Journal of Archaeological Science 139 (2022) 105556

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Journal of Archaeological Science

journal homepage: www.elsevier.com/locate/jas

Canine companions or competitors? A multi-proxy analysis of

dog-human competition
Patricia Pillay a, b, *, Melinda S. Allen a, b, Judith Littleton a
a
Anthropology, School of Social Sciences (Te Pokapū Pūtaiao Pāpori), University of Auckland, Auckland, 1023, Aotearoa, New Zealand
b
Te Pūnaha Matatini, Centre of Research Excellence for Complex Systems, The University of Auckland (host), Auckland, New Zealand

A R T I C L E I N F O A B S T R A C T

Keywords: Globally, the place of dogs in the anthropogenic niche is varied, with dogs often tightly integrated into human
Canis lupus familiaris communities, but sometimes pushed to the margins, and occasionally persisting as independent feral pop­
Polynesian dog (kurı̄) ulations. Dog-human symbioses are correspondingly diverse, ranging from mutualistic to commensal to
Dental markers
competitive. The Pacific Islands, and Polynesia in particular, offer a useful context in which to consider dog-
Anthropogenic niche
Competitive release
human symbioses across varied socio-environmental settings. The translocation of domestic dogs across this
Interspecific (dog-human) competition oceanic region was underway more than two millennia ago, if not earlier, with dogs established on numerous
Multi-proxy analyses Pacific Islands. However, occasionally dogs were subsequently extirpated, a situation often attributed to
Aotearoa New Zealand competition between dogs and their human managers. Here we focus on how the dog-human symbiosis shifted as
colonists moved from the small, environmentally-circumscribed, islands of tropical central Polynesia, to the
largest, most ecologically diverse landmasses in the region—the islands of Aotearoa New Zealand. We hypoth­
esize that the mid-13th century settlement of Aotearoa New Zealand initially resulted in competitive release for
Polynesian dogs (i.e., relaxation of competition with humans) but as large native prey were depleted, and human
communities economically reorganised, dog-human competition arose anew. These two hypotheses are evalu­
ated using: a) country-wide data on dog distributions and abundance over time; and b) a regionally focused
analysis of dental markers relating to dog diet and health. Our results support the hypothesis of competitive
release on entry to Aotearoa New Zealand; dogs were quickly distributed across the two main islands, onto many
large offshore islands, and into varied ecological niches—where they were generally well represented and
associated with human occupations. This situation appears to have been followed by interspecific competition
midway through the Māori sequence (ca. AD 1450–1650), when both dog assemblages and dog abundances are
poorly represented. From the mid-17th century, dog population rebound is suggested, possibly accompanied by
new husbandry practices. These trends are not, however, well reflected in the regionally focused dental marker
analysis, where good oral health and adequate nutrition are indicated. Published studies of dog coprolites and
stable isotopes analyses help flesh out the dental analyses and point to avenues of future study. Our research
gives new insights into variability in dog-human symbioses across the Pacific Islands and potentially elsewhere,
with a particular focus on the conditions that give rise to competition and the value of multi-proxy analyses in
unravelling these complex entanglements.

1. Introduction formal husbandry and on-going co-evolutionary relationships (e.g.,

Dogs (Canis lupus familiaris) are the world’s oldest animal domesti­
cate and have long been an important component of the human niche (e.
g., Boschin et al., 2020; McKechnie et al., 2020; Perri, 2016; Wang et al.,
2016; Yeomans et al., 2019). Global studies demonstrate a range of
human-canid relationships, from semi-commensal engagements with
wild or feral populations, to fully domesticated partnerships shaped by
Clutton-Brock, 1980, 1995; Hixon et al., 2021; Koungoulos and Fillios,
2020; MacKinnon, 2010; McKechnie et al., 2020; Yeomans et al., 2019;
Zeder, 2012). Recent archaeological studies have focused on mutualistic
outcomes of the human-dog symbiosis, including the use of dogs as
hunting aids, for haulage, in herding, as sources of wool and food, in
rituals and otherwise (e.g., Hixon et al., 2021; Losey et al., 2018; Lupo,
2017; McKechnie et al., 2020). The “Canine Surrogacy Approach”

* Corresponding author. Anthropology, School of Social Sciences (Te Pokapū Pūtaiao Pāpori), University of Auckland, Auckland,1023, Aotearoa, New Zealand.
E-mail address: ppil534@aucklanduni.ac.nz (P. Pillay).

https://doi.org/10.1016/j.jas.2022.105556
Received 23 August 2021; Received in revised form 13 December 2021; Accepted 23 January 2022
Available online 1 February 2022
0305-4403/© 2022 Elsevier Ltd. All rights reserved.
P. Pillay et al. Journal of Archaeological Science 139 (2022) 105556

(Guiry, 2013; Hillis et al., 2020), where dogs serve as proxies for human
conditions, has also gained traction, although a growing number of
studies demonstrate that human-dog dietary overlap is often only partial
(e.g., Ames et al., 2015; Laffoon et al., 2019; Loponte et al., 2021; Perri
et al., 2019; Witt et al., 2021). Others highlight the analytical value of
using dogs (and other domestics) as alternative vantage points from
which to gain differing perspectives on the anthropogenic niche (Allen
and Craig, 2009; Craig, 2009; Littleton et al., 2015, 2020). Here we
consider another dimension of the human-dog symbiosis—the potential
for interspecific competition (between dogs and humans), and the
potentially adverse effects of this on dogs, humans, or both (e.g., Smith
and Litchfield, 2015).
Interspecific competition in general may arise from niche overlap,
especially when two species rely on the same food resources. This can
lead to competitive exclusion (Hardin, 1960), where the inferior
competitor shifts to an alternative ecological niche or, over longer time
frames, character displacement occurs under natural selection, or there
is species extinction. Competition is not, however, limited to dietary
overlap. In the case of dogs, free-roaming individuals may predate on
the eggs or juveniles of other domestic species, or those of shared wild
prey (e.g., Terry and Thaman, 2020:282). Zoonotic diseases may also be
a problem, transmitted through intestinal and external parasites or via
faecal matter (e.g., Collins, 2013; Chiodo et al., 2006). Dog attacks can
also endanger human health and feralization may give rise to dangerous
roving packs.
Interspecific competition can be reduced through several processes Fig. 1. Locations of study areas. Ocean Basemap sources: Esri, GEBCO, NOAA,
(e.g., Bolnick et al., 2010; van Valen, 1965). Niche partitioning may National Geographic, DeLorme, HERE, Geonames.org, and other contributors.
arise, with species developing divergent food or habitat preferences.
Competition may also be reduced by relocation into novel or newly arrival in Aotearoa; and b) dog-human competition increased over time.
available environments (e.g., following natural disturbances), where These two hypotheses are initially evaluated through an assessment
shared resources are potentially more abundant or new ones found. In of the temporal distribution and abundance of dogs across the whole of
the case of domestics, animal husbandry practices can alleviate Aotearoa. This is paired with an in-depth analysis of dog dentition,
competition in other ways, as for example, through physical constraints drawing on assemblages from the northern half of the North Island (Te
or managed feeding regimes. But rarely are domestics completely Ika-a-Māui), a region where human population densities were arguably
extirpated. the highest, and inter-tribal competition the most intense, in the cen­
Some of the clearest demonstrations of intensified dog-human turies leading up to western contact (AD 1769) (Anderson, 2016). Five
competition derive from the Pacific Islands, where domesticated mam­ dental markers that have previously been linked to variation in
mals (dogs and pigs) were introduced, established, but sometimes sub­ mammalian diet and/or health are considered: caries, calculus, peri­
sequently extirpated. Given that early Polynesians carried these odontal disease, tooth wear, and enamel hypoplasia. Notably, we as­
domestics on long-distance voyages across thousands of kilometres of sume that dog and human diets were not necessarily commensurate; by
open ocean, and successfully established them on dozens of islands, their extension, dogs are not being used here as proxies or surrogates for
subsequent extirpation is perhaps unexpected (Allen et al., 2001; Allo, inferring human dietary patterns. Rather, our focus is on understanding
1970; Allo Bay-Petersen, 1983; Clark et al., 2013; Cramb, 2021; Giovas, how dog diets (as inferred from dental markers) might align with or
2006; Kirch, 2000; Titcomb, 1969). Allo Bay-Petersen (1983) argued diverge from those of their human managers, across different geographic
these patterns reflect the high costs of domestic maintenance and, in settings and over time. We also use published coprolite and stable
settings where domestics were fed agricultural produce, direct compe­ isotope studies to evaluate the outcomes of our distributional and dental
tition with human populations, especially on the region’s smallest analyses.
islands (those less than 50 km2). The relationship between island size
and extirpation has been rigorously evaluated for pigs (Giovas 2006), 2. Background
who also tested for other geographic variables. Dogs, in contrast, are
often not well represented in Pacific archaeological sites and differen­ 2.1. Dogs of Aotearoa New Zealand and their socio-natural contexts
tiating faunal loss from sampling issues can be challenging. Nonetheless,
a recent analysis by Cramb (2021) identified high rates of dog extirpa­ Genetic research suggests that Pacific dogs originated in mainland
tions (>60%) across 23 island groups of Remote Oceania (Fig. 1); he Southeast Asia and were introduced to Remote Oceania around 2000
found that extirpations were particularly common on small, low and/or years ago (Greig, 2017; Greig et al., 2015, 2018). A considerable chal­
isolated islands that tended to be more spatially constrained and lenge to Pacific colonists and their dogs was the diminishing size and
vulnerable to climatic hazards. productivity of islands as they moved eastward. These geographic cir­
Our analysis focuses on the islands of Polynesia, with the aim of cumstances were especially marked with dispersal into central East
evaluating interspecific competition between dogs and humans using Polynesia, a process that commenced around the 10th to 12th centuries
assemblages from Polynesia’s largest island group, Aotearoa New Zea­ AD (Kirch et al., 2017; Sear et al., 2020). As the terminus of the East
land (hereafter Aotearoa) (Fig. 1). The abundance of dogs in the Polynesian diaspora, Aotearoa was settled somewhat later than the
Aotearoa archaeological record provides a unique opportunity to central archipelagos, from around the mid-13th century AD (Schmid
consider the occurrence of competition as a facet of the dog-human et al., 2018). These ancestors of indigenous Māori populations encoun­
symbiosis, and also assess the utility of different proxies for tracking tered comparatively vast landscapes (268,021 km2), and protein re­
such relations over space and time. We aim to test two hypotheses, sources unparalleled in diversity and abundance relative to those of their
specifically that: a) Polynesian dogs underwent competitive release on

2
P. Pillay et al.
central East Polynesian homelands. Oral traditions recount that Pacific
Island explorer Kupe, often credited with discovering Aotearoa, was
accompanied by his dog on the long and challenging voyage (Colenso,
1877). Archaeology provides further evidence that dogs were intro­
duced by the earliest colonists, along with the Polynesian rat (kiore,
Rattus exulans) (Greig et al., 2018).
George Förster (1778: 219), naturalist aboard Captain James Cook’s
HMS Resolution, provides one of the earliest written accounts of Māori
dogs, known to Māori as kurı̄, as observed in Queen Charlotte Sound: “A
good many dogs were observed in their canoes, which they seemed very
fond of, and kept tied with a string round their middle; they were of a
rough long-haired sort with pricked ears, and much resembled the
common shepherd’s cur.” Other historical accounts make clear the sig­
nificant roles dogs held in Māori society prior to European arrival (e.g.,
Barrow, 1984; Best, 1924; Buck 1949; Colenso, 1877; Crozet, 1891;
Luomala, 1960; Schwimmer, 1963). Kurı̄ were important hunting aids,
their meat a source of food, their bones used for tools, and at western
contact their skin and fur highly valued for chiefly clothing and orna­
mentation (Allo Bay-Petersen, 1979; Anderson, 1981; Coutts and
Jurisich, 1973; Greig and Walter, 2021; Taylor and Irwin, 2008).
The large land masses and abundant protein resources of Aotearoa,
coupled with the lack of native mammals, initially may have allowed for
niche expansion in both dogs and their human managers. Varied lines of
evidence show that early dogs consumed flightless moa (order Dinor­
inthiformes), fur seals (Arctocephalus forsteri), numerous species of
avifauna, as well as marine fish and shellfish (e.g., Anderson and Clark,
2001; Clark, 1997; Davidson, 1984; Horrocks et al., 2003; Nagaoka
et al., 2008; Wood et al., 2016). Presumably much of this food was
provided by human managers in the context of hunting and foraging. In
warmer regions of the country Māori communities cultivated a small
number of Polynesian crop plants, sometimes on a considerable scale,
with cultivation focused largely on annual production of sweet potato
(kūmara, Ipomoea batatas), but also of taro (Colocasia esculenta), gourd
(hue, Lagenaria siceraria) and four other minor crops (Anderson, 2016).
Over time, however, wild fauna, were reduced by hunting, habitat
alteration, and the newly introduced predators (humans, dogs, and rats).
Within a few centuries of human arrival many taxa some thirty land and
freshwater birds were extinct, including all nine species of moa (Allen­
toft et al., 2014; Nagaoka, 2002; Oskam et al., 2012; Smith, 1981, 2005,
2013). Anderson (1981) outlines several ways that kurı̄ potentially
contributed to the extinction of moa (and presumably other flightless
and ground-nesting species), including increasing the effectiveness of
human hunting, predation on chicks and eggs, and through the estab­
lishment of feral dog populations.
From the 15th century onward, Aotearoa became colder and drier
with the onset of Little Ice Age conditions. Some originally important
gardening zones in the south became unproductive and were ultimately
abandoned, and there was marked population movement northward
(Anderson, 2016; Leach and Leach, 1979). Along with intensification of
agricultural activities in the warmer north (Anderson and Petchey,
2020; Barber, 2010), there is evidence of increasing use of marine re­
sources by many communities at this time (Leach, 2006; Smith and
James-Lee, 2010). Intertribal conflict is also well attested, with the
appearance of fortified sites from the 16th century onward, largely in the
northern half of the North Island where agriculture was especially
productive (Anderson, 2016). These marked economic and social
changes in Māori societies undoubtedly had flow-on effects for kurı̄,
potentially including increasing dog-human competition. Even so, kurı̄
continued to have an important place in Māori society well into the
western contact period, being widely observed by early explorers.
Nonetheless, by the mid-19th century kurı̄ were functionally extinct—an
outcome often attributed to inter-breeding with, or competition from,
newly introduced European dogs (Allo Bay-Petersