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Human and Canid Dietary Relationships: Comparative Stable
Isotope Analysis From the Kodiak Archipelago, Alaska
Author(s): Catherine F. West and Christine A. France
Source: Journal of Ethnobiology, 35(3):519-535.
Published By: Society of Ethnobiology
https://doi.org/10.2993/etbi-35-03-519-535.1
URL: http://www.bioone.org/doi/full/10.2993/etbi-35-03-519-535.1
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HUMAN AND CANID DIETARY RELATIONSHIPS:
COMPARATIVE STABLE ISOTOPE ANALYSIS FROM THE
KODIAK ARCHIPELAGO, ALASKA
Catherine F. West1* and Christine A. France2
Stable carbon and nitrogen isotopes are used to address the dietary relationship between humans and two
canid species at the Uyak site (KOD-145) on Kodiak Island, Alaska: dog (Canis familiaris) and red fox (Vulpes
vulpes). We assess the relative contribution of marine and terrestrial protein to each species’ diet as a measure of
their dietary relationship to people, using zooarchaeological data, food web data, and ethnohistoric observations
to interpret the results. The results suggest that dogs and foxes had different diets: the dogs are consistently
enriched in both 13C and 15N, which indicates a heavy dependence on marine protein, while the fox samples
produced both marine and terrestrial isotope values. Data from this project have the potential to expand our
understanding of human-canid relationships in this island environment and in the greater context of island
ecology, and contribute some of the first isotopic data for small terrestrial mammals in the Gulf of Alaska.
Keywords: Stable isotope analysis, Canis familiaris, Vulpes vulpes, island ecology
Introduction
During the last 10,000 years, dogs (Canis familiaris) have been found
throughout North America, including on many offshore islands (e.g., Brown
et al. 2013; Cannon et al. 1999; Haag 1948; Laffoon et al. 2013; Morey 2006; Park
1987; Rick et al. 2008; West and Jarvis 2012). The symbiotic relationship between
dogs and humans has been explored at great length, with dogs thought to have
been important companions that provided a range of benefits to ancient peoples
(for example, traction, protection, assistance in hunting, food, and ritual activity)
(Morey 2010). More recently, researchers have begun to explore the potential
impact of ancient dogs on ecosystems and organisms (Fiedel 2005; Rick et al. 2008).
One of the key issues in domestic dog research is to understand how dogs
may serve as dietary proxies for humans in an archaeological context. To explore
ancient dog diets, researchers have studied stable isotopes of carbon (d13C) and
nitrogen (d15N) in collagen from archaeological bone (e.g., Cannon et al. 1999;
Clutton-Brock and Noe-Nygaard 1990; Fischer et al. 2007; Guiry 2012, 2013; Rick
et al. 2011). These isotope values have been compared to isotope values from
humans, to faunal remains identified in archaeological sites, and to other canids
found in archaeological sites.
In coastal and island regions, where there is often a unique mix of marine
and terrestrial foods available, studies of dog remains have the potential to shed
1 Department of Archaeology, Boston University, 675 Commonwealth Avenue, Boston, MA 02215
2 Museum Conservation Institute, Smithsonian Institution, 4210 Silver Hill Road, Suitland, MD 20746
* Corresponding author (cfwest@bu.edu)
Journal of Ethnobiology 35(3): 519–535 2015
light on the interactions among dogs, humans, and both terrestrial and marine
ecosystems. In this paper, we explore the dietary relationship between dogs and
people in the Kodiak archipelago, Alaska, by asking the following questions: do
dogs and people consume the same trophic level prey in this island environment,
and is this reflected in the zooarchaeological and isotope records? How does this
compare to other canids in this context, in particular the red fox (Vulpes vulpes)?
And finally, can these data be used to interpret how humans and canids
interacted with one another in this island environment? The Kodiak archipelago
is particularly well suited to examine these dietary relationships, given the
intense maritime adaptation of the hunter-gatherers, the presence of dogs in the
archaeological record, and the suggestion in the ethnohistoric record that foxes
may have been tamed or kept as pets by Native people. The results of this study
are among the first isotopic data for small terrestrial mammals in the Gulf of
Alaska, and while the data presented here come from the Kodiak archipelago, the
questions and techniques are broadly applicable in various island contexts where
human-canid relationships are of interest.
Background
The Kodiak Archipelago
The Kodiak archipelago is a mountainous island group located in the central
Gulf of Alaska in the Northeastern Pacific Ocean (Figure 1). Glaciation and
tectonic activity have created a dynamic landform that is characterized by
a complex coastline with many bays, lagoons, and river mouths. These areas
provide protected environments for rich biological resources and human
occupants, which are supported by a productive marine environment. The
earliest people arrived on Kodiak by boat, where they built large villages,
complex social and political systems, and established seasonal rounds and
storage systems designed to support substantial populations year-round
(Fitzhugh 2003; Saltonstall and Steffian 2010; Steffian et al. 2006).
