications 10.1177/0959683608095579 *Author for correspondence (e-mail: [email protected]) 07-095579-Rick.qxd 7/24/2008 9:20 PM Page 1

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Page 1: Dogs, humans and island ecosystems: of domestic dogs ......Dogs, humans and island ecosystems: the distribution, antiquity and ecology of domestic dogs (Canis familiaris) on California’s

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Kennett and Winterhalder, 2006). Developments in the domesti-cation of plants and animals significantly contributed to the accel-eration of human population growth and the growing impacts ofpeople on Earth’s natural ecosystems. A major source of suchimpacts is related to the translocation (intentional or uninten-tional) of both wild and domestic animals to new regions of theworld. Such biotic introductions often result(ed) in significantalterations in natural landscapes and ecological communities,including numerous extinctions (Kirch and Hunt, 1997; Grayson,

Introduction

The domestication of animals and plants was a major milestone inhuman history, when numerous species around the world wereselectively bred to increase their value (food, labour, protection,etc.) for people (see Bellwood, 2004; Barker, 2006; Zeder, 2006;

Dogs, humans and island ecosystems:the distribution, antiquity and ecologyof domestic dogs (Canis familiaris) onCalifornia’s Channel Islands, USATorben C. Rick,1* Phillip L. Walker,2 Lauren M. Willis,1 Anna C.Noah,3 Jon M. Erlandson,4,5 René L. Vellanoweth,6 Todd J. Braje5

and Douglas J. Kennett5

(1Department of Anthropology, Southern Methodist University, Dallas TX 75275-0336, USA;2Department of Anthropology, University of California, Santa Barbara CA 93106-3210, USA; 3Cotsen

Institute of Archaeology, University of California, Los Angeles CA 90095-1510, USA; 4Museum of

Natural and Cultural History, University of Oregon, Eugene OR 97403-1224, USA; 5Department of

Anthropology, University of Oregon, Eugene OR 97403-1218, USA; 6Department of Anthropology,

Humboldt State University, Arcata CA 95521-8299, USA)

Received 23 October 2007; revised manuscript accepted 8 April 2008

Abstract: Archaeologists have made significant contributions to our understanding of ancient island environ-ments, including the timing and implications of the introduction of non-native animals (pigs, chickens, rats,etc.) by humans. Here, we focus on the historical ecology and biogeography of domestic dogs (Canis familiaris)on California’s Channel Islands during the Holocene. Dogs are the only animal known unequivocally to havebeen introduced by Native Americans to the islands, but relatively little is known about their distribution, antiq-uity or influence on native island fauna and flora. We identified a minimum of 96 dogs from 42 archaeologicalsites on six of the eight islands. Dogs were present for at least 6000 years and appear to have increased in abun-dance through time. Our analysis suggests that dogs, along with humans and island foxes (Urocyon littoralis),would have had an impact on native animals and ecosystems, especially breeding birds and marine mammals.Dogs and island foxes likely competed with one another for food, however, and the impacts of dogs on islandecosystems may have been reduced by the presence of island foxes and the symbiotic relationship between dogsand humans. Dogs have been removed from all but one of the islands today, eliminating one of the few terres-trial carnivores present for most of the Holocene.

Key words: Dogs, palaeoecology, human environmental impacts, zooarchaeology, Canis familiaris, ChannelIslands, Holocene.

The Holocene 18,7 (2008) pp. 01–11

© 2008 SAGE Publications 10.1177/0959683608095579

*Author for correspondence (e-mail: [email protected])

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LY2001; Kirch, 2005), a process that continues to pose costly andglobal ecological challenges for human societies worldwide.