Based on zooarchaeological data, this seasonal round seems to have been
focused primarily on the marine environment and on the anadromous fish taken
from the rivers (Amorosi 1986; Clark 1974; Etnier 2002, 2011; Foster 2006; Hays
2007; Kopperl 2003; Partlow 2000; West 2009, 2014). Based on these previously
analyzed datasets, it is clear that people relied heavily on large-bodied marine
fish, particularly migratory Pacific salmon species (Oncorhynchus spp.) and cod
species (Gadidae). In addition, flatfish (Pleuronectidae), greenlings (Hexagram-
midae), sculpins (Cottidae), and occasionally herring (Clupea pallasii), are found
in smaller numbers. While both gastropods and molluscs are found at some of
the archaeological sites, shellfish remains tend to be dominated by the butter
clam (Saxidomus gigantea) when recovered. The mammalian fauna are dominated
by marine species: harbor seal (Phoca vitulina) are the most commonly recovered
marine mammal, but whales (Cetacea), sea lion (Eumetopias jubatus), northern fur
seal (Callorhinus ursinus), dolphin (Delphinidae), and very few sea otters (Enhydra
lutris) have been identified as well. Bird remains have been understudied in
Kodiak, but there is evidence for seabirds in the zooarchaeological material
(Partlow 2000; West 2014; Yesner 1989).
520 WEST and FRANCE Vol. 35, No. 3
Figure 1. The Kodiak archipelago, showing locations mentioned in the text. Map courtesy of the
Alutiiq Museum and Archaeological Repository.
2015 JOURNAL OF ETHNOBIOLOGY 521
In contrast, there were relatively few terrestrial mammal species available to
humans before the Russian and American introductions. These introductions
include a variety of small mammals (beaver, hare, red and Arctic ground
squirrel, muskrat, and Arctic fox) and game (mountain goat, Sitka black-tailed
deer, Roosevelt elk, reindeer/caribou, and moose). There is little evidence that
the Kodiak brown bear (Ursus arctos) played an important role in subsistence, but
people did harvest birds and some small mammals (red fox and river otter) from
the terrestrial environment. In sum, the human diet on Kodiak seems to have
been dominated by marine fish, with some consumption of marine mammals,
birds, and shellfish. Unfortunately, the relative contribution of plant taxa, such as
berries and seaweed, is unknown (see Mischler 2003 for an ethnographic
perspective).
Dogs and Foxes on Kodiak
Both dog and red fox have been recovered from archaeological sites around
the Kodiak archipelago (Clark 1974; Kopperl 2003; Partlow 2000; West and Jarvis
2012). Dogs are frequently associated with northern communities, and these
animals have become part of the identity and symbolism of many Native groups.
However, while dogs are commonly found in the archaeological record on
Kodiak, their role in society, their relationship to humans, and how humans
treated dogs remain unclear. As Shelikof (in Hrdlicˇka 1944:475) notes about the
dogs that lived among Kodiak’s Native people, “although there are many dogs
they do not use them.” This observation is echoed by Petroff, who wrote that, in
contrast to the northern people who use dogs for traction, “the native of Kadiak
[sic] seldom harnesses up his dogs…[t]he people of Kadiak have few sleds…”
(in Hrdlicˇka 1944:475). Similarly, Clark (1974:44) writes that he has “not
encountered any references to the Koniags using dogs for hunting or as
watchdogs,” as has been seen among the Eyak and Chugach on the coast of
mainland Alaska (Birket-Smith 1953; Birket-Smith and deLaguna 1938). Despite
the lack of ethnographic information, the archaeological record contains
substantial numbers of dog bones, as mentioned above, and many of these
specimens exhibit cut marks. Based on this evidence, both Clark (1974) and West
and Jarvis (2012) argue that the dogs may have been used as sources of food or
fur. This is supported by ethnographic and archaeological data from mainland
Alaska and the Northwest Coast that reference dogs as food and as a source of
fur, in addition to their role as work animals and in ritual activity (Birket-Smith
and deLaguna 1938; Crockford 1997; Lantis 1980).
Red foxes are also abundant in the archaeological record on Kodiak; Clark
(1974:42) argues that “the prehistoric Kodiak emphasis on foxes is exceptional.”
Like the dog, it is unclear whether they were used for food, harvested for their
pelts, or even kept as pets. The red fox is thought to be indigenous to Kodiak,
meaning that it likely colonized the area in the early post-glacial period and was
present when people arrived in the archipelago (Clark 1958; Rausch 1969). The
ethnographic record suggests foxes were taken primarily for their fur,
particularly when the practice was industrialized in the historic period (Bailey
1993; Clark 1974; Huggins 1981; Isto 2012). Clark (1974:42) notes Glottof’s 1763
observation that he traded for fox pelts during an early visit to Kodiak and that
522 WEST and FRANCE Vol. 35, No. 3
“Koniags wore garments of the skins of birds, foxes, sea otters, young caribou,
and ground squirrels,” while both Sauer and Father Gedeon observed that foxes
were hunted and their pelts traded with the Russians (Clark 1974). On the other
hand, Hrdlicˇka suggests that foxes were “domesticated” (1944:220) and
“doubtless kept as pets” (1944:479) and goes as far to suppose that Native
people had “[e]vidently some fox cult” (1944:245) based on his excavations at
Uyak. In his diary, Huggins (1981:15) also notes that Russian explorers observed,
“natives on some of the islands had tame foxes.”
Today, red foxes are common throughout the archipelago, where they are
hunted for fur. There has been relatively little research on the Kodiak red fox
itself, perhaps because the population is so healthy (Larry VanDaele, personal
communication); however, their life history has been studied across the Gulf of
Alaska, where they are considered an invasive species (Bailey 1993; Ebbert and
Byrd 2002). In general, red foxes are adaptable and can be found in many
environments, where they are omnivorous hunters and scavengers. This
flexibility means their diet is varied and may include both terrestrial and marine
animals (hunted or scavenged) and plants, although there is some suggestion
that they prefer to hunt small animals (Haltenorth and Roth 1968). On the islands
across the Gulf of Alaska, introduced red foxes are known to prey on bird species
(adults, young, and eggs) and they have been blamed for the widespread
elimination and suppression of local bird populations (Bailey 1993). Red foxes are
frequently found near human settlements and in urban areas, where they
scavenge in refuse dumps and tend to be gregarious and commensal in some
settings (Gloor et al. 2001). Their potential as tame animals or as pets has been
demonstrated in Russia, where foxes were successfully domesticated (Belyaev et
al. 1985; Trut et al. 2013).
Stable Isotopes in Dietary Reconstruction
Given the abundance of both dog and red fox remains in the archaeological
record and the ambiguous understanding of their relationship to people, stable
isotope data have the potential to illuminate the dietary relationship. In most
animals, the stable isotopes of carbon (12C, 13C) and nitrogen (14N, 15N) fractionate
during digestive processes such that the heavier isotope is concentrated in the
body tissues, and higher levels of any given food chain are more enriched in 13C
and 15N. Stable isotopes are reported in standard delta notation relating the ratio
of heavy to light isotopes compared to an international standard:
dX~ Rsample{Rstandard
 
=Rstandard
   1000
where X is the system of interest (i.e., 13C or 15N), R is the isotope ratio (i.e.,
13C/12C or 15N/14N), units are per mil (%), and the standards are V-PDB and
atmospheric air for carbon and nitrogen respectively. In general, the isotopic
enrichment at each level of the food chain is ,1% for d13C and ,3-4% for d15N
(Bocherens and Drucker 2003; DeNiro and Epstein 1981; Minagawa and Wada
1984; Post 2002; Schoeninger and DeNiro 1984; Sutoh et al. 1987). In marine
ecosystems where the food chain length tends to be much longer, these
2015 JOURNAL OF ETHNOBIOLOGY 523
fractionations are additive and the d15N and d13C of terminal marine consumers
are significantly more positive than consumers in a terrestrial food chain.
Mammalian diets derived from marine protein are generally reflected by d15N
values between +10 and +20 % and d13C values between 220 and 210 %, while
terrestrial diets produce d15N values between 0 and +10 % and d13C values
between –30 and 217% (Burton et al. 2001; Hobson and Welch 1992; Hobson et
al. 1997; Schoeninger and DeNiro 1984). The absolute abundance of the heavier
15N and 13C incorporated in body tissues depends on the amount of marine food
consumed. In the case of dogs and foxes on Kodiak Island, those with a diet most
similar to humans will likely consume a relatively high marine input given the
maritime focus on Kodiak.
The Uyak Site
To address our questions about the dietary relationship between canids and
people, we used faunal material collected at the Uyak site (KOD-145) on the west
side of Kodiak Island. Physical anthropologist Alesˇ Hrdlicˇka excavated the Uyak
site from 1933 to 1936 (Hrdlicˇka 1944; Figures 1 and 2). Also known as “Our
Point,” the Uyak site was a substantial midden that formed a peninsula
extending into Larsen Bay (Figure 2). The midden deposits were at least three
meters deep, and Hrdlicˇka and his team divided the deposits into three layers
(lower, middle, and upper) based on the artifact types recovered and on rough
deposit descriptions. In addition to extensive faunal remains, the midden
consisted of organic and stone artifacts, human remains, slate hearths, and house
structures. Because the focus of the excavation was human remains, the team
collected select whole mammal bones, including those from dogs and foxes
(Allen 1939). Hrdlicˇka (1944:325) and others (Heizer 1956; Steffian 1992) have
estimated that the occupations date from 2000 years ago until Russian contact in
the mid-eighteenth century. West and Jarvis (2012) present recent Accelerator
Mass Spectrometry (AMS) dates taken directly on the dog bones, which confirm
that the site was occupied during this time period (Table 1).
Figure 2. Larsen Bay, showing the location of the Uyak site (KOD-145) and KOD-029 (Yesner 1989).
524 WEST and FRANCE Vol. 35, No. 3
Methods
Faunal Remains Identification and Sampling
As described above, Hrdlicˇka (1944) and his team divided the Uyak midden
into three broad levels. Those from the “Deep” or lowest level of the midden
were assigned to the Early Kachemak phase (4000-2700 cal BP), the
“Intermediate” or middle level contained Late Kachemak artifacts (2700-900 cal
BP), and the “Upper” level was interpreted as the latest phase in Kodiak’s
prehistory, or the Koniag phase (900-200 cal BP) (Hrdlicˇka 1944; Saltonstall and
Steffian 2010). Hrdlicˇka (1944) collected dog and fox remains from each of these
levels; however, their context was poorly recorded and it is unclear if these
animals were intentionally buried or not (Heizer 1956).
We selected dog and fox mandibles for stable isotope analysis and
radiocarbon dating for two reasons: 1) because they were well preserved and
suitable for collagen extraction, and 2) because of their prevalence in the Uyak
faunal assemblage. While the mandibles had been previously identified as C.
familiaris and V. vulpes, we re-identified the specimens. To distinguish between
dog and fox, we used comparative materials in the Archaeobiology Lab at the
National Museum of Natural History and identification guides (Crockford,
personal communication; Crockford 1997; Gilbert 1990; Miller et al. 1964; Olsen
1980). Specifically, dog and fox mandibles in this region can be distinguished
using the following characteristics: 1) the posterior margin of the ramus in dogs is
curved and forms a prominent hook on the coronoid process, whereas foxes have
a straight margin and a straight coronoid process; 2) the body curves up at
a sharp angle in dogs, while the body is straight in foxes; 3) there is a larger gap
between the third molar (M3) and the ramus in foxes than in dogs, which show
tooth crowding; 4) dogs are frequently missing the first premolar (P1) and
occasionally the second and fourth premolars (P2 and P4). Also, the mature dogs
are overall more robust than the mature foxes in this assemblage.
Table 1. AMS dates produced by NOSAMS for the Uyak Site on C. familiaris left adult mandibles,
adapted from West and Jarvis (2012).
NMNHa
Catalog # Provenienceb NOSAMSc # Material
Radiocarbon
age range (BP)
d13C
(%,V-PDB)
Calibrated
age range
(cal BP; 2s)d
560028-27 Upper Level OS-86795 Bone 1270 ± 30 -13.9 560 ± 80
560028-29 Upper Level OS-86785 Bone 2010 ± 50 -14.7 1230 ± 150
560028-30 Middle Level OS-86791 Bone 2220 ± 30 -13.6 1440 ± 140
560028-8 Middle Level OS-86796 Bone 2630 ± 30 -12.8 1940 ± 150
560028-15 Lower Level OS-86783 Bone 2510 ± 25 -14.7 1780 ± 150
560028-24 Lower Level OS-86788 Bone 2560 ± 30 -14.6 1840 ± 150
a National Museum of Natural History.
b Provenience refers to the broad stratigraphic levels defined by Hrdlicˇka (1944).
c National Ocean Sciences Accelerator Mass Spectrometry Facility.
d Because the stable isotope analysis done for this study indicates the Uyak dogs ate a primarily marine diet similar
to other high-level marine predators (Hirons et al. 2001; Misarti et al. 2009), the dates were calibrated using Calib
version 6.0 and corrected using the Marine Reservoir Correction Database where DR5320±50 for the west side of
Kodiak Island (McNeely et al. 2006; Reimer et al. 2004; Stuiver and Reimer 1993) and % marine carbon was 100.
2015 JOURNAL OF ETHNOBIOLOGY 525
To avoid sampling the same individual twice, we selected only left
mandibles (Table 2). We chose those that showed full tooth eruption to avoid
sampling immature animals whose isotope values could be affected by nursing
or weaning (Fogel et al. 1989; Katzenburg et al. 1996). We selected a total of 30
dogs and 20 foxes (N550): 10 dogs from each stratigraphic level, 6 foxes from the
lower and middle levels, and 10 foxes from the upper level.
Stable Isotope Methods
Solid bone chunks (,500 mg) were separated from mandibles using a rotary
tool and coring bit. Collagen was extracted from solid bones pieces using
Table 2. Carbon and nitrogen isotope values for dog (C. familiaris) and red fox (V. vulpes) from the
Uyak Site (KOD-145).
Sample #a Taxon Provenienceb d15N (%, air)c d13C (%,V-PDB) C:N
Uyak 560028-1 C. familiaris Upper level 15.4 -15.4 3.4
Uyak 560028-2 C. familiaris Upper level 16.6 -15.0 3.5
Uyak 560028-4 C. familiaris Upper level 15.2 -14.6 3.4
Uyak 560028-8 C. familiaris Middle level 17.1 -12.9 3.4
Uyak 560028-9 C. familiaris Middle level 15.4 -15.2 3.5
Uyak 560028-12 C. familiaris Middle level 15.8 -14.6 3.4
Uyak 560028-13 C. familiaris Middle level 16.9 -14.1 3.6
Uyak 560028-14 C. familiaris Lower level 14.5 -14.7 3.2
Uyak 560028-15 C. familiaris Lower level 16.0 -14.8 3.5
Uyak 560028-17 C. familiaris Lower level 15.5 -15.2 3.6
Uyak 560028-18 C. familiaris Lower level 16.3 -14.8 3.5
Uyak 560028-21 C. familiaris Middle level 16.0 -13.9 3.4
Uyak 560028-22 C. familiaris Lower level 16.4 -13.0 3.4
Uyak 560028-23 C. familiaris Lower level 15.2 -15.5 3.5
Uyak 560028-24 C. familiaris Lower level 14.7 -14.7 3.4
Uyak 560028-25 C. familiaris Lower level 16.0 -14.3 3.4
Uyak 560028-26 C. familiaris Lower level 15.9 -15.1 3.5
Uyak 560028-27 C. familiaris Upper level 16.7 -14.0 3.5
Uyak 560028-28 C. familiaris Upper level 15.9 -14.8 3.4
Uyak 560028-29 C. familiaris Upper level 15.0 -14.8 3.4
Uyak 560028-30 C. familiaris Middle level 16.4 -13.8 3.4
Mean 15.8 -14.5
Standard Deviation 0.7 0.7
Sample #a Taxon Provenienceb d15N (%, air)c d13C (%,V-PDB) C:N
Uyak 560028-32 V. vulpes Lower level 8.1 -19.6 3.3
Uyak 560028-35 V. vulpes Upper level 16.7 -11.9 3.4
Uyak 560028-37 V. vulpes Middle level 7.6 -19.7 3.4
Uyak 560028-39 V. vulpes Middle level 12.2 -17.9 3.6
Uyak 560028-40 V. vulpes Middle level 13.8 -14.6 3.5
Uyak 560028-41 V. vulpes Lower level 14.2 -15.3 3.5
Uyak 560028-42 V. vulpes Lower level 7.0 -20.8 3.6
Uyak 560028-43 V. vulpes Lower level 16.0 -13.6 3.5
Uyak 560028-45 V. vulpes Middle level 10.7 -17.6 3.3
Uyak 560028-49 V. vulpes Upper level 13.0 -15.9 3.5
Mean 11.9 -16.7
Standard Deviation 3.4 2.9
a Only well-preserved samples are included in the table.
b Provenience refers to the broad stratigraphic levels defined by Hrdlicˇka (1944), which date between 560±80 and
1940±150 cal BP (see Table 1; West and Jarvis 2012).
c All d15N and d13C values are reported with an error of ±0.2% (1s).
526 WEST and FRANCE Vol. 35, No. 3
a modified Longin (1971) method. Bone pieces were sonicated in ultra-pure water
to remove sediments and labile salts, then rinsed. Samples were decalcified in
0.6M HCl at 4uC for 24 hour increments (acid replaced daily) until reaction
ceased. Samples were rinsed to neutrality then soaked in 0.125M NaOH for 24
hours at room temperature to remove humic and fulvic contaminants. The crude
gelatin was rinsed and reacted in 0.03M HCl at 95uC for 24 hours to denature the
protein. The resulting supernatant was freeze-dried to produce a purified
collagen extract.
The extracted collagen samples were weighed into tin cups and combusted in
a Costech 4010 Elemental Analyzer (EA) with a zero blank autosampler. The
resulting N2 and CO2 gases were analyzed for d15N and d13C values in a Thermo
Delta V Advantage mass spectrometer coupled to a Conflo IV interface. Raw data
was linearly corrected to a calibrated acetanilide and urea standard. All d15N and
d13C values are reported with an error of ±0.2 % (1s) which is based on
reproducibility of repeated standard and sample measurements. The atomic C:N
ratios of the collagen were also calculated based on calibration to a homogeneous
acetanilide standard. C:N ratios between 2.8 and 3.6 indicate good preservation
for isotopic analysis (Ambrose 1990; DeNiro 1985; Table 2). Those with values
outside of this range were eliminated from the analysis.
Results
The results of the stable isotope analysis are presented in Table 2 and in
Figure 3, showing a difference in variability between the dogs and the foxes.
Among the 30 dog specimens sampled, 21 produced viable collagen; 10 of the 20
red fox specimens produced viable collagen. Samples in this study showing good
preservation had a C:N value range of 3.2-3.6. The isotope results show the
following ranges: dog d15N 5 +14.5 to +17.1 %, fox d15N 5 +7.1 to +16.7 %, dog
d13C 5 215.5 to 213.0 %, fox d13C 5 220.9 to 211.9 % (Table 2). The error
reported is ±0.2 % (1s).
The results presented in Table 2 and Figure 3 show the differences in domestic
dog and red fox d15N and d13C values. Dogs show relatively little variability in both
carbon and nitrogen isotope ratios (1s 5 0.7 for both d15N and d13C) and reveal
a distinct marine signature. The foxes, in contrast, show significant variability and
produced values along the continuum from an exclusively terrestrial diet to one
dominated by marine protein (1s5 3.4 and 2.9 for d15N and d13C, respectively). A
Mann-Whitney U Test suggests the distributions of d13C and d15N values between
dogs and fox differ significantly (Mann-Whitney U5 155.5 and 178.5 for d13C and
d15N respectively, ndog 5 21, nfox510, P , 0.03 in all comparisons). Given the
excavation and sampling methods, it is impossible to assess change over time in
stable isotope values at this site.
Discussion
The stable isotope data statistical comparisons presented above suggest that
the dog diet at the Uyak site was relatively homogeneous when compared to the
2015 JOURNAL OF ETHNOBIOLOGY 527
red fox. More specifically, dogs were likely consuming a large proportion of
marine resources, while foxes were consuming resources from both the terrestrial
and marine environments. Because stable isotope data alone are inadequate to
make detailed dietary reconstructions for each species, we turn to existing stable
isotope data for Gulf of Alaska fauna and human stable isotope data from the
Larsen Bay region to interpret the results. In combination with the zooarchaeo-
logical data and canid stable isotope results presented above, this information
may be used to address our questions about the dietary relationship between
canids and people.
To estimate which resources are reflected in the dog and fox isotope data, we
must understand the isotopic profile of the local food web. Because we are not
aware of a detailed published dataset for Kodiak, we turn to isotope data
produced using modern samples across the Gulf of Alaska and the Bering Sea,
which include marine mammals, fish, and primary producers (Burton and Koch
1999; Hirons et al. 2001; Hobson et al. 1997; Misarti 2007; Misarti et al. 2009;
Satterfield and Finney 2002; Figure 3). Figure 3 displays these values as reported
by Misarti (2007) and Misarti et al. (2009), which have been corrected for the
Suess effect and for the isotopic differences between bone and tissue (Misarti
2007), as well as values reported by Satterfield and Finney (2002). The results are
certainly influenced by geographic variables, inter-lab error, and differences in
sex, age, and migration patterns within individual species (Misarti et al. 2009;
Newsome et al. 2007); however, these values provide a rough estimate of what
a generalized Kodiak food web might look like isotopically. According to the
authors of these studies, these data suggest that in this region an animal’s d13C
value reflects near shore versus offshore feeding habits, and that the d15N value
reflects trophic level, as expected. Lacking from the literature, however, are
samples of terrestrial mammals commonly found on Gulf of Alaska islands
(i.e., ground squirrel (Spermophilus parryii), tundra vole (Microtus oeconomus), and
ermine (Mustela erminea); therefore, we are unable to produce a quantitative
mixing model for the dog and fox diets and rely on the qualitative understanding
that terrestrial resources are less enriched in both 13C and 15N than marine
resources.
Using the isotopic data presented in Figure 3 and the assumption that with
each trophic level carbon enriches by ,1 % and nitrogen by ,3-4 %, the Uyak
dogs appear to have been eating a relatively large proportion of high trophic
level foods and are consuming primarily marine protein, which is consistent with
the stable isotope values for marine mammal and marine fish available in the
Kodiak archipelago. However, the variability in d15N and d13C among the dog
samples suggests that some are eating at least one full trophic level higher than
others, and dogs have enriched d13C values. As Guiry (2012) argues, this
variability may be explained by cultural factors that influence human-dog
relationships. Ethnographic data suggest it is common practice among northern
people to feed dogs salmon (Anderson 1992; Lantis 1980), though some Alaskan
dogs are reported to have been fed bone (Birket-Smith and Laguna 1938) and
“parts of the game not destined for human consumption,” such as guts and skins
(Spencer in Lantis 1980), as well as feces (Lantis 1980). Misarti et al. (2009)
observe that in both cod and salmon, the d13C values of bone collagen are almost
528 WEST and FRANCE Vol. 35, No. 3
2.5 % higher than muscle, and Schoeninger and DeNiro (1984) report that
mammal collagen d13C is 4% higher than muscle. Therefore, dogs with enriched
d13C values may have been consuming substantial amounts of bone in addition to
muscle or other flesh, while those with enriched d15N values may have been
consuming more marine mammal than fish.
We can use these results to address the first question posed in this paper: do
dogs and people consume the same trophic level prey in this maritime-oriented
community, and is this reflected in the isotopic and zooarchaeological records?
To answer this question, we first compare the dog and food web isotope values to
the d13C and d15N values of a single human element from nearby archaeological
site KOD-029 in Larsen Bay (Yesner 1989; Figures 2 and 3). The results suggest
this person’s diet was heavy in marine protein: d13C5215.2% and d15N5 +17.3%.
As described above, the Kodiak zooarchaeological record is dominated by marine
fish, particularly Pacific salmon (Oncorhynchus sp.) and cod species (Gadidae), as
well as some marine mammal. Mixing these resources would result in the human
signature seen in Figure 3: less enriched in both 13C and 15N than marine mammals
and more enriched than marine fish.
Figure 3. Carbon and nitrogen isotopes values for the archaeological samples of dog (Canis familiaris)
and fox (Vulpes vulpes) collagen produced by this study, the single archaeological human collagen
sample from KOD-029 (Yesner 1989), as well as values for modern marine mammal and fish muscle
samples reported by Misarti (2007), Misarti et al. (2009), and Satterfield and Finney (2002). These
include: Steller sea lion (Eumetopias jubatus), harbor seal (Phoca vitulina), northern fur seal (Callorhinus
ursinus), sea otter (Enhydra lutris), Pacific cod (Gadus macrocephalus), walleye pollock (Theragra
chalcogramma), king salmon (Oncorhynchus tshawytscha), chum salmon (O. keta), pink salmon (O.
gorbuscha), and sockeye salmon (O. nerka).
2015 JOURNAL OF ETHNOBIOLOGY 529
Based on the data presented in Figure 3, the human bone is similar
isotopically to the dog remains from Uyak. We know very little about this single
human sample (for instance, its age, sex, condition, and analytical methods), but
the results do suggest that while dogs and the single human were consuming
foods from a similar trophic level overall, many of the dogs have less negative
d13C values. Given the differences between bone and muscle values discussed
above, this may be because humans were eating the filets and feeding their
dogs the bony remnants after fish processing (or dogs were scavenging bone
from the middens); similarly, the dogs show less positive d15N values, which
may mean they had access to less marine mammal protein or large-bodied
marine fish.
To address our second question, how dogs compare to other canids in this
context, we turn to the results of the fox analysis. In contrast to the dogs, the fox
remains show significantly more isotopic variability, as seen in Table 2 and
Figure 3. While some foxes are clearly eating marine foods similar to dogs and
the single human, others are eating exclusively terrestrial protein. Some of the
foxes are consuming a mix of marine and terrestrial protein (Figure 3).
Finally, to address our third question: can these data be used to interpret
how humans and canids interacted with one another in this island environment?
Given the red fox behaviors, habitat requirements, and flexibility described
above, the red fox data suggest several types of human-animal interaction. First,
the isotope results may reflect natural variability in Kodiak red fox diet: some
foxes live inland where they hunt small mammals (the tundra vole [M.
oeconomus], primarily) and others live closer to the coastline where they scavenge
a variety of marine and terrestrial resources. This behavior is observed in the
Arctic fox (Vulpes lagopus) in other northern island environments, where this
comparable species has adapted to both coastal and terrestrial ecological niches
and this adaptation is reflected in their isotope profiles (Dalerum et al. 2012;
Tarroux et al. 2012). Red foxes tend to have a home range of 2-5 km2 and are
unlikely to travel long distances (Haltenorth and Roth 1968); therefore, these fox
diets may actually reflect human hunting strategy and suggest that people
traveled around the landscape to collect the foxes. The position of the Uyak site
at the mouth of Larsen Bay inside the larger Uyak Bay afforded people easy
access to both inland and coastal environments. The second possibility is that
some of these foxes lived in direct contact with people and were not simply prey
items. Those foxes with diets enriched in 15N and 13C ate a high proportion of
marine foods. Together with the ethnohistoric data, this suggests that they could
have been fed by people and possibly kept as pets, or they were scavenging
marine resources from human-deposited middens. Given the high enrichment of
some of the foxes and their reputation as scavengers, the most parsimonious
explanation is that some foxes took advantage of the massive midden at Uyak for
food.
In terms of the dogs, the ethnographic data for coastal Alaska and the dietary
data produced by this study together suggest that dogs were likely somewhere
on the commensal spectrum, whether as pets, companions, or in another
role. While the dietary data do not offer a clear explanation of the nature of the
dog-human interaction in this region, it is clear that dogs were brought to Kodiak
530 WEST and FRANCE Vol. 35, No. 3
by people and that the Uyak dogs maintained a homogenous marine diet
relatively similar to people. Given our broader understanding of dog-human
interactions in the Gulf of Alaska, therefore, the dogs were likely fed by people
directly or scavenged from the middens deposited by the people living at the
Uyak site. The cut marks on the dog remains suggest this symbiotic relationship
may have been complicated, and will be the focus of future research.
Conclusions
The results presented above suggest that the dogs and foxes sampled in the
Uyak assemblage had different diets: the carbon and nitrogen isotope data
indicate that the dogs sampled were all eating a similar high-trophic, marine diet,
while the foxes sampled represent broad diets that included both terrestrial and
marine resources. We interpret these results to suggest that these canid species
had different dietary relationships with people at the Uyak site: the dogs were
living in close proximity to people and were either being fed by people from their
own set of marine resources, or they were scavenging in the Uyak midden.
Further, among the dogs there was variable resource consumption, although they
were all eating a high proportion of marine foods. On the other hand, while their
diets suggest that some of the foxes may have been living closely with people
either as tame pets or scavengers, others were relying primarily on scavenged or
hunted terrestrial resources.
This broad difference is significant because it illustrates the possible
relationships between humans and individual animal species on this island
landscape, as visible through dietary reconstruction. The relationship between
dogs and people on Kodiak has been difficult to understand, and these data
provide a glimpse into the close relationship of these two groups with each other
and with the landscape; further research on the dog skeletal elements will help to
illuminate the nature of this close relationship (e.g., West and Jarvis 2012). On the
other hand, the foxes both underscore what we see in the dogs and offer
a contrast: those foxes that are similar to dogs isotopically were potentially
scavenging only from the marine environment; however, given their presence in
the archaeological record, their gregarious behavior, and the historic references to
foxes on Kodiak, it is possible that they were living closely with people. In
contrast, those that recorded a terrestrial diet may have simply been a prey
species. Again, further analysis of the skeletal remains, including cut marks,
body size, and pathologies, will help to inform this interpretation.
From a regional perspective, the Kodiak sample supports conclusions made
by Cannon et al. (1999) and Rick et al. (2011), who found that dogs and humans
were eating similar marine diets, and it adds to the growing body of literature
that compares dogs and humans in the North Pacific. In great contrast to the
California Channel Islands foxes sampled by Rick et al. (2011), these data suggest
that humans and foxes have lived in close proximity on Kodiak for at least
2000 years and that foxes took advantage of the human presence. Further analysis
should combine zooarchaeological data and stable isotope analysis at Uyak with
other sites on Kodiak and elsewhere in the North Pacific to expand on this
regional perspective.
2015 JOURNAL OF ETHNOBIOLOGY 531
Acknowledgments
Our gratitude goes first to the Alutiiq Museum and Archaeological Repository of
Kodiak, Alaska for encouraging and supporting work with the Uyak Site collections.
Thanks to the Smithsonian Office of Fellowships, the National Museum of Natural History
Archaeobiology Program, the Smithsonian Museum Conservation Institute, and Torben
Rick, R. Jeff Speakman, and Melinda Zeder for the opportunity and funding to perform
this research. Thanks to the National Ocean Sciences Accelerator Mass Spectrometry
(NOSAMS) facility at the Woods Hole Oceanographic Institute for providing the AMS
analysis, and to the three anonymous reviewers for their comments.
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