Some of the best documented cases of ancient animal introduc-tions and their environmental impacts come from islands, specifi-cally analyses of island zooarchaeological and palaeontologicalcollections. On Pacific Islands in Polynesia, for example, thetransport of chickens, dogs, pigs and rats had a devastating effecton native floral and faunal communities (Steadman, 1995, 2006;Kirch, 2000, 2005). Rats, often transported unintentionally asstowaways on long ocean voyages and possibly sometimes inten-tionally introduced, had particularly devastating impacts on localbird populations and played a role in avian extinctions. InSoutheast Asia and western Melanesia, hunter-gatherers and lateragriculturalists also appear to have carried animals to islands (eg,the Bismarcks, Solomons, Moluccas, Sulawesi and LesserSundas) for at least 20 000 years, contributing to very early, butstill poorly understood transformations of their endemic land-scapes (White, 2004). Introduced domestic and wild animals havealso been documented on islands in the Caribbean (dogs, guineapigs, agouti and hutia), Mediterranean (red deer, wild goats, etc.),North Atlantic (dogs, sheep, pigs, cattle, horses) and beyond,demonstrating similar practices around the world (Grayson, 2001;Newsom and Wing, 2004: 204–208). While domestic dogs mayhave had less profound ecological consequences than rats andsome other animals, they were a predator that often had an impacton naïve insular fauna. In New Zealand, for example, a single feraldog killed over half of a brown kiwi (Apteryx australis) colony of900 birds in about six weeks (Taborsky, 1988), and on theGalapagos Islands feral dogs have been observed to prey heavilyon marine iguanas (Kruuk and Snell, 1981). The middle- to late-Holocene introduction of the dingo (Canis lupus dingo) toAustralia also appears to coincide with significant ecologicalchanges, including the extinction of three vertebrates (see Corbett,1995; Johnson and Wroe, 2003; Savolainen et al., 2004)

In this paper, we provide evidence for the transport of domesticdogs (Canis familiaris) by hunter-gatherers to California’sChannel Islands during the Holocene (Figure 1). Dogs are one ofthe few domestic animals to be found in most areas of the worldamong hunter-gatherers and agriculturalists. In the Americas, theyhave a long presence probably spanning at least 10 000–8500

years (Morey and Wiant, 1992; Lupo and Janetski, 1994; Fiedel,2005; Snyder and Leonard, 2006). Recent mtDNA analysis ofmodern and ancient dogs around the world points to their domes-tication about 15 000 years ago in East Asia, with New Worlddogs currently thought to be derived from Old World populationsrather than independently domesticated from American greywolves (Leonard et al., 2002; Savolainen et al., 2002; Wayneet al., 2006). The available data suggest that dogs were presentthroughout much of the Americas (Schwartz, 1997) by the earlyHolocene, including a few potential early Holocene dogs incoastal California (Erlandson, 1994:194, 220, 222).

Owing to excellent archaeological integrity and an archaeologi-cal record spanning some 13 000 calendar years, California’sChannel Islands provide an important laboratory for investigatinganimal translocation to islands by foragers, the historical biogeog-raphy of domestic dogs, and the effects that dogs may have had onisland ecosystems. Archaeologists have known for some time thatNative Americans brought dogs to the islands (Schumacher, 1877:48; Bowers, 1890; Wagner, 1929; Nidever, 1937), but the distribu-tion, antiquity and potential effects of dogs on island ecology havereceived limited attention. A few researchers have discussed therole of dogs in Channel Island ritual (eg, Collins, 1991a; Raabet al., 1994; Hardy, 2000; Hale and Salls, 2000), but there has beenno attempt to assess when dogs were introduced or how wide-spread they became on these islands. To help fill this void, we syn-thesize published and unpublished occurrences of Channel Islanddogs, including specimens from southern California museums.

Cultural and environmental context

The eight Channel Islands are located between 20 and 98 km offthe southern California Coast, and range in size from about 2.6 to249 km2 (Table 1). The islands are divided into northern(Anacapa, Santa Cruz, Santa Rosa and San Miguel) and southern(San Clemente, Santa Catalina, San Nicolas and Santa Barbara)groups, which were never connected to the mainland during theQuaternary. The northern islands are an east–west trending chainalong the Santa Barbara Channel, but the southern islands are con-siderably more dispersed and isolated. All of the islands have a

2 The Holocene 18,7 (2008)

Figure 1 The Channel Islands and southern California Coast

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Mediterranean climate, with mild summers and generally cool,wet winters, but climatic conditions fluctuated throughout theHolocene (Kennett and Kennett, 2000; Kennett et al., 2007a).

The Channel Islands have a limited terrestrial flora and fauna,lacking many plants and animals found on the mainland(Schoenherr et al., 1999: 7–17). The marine environment sur-rounding the islands, in contrast, is rich in marine mammals,seabirds, fish and shellfish. Except for the diminutive (roughlyhouse-cat sized) island fox (Urocyon littoralis) and spotted skunk(Spilogale gracilis), and a few rodents (eg, deer mouse(Peromyscus maniculatus)), the islands lack terrestrial mammals.Subspecies of the island fox occur on each of the islands exceptAnacapa and Santa Barbara. Skunks are currently only found onSanta Cruz and Santa Rosa. Pygmy mammoths roamed the north-ern islands during the Pleistocene, but it is unclear if they persistedinto the human era (see Erlandson et al., 2004; Agenbroad et al.,2005). Vegetation communities on the islands are also distinct,including a number of endemic and relict species, but are gener-ally impoverished compared with mainland communities. Thelimited terrestrial fauna on the Channel Islands suggests thatdomestic dogs, along with the people who introduced them, wouldhave had significant effects on potentially naïve native animals,especially breeding marine mammals and seabirds. Collins (1991a,b), Vellanoweth (1998), Kennett (2005: 49) and Rick et al. (2008a)have also suggested that island foxes may have been introduced byhumans to some or all of the islands, raising further questionsabout the role of introduced animals in island ecology duringthe Holocene.

While terrestrial resources are generally limited, Channel Islandmarine environments are exceptionally productive, with theupwelling of nutrient-rich waters supporting large populations ofpinnipeds, cetaceans, seabirds, shellfish and fishes. These richmarine communities also include scores of breeding seabirds,seals and sea lions. Located on a boundary between colder cur-rents to the north and warmer currents to the south, the waters sur-rounding the Channel Islands contain a mix of cold- andwarm-water marine fauna.

These productive and diverse marine environments fosteredhuman occupation spanning the last 13 000 years (Erlandsonet al., 1996; Johnson et al., 2002). Thousands of archaeologicalsites, ranging from large shell middens and villages to small lithicscatters, have been documented on the eight islands (Kennett,2005; Rick et al., 2005). At the time of European contact, thenorthern Channel Islands were inhabited by Chumashan-speakingpeoples, while the southern islands were inhabited by Uto-Aztecan-speaking peoples (Kennett et al., 2007a, b). Although lin-guistically and culturally distinct, both groups had a similarmaritime technology and kept domestic dogs. Although theChannel Islands contain a lengthy record of human occupation,

analyses of dog skeletal remains from the Channel Islands suggestthat they are not a relict population of animals brought out by theearliest inhabitants. Instead many of these dogs appear to fallwithin the ‘small short-faced pueblo dog’ size category (cf. Colton,1970; Olsen and Olsen, 1970), which is considerably smaller thanthe earliest known North American dogs such as those found at theKoster site (Morey and Wiant, 1992). The size of these island dogssuggests that there was at least sporadic exchange of dogs betweenthe islands and mainland as part of the highly developed islandmainland exchange systems. The exchange of dogs also likelyworked against the development of a distinctive breed of island dogsuch as those in other island contexts (eg, Busuttil, 1969).

Methods

For this study, we systematically reviewed the published andunpublished literature for accounts of dogs from the CaliforniaChannel Islands. The available literature on Channel Island dogsis widely scattered, with few researchers consistently noting theamount or types of recovered dog remains. Nonetheless, we haveassembled a comprehensive data set. We also included unpub-lished data from archaeological specimens housed at the SantaBarbara Museum of Natural History and the Fowler Museum atthe University of California, Los Angeles. These and other speci-mens are part of our ongoing analysis of ancient southernCalifornian dog populations.

Table 2 presents the available domestic dog data, including siteor location, minimum number of individuals (MNI) and the ele-ments recovered or degree of skeletal completeness. We were asspecific as possible, but in many cases some or all of these data werenot available. We caution that researchers were not always clear onhow they determined if specimens were from dogs rather than otherspecies of Canis (eg, C. latrans (coyotes)) or hybrids, but we sus-pect that these species would be very rare on the Channel Islands.There are likely dog remains from early antiquarian projects on theislands during the late eighteenth and early nineteenth centurieshoused at the Smithsonian and other institutions that lack detailedprovenance and are not included in this study.

To our knowledge, there are currently no direct 14C dates forarchaeological dog remains from the Channel Islands.Consequently, in Table 2 all of the dates for the various dogremains are from 14C associations. In some cases we listed generaltime frames (eg, late Holocene) based on the presence of multiple14C dates from a single site. These dates are generally for a site orsite component rather than directly for a dog burial or dogremains. These dates provide a reasonable estimate of the age ofthe dog remains, but future direct dating of dog specimens couldsignificantly enhance their chronology. Our estimates of the age of

Torben C. Rick et al.: Dogs, humans and island ecosystems, California 3

Table 1 General attributes of the Channel Islands and number of sites known to contain dogsa

Island Area (km2) Max Distance from No. of land Native No of sites Dogelevation(m) mainland (km) mammals plant taxa wia dogs MNI b

Anacapa 2.9 283 20 2 190 – –Santa Cruz 249 753 30 12 480 11 21Santa Rosa 217 484 44 4 387 3 10San Miguel 37 253 42 3 198 7 10Santa Barbara 2.6 149 61 2 88 – –Santa Catalina 194 648 32 9 421 1 1San Nicolas 58 277 98 2 139 13 29San Clemente 145 599 79 6 272 7 25

a Physical characteristics based on Schoenherr et al. (1999:7).b MNI, minimum number of individuals.

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Torben C. Rick et al.: Dogs, humans and island ecosystems, California 5

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LYeach of these dog specimens suggest that, as elsewhere in NorthAmerica, dogs have a long history on the Channel Islands, proba-bly spanning most of the Holocene, and their presence appears toincrease significantly through time.

Results

At least 42 Channel Island archaeological sites or localities areknown to contain dog remains, with a number of other known dogparts assigned to an island but to no known location. This includesat least 96 individual dogs, a conservative value given the absenceof MNI data from many reports. A number of sites (eg, SRI-2,SCRI-240, SCLI-1524 and SNI-25) also produced multiple dogremains or sometimes multiple dog burials. Dogs were found onsix of the eight Channel Islands, with only the smallest islands(Anacapa and Santa Barbara, each about 2.5–3 km2 in total area)lacking documented dog remains (Figures 2 and 3). These twoislands have also seen the least amount of archaeological research.The larger islands had relatively sizeable human settlements dur-ing the middle and late Holocene, and most of these have a num-ber of sites with dog bones. Santa Catalina, the largest and closestto the mainland of the southern islands, has only one site with doc-umented dog remains, but it has seen little archaeological work.Lupo and Janetski (1994: 201) noted that for the entire easternGreat Basin (the state of Utah) only 29 dogs have been identifiedfrom deposits spanning the Holocene, suggesting that the densityof dogs on the Channel Islands was relatively high.

The largest number of sites or localities with dog remainsoccurs on San Nicolas Island (n = 13), followed by Santa Cruz(n = 11), San Clemente (n = 7), San Miguel (n = 7), Santa Rosa(n = 3) and Santa Catalina (n = 1). San Nicolas, at roughly 98 kmfrom the mainland, has the most sites with dogs, including severalsites with dog burials. Similarly, San Clemente Island, locatedabout 79 km offshore, has seven sites with dog remains, includinglate Holocene sites such as SCLI-1524 that produced six dog buri-als alone (Hale and Salls, 2000).

The oldest dog remains from the Channel Islands appear to bea left mandible fragment from Daisy Cave on San Miguel Island,the depth of which may correlate with the site’s early Holocene(c. 8600–10 000 cal. yr BP) deposits (Walker et al., 1978;Erlandson, 1994: 194). An early-Holocene date would make theseremains among the oldest dog bones in the Americas (see Morey,2006), but the specimen has not been directly dated and it couldbe from one of the younger site components. The possible early-Holocene age of the Daisy Cave specimen may be supported bydog remains identified in a few early mainland sites (Erlandson,1994), but stratigraphic mixing is an issue in many of these sitesand mainland specimens have not been directly dated. Four sitescontain dog remains that are either middle or early late Holocenein age. Two dog bone tools identified by Glassow et al. (2008)from Punta Arena on Santa Cruz Island date to the middleHolocene, but it is unclear if these were made from a dog that livedon the island or from tools traded or transported from the mainland.Two dogs identified from SNI-11 were from Stratum 3, datedbetween about 6500 and 4500 years ago, providing a reasonably

6 The Holocene 18,7 (2008)

Figure 2 Distribution of dog remains on the southern Channel Islands. Triangles indicate dog remains and circles indicate definitive dog burials

Figure 3 Distribution of dogs on the northern Channel Islands. Triangles indicate dog remains and circles indicate definitive dog burials

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secure middle-Holocene age for these materials. Dog remainsfrom SRI-41 on Santa Rosa Island and SCA-17 on Santa CatalinaIsland may also be middle or early late Holocene in age. Themajority of sites (n = 38) date to the late Holocene, indicating thatdog populations, similar to human populations, increased throughtime on the islands. More late-Holocene sites have been excavatedthan middle- and early-Holocene sites, however, which may con-tribute to the dearth of earlier dog evidence.

Dog remains are found in a wide range of archaeological con-texts, including shell middens and formal burials. The latter arebetter documented on the southern islands, where several sites onSan Clemente (eg, Eel Point, Lemon Tank and Big Dog Cave) andSan Nicolas islands (SNI-25) have produced formal dog burials,some with grave goods. On San Clemente, researchers have sug-gested that these dog burials, including apparently dismembereddogs, may be part of ancient religious ceremonies (Raab et al.,1994; Hardy, 2000; Hale and Salls, 2000). On the northern islandsformal dog burials are less common, but dogs have been foundfully articulated in midden deposits, where they were either dis-carded, intentionally buried or died naturally in place. Althoughsome of the San Clemente dogs may have been ritually modifiedand perhaps decapitated (Hardy, 2000), signs of butchery or otherindications of human consumption are rare. At SNI-11 a burneddog bone was identified (Bleitz, 1993), and a few San Clementesites also contained burned dog bones (Garlinghouse, 2000).However, this burning may be a product of inadvertent burning ofbones buried near to hearths or the use of fire to dispose of theremains of dogs who were not consumed. Overall, these data sug-gest that dogs were generally not consumed, except perhaps dur-ing times of scarcity. The two dog-bone pins identified byGlassow et al. (2008) from middle-Holocene deposits on SantaCruz Island suggest that dog bones were occasionally used to makeutilitarian objects. Gnaw marks that are probably from dogs, andpossibly foxes, have been noted on human remains from Eel Point(SCLI-43) on San Clemente Island (Titus and Walker, 2000: 85)and on non-human faunal remains from three Santa Cruz Islandsites (SCRI-192, -236 and -240; Noah, 2005: 120) and at SRI-2 onSanta Rosa Island. These data demonstrate that canids were alsoscavengers and agents of taphonomic disturbance.

Discussion

Dogs were a significant component of Holocene Channel Islandecosystems and were widespread during the late Holocene.Humans and dogs had a symbiotic relationship, with dogs proba-bly functioning as hunting companions, sentinel animals, pets,offal scavengers, food sources and potentially as status symbols asthey did elsewhere in the world. During periods of high interper-sonal violence, dogs may have also served an important functionby warning people of intruders (Walker and Snethkamp, 1984:142; Walker, 1989). Ethnographic data for the Chumash(Blackburn, 1975: 242), Hopi (Titiev, 1972), Navajo (Downs,1972), Chippewa (Sharp, 1976) and Mundurucú (Murphy, 1985)suggest that dogs also consume human excrement, which isimportant for disposing of waste. Dogs likely did the same on theChannel Islands.

Like their wolf ancestors, dogs are pack animals that generallyhave good hunting skills. A few ethnographic accounts ofChumash and Gabrielino peoples on the mainland, however, sug-gest that some Chumash groups may not have employed dogs forhunting, but occasionally used them for food (Kroeber, 1941;Harrington, 1942: 6–7). Given the dearth of terrestrial animals onthe Channel Islands, dogs probably were of limited use in hunting.They would hinder hunting sea mammals hauled out along thecoast, for example, because they would likely scare them into the

water before they could be taken. With their presence on theislands for at least 6000 years, we argue that dogs, together withfoxes and humans, negatively affected breeding bird and seamammal populations on the mainland portions of the islands,likely driving these animals to offshore islets or other isolatedareas. If feral dog populations were present, as they were histori-cally (see below), these impacts would have been more pro-nounced. Guthrie (1993) has identified the fossil remains ofChendytes lawi, a ground-nesting, flightless scoter, on San MiguelIsland dated to c. 12 000 RYBP (~14 000 cal. yr BP). ExtensiveChendytes breeding colonies would not have been possible withthe regular presence of dogs, foxes and humans. It is conceivablethat dogs contributed to the extinction of the flightless goose nearthe end of the middle Holocene or beginning of the late Holocene(see Guthrie, 1993; Jones et al., 2008; Rick et al., 2008b).

One of the most significant ecological impacts of ChannelIsland dogs was probably related to their propensity to disturbaggregations of breeding animals. A recent study on disturbanceto western snowy plovers (Charadrius alexandrinus nivosus), athreatened shore bird, and other birds near the Devereux Sloughon the Santa Barbara mainland demonstrated that unleashed dogshad some of the highest impacts on plovers, probably influencingreproduction, survivorship and distribution (Lafferty, 2001a, b).Since snowy plovers are also found on the Channel Islands,including a protected area near Skunk Point on Santa Rosa Island,Native American dogs may have had much the same impact onthese and other birds in the more distant past. This is especiallytrue since no disturbances to snowy plovers were observed at theprotected area of Santa Rosa today (Lafferty, 2001a, b), but in thepast Skunk Point was close to numerous large centres of NativeAmerican habitation, including the historic Chumash village ofQshiwqshiw that would have almost certainly had dogs.

Dogs would also have competed directly with island foxes forfood and other resources. Although a great deal of attention hasbeen given to the biology and ecology of island foxes (eg, Collins,1991a, 1991b; Roemer et al., 2001, 2002; Coonan et al., 2002,2005), few studies have examined the possible interactionsbetween dogs and foxes in the distant past. Collins (1991a) dis-cussed the burial of both dogs and foxes in island sites, demon-strating that people often gave the same ritual treatment to theseanimals. Collins (1991b) and Vellanoweth (1998) argued thatNative Americans introduced island foxes to the southern ChannelIslands but, based on a single ‘Pleistocene’ specimen from SantaRosa Island recovered in the 1950s by Phil Orr, it has been sug-gested that foxes arrived naturally on the northern islands prior tohuman arrival (see Collins, 1991b). However, recent direct 14Cdating of this and several other fox specimens, previously thoughtto be as much as 38 000 years old, suggests that they are all mid-dle or late Holocene in age, raising significant questions about thenatural dispersal of foxes to the northern islands (Shelley, 2001;Rick et al., 2008a). As Collins (1991a) noted, island fox behaviourmeshes well with humans, and their presence in formal burialsclearly demonstrates their importance in Island Chumash society.

Dogs and foxes appear to have lived along side one anothersince at least the middle Holocene, but it is still unclear how theywould have influnced each others’ lives. They would have com-peted for food and the small size of island foxes would have madethem potential prey for feral dogs (eg, Ralls and White, 1995).Dog pups could also have been prey for island foxes, but we sus-pect direct predation may have been limited, especially if peopleactively discouraged it. In this context, both island foxes and dogsmay have provided an additional benefit to people by eating dis-carded food and human waste, preying on rodents and other pestsaround habitation sites and providing protection from intruders.The introduction of dogs to the Channel Islands and occasionalprehistoric transport of mainland dogs to the islands could also

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have had deleterious consequences by transmitting potential lethalpathogens such as canine distemper, canine adenovirus, canineparvovirus and toxoplasma to endemic island foxes (Cliffordet al., 2006).

Despite these potential negative consequences for foxes, weknow from archaeological evidence that both canids coexisted forseveral thousand years, making it clear that potential niche overlapbetween the two species resulted in some form of equilibriumrather than competitive exclusion. The key adaptive difference thatmade this possible is likely the symbiotic relationship betweenhumans and their domestic dog companions. Interestingly, the ear-liest secure date for island foxes is about 7000 cal. yr BP, a datecomparable with the earliest secure dates available for dogs. Futureresearch on the antiquity and distribution of dogs and island foxesshould prove important for delineating the relationship betweenthese two animals and their ecological impacts.

Historically, dogs were present on all the islands and wereintroduced during successive ranching occupations since the nine-teenth century (Schoenherr et al., 1999). The fate of NativeAmerican dogs on the islands during the Historic period is poorlydocumented. Many dogs were probably left behind when Nativepeoples left the islands for mainland missions, and a few accountssuggest that ‘wild dogs’ were present historically (Schumacher,1877). Juana Maria, the lone woman of San Nicolas Island, whosurvived alone on the island for many years in the mid nineteenthcentury, had several dogs (Nidever, 1937). Juana Maria even sug-gested that her child was eaten by wild dogs (Hardy, 2000: 95).Schoenherr et al. (1999: 41) noted that feral dogs descended fromNative American dogs were removed from San Nicolas Island bysheepherders because they were impacting sheep populations.Native American dogs may have survived on the other islands aswell, but the fate or abundance of these dogs is poorly known. Inrecent years, dogs were removed and banned from some of theislands by the National Park Service and US Navy, with dogspresent only on Santa Catalina today. Canine diseases and para-sites were implicated as one component of the dramatic reductionof island fox populations during the last decade, although preda-tion by golden eagles appears to be the major cause of the foxdecline (Coonan et al., 2002, 2005; Roemer et al., 2002). Theintroduction of dogs in the distant past by Native Americans raisesquestions about the nature of such dog/fox interactions. Caninediseases may have passed back and forth between the two popula-tions in the past, but both foxes and dogs appear to have co-existedfor much of the Holocene and we argue played a role in islandecology, biogeography and Native American culture.

Conclusion

The evidence we have presented in this paper for the translocationof dogs to the Channel Islands adds to our growing understandingof the influence of Native Americans and other pre-Industrial peo-ples on ecology and biogeography (Redman, 1999; Grayson,2001; Kirch, 2005). These impacts are particularly significant onislands, which contain unique fauna and ecosystems that are gen-erally more vulnerable to human activities than continental land-masses. Although we are just beginning to understand the role ofpeople in shaping ancient Channel Island marine and terrestrialecosystems (Erlandson et al., 2004, 2005; Kennett, 2005; Braje,2007; Rick, 2007; Braje et al., 2007; Rick et al., 2008a, b), theintroduction of domestic dogs to the islands is part of a muchlarger ‘domestication’ of the Channel Islands landscape. TheChannel Islands contain unique ecosystems distinct from thenearby mainland, with a number of island endemic and relictspecies (Schoenherr et al., 1999). It is clear, however, that theseenvironments were also influenced by the direct and indirect

actions (eg, hunting, setting of wildfires, dune stabilization andde-stabilization, and animal translocations) of Native peoples forat least 13 000 calendar years.

Understanding and documenting the activities of ancient peo-ple, and the animals they introduced, we can improve models ofancient island ecosystems and significantly enhance the manage-ment and restoration of these habitats by providing importantbaseline data on how island ecosystems may have been structuredand functioned in the past. Dogs were a predator/scavenger intro-duced to the Channel Islands by humans that, along with islandfoxes and humans, influenced the biogeography and breedingbehaviour of birds, marine mammals and other animals that,largely free from predation today, breed in large numbers on sev-eral of the islands, including over 100 000 seals and sea lions onSan Miguel Island alone (Delong and Melin, 2002). The greatestecological influence of dogs would have been in the areas closestto human communities. Since Native villages and their dogs werepresent across much of the islands, especially around the coast-lines and near good water sources, these impacts may have beenfairly widespread.

There is still much to be learned about the antiquity and effectsof the prehistoric introduction of domestic dogs on the ecology ofCalifornia’s Channel Islands. At the very least, dogs may havescared marine mammals and birds off island beaches and otherlandforms, driving them to breed, roost and haul-out elsewhere. OnSan Miguel Island, all foxes were pulled into captivity for breedingstarting in the late 1990s, which resulted in the rapid spread ofground-breeding northern harriers (Circus cyaneus), western gulls(Larus occidentalis) and Brandt’s cormorants (Phalacrocoraxpenicillatus) in vulnerable and conspicuous locations around theisland (Drost et al., 2008). This significant expansion of bird breed-ing habitat provides a possible glimpse of what seabird populationsmay have looked like prior to the late Pleistocene/Holocene arrivalof foxes, dogs and humans. Given the lengthy presence of peopleand dogs on the islands, the modern situation appears to be radi-cally different from much of the Holocene.

Acknowledgements

We thank Ann Huston, Kelly Minas, Don Morris, Mark Senningand Ian Williams of Channel Islands National Park for supportingour research and enhancing our understanding of island ecologyand natural history. Steve Schwartz supported our research on SanNicolas and San Miguel islands and our work on the northernChannel Islands has been supported by Channel Islands NationalPark, the National Science Foundation, Western Parks andMonuments Association, the University of Oregon and SouthernMethodist University. Roger Colten provided data on some of thedog remains from Santa Cruz Island. We are also indebted to PaulCollins and Dan Guthrie for discussions that deepened our under-standing of island ecology and palaeoecology. We thank VirginiaButler and William Keegan for thoughtful comments on an earlierversion of this manuscript. Finally, we thank anonymous review-ers and the editors and staff of The Holocene for help in the reviewand production of this manuscript.

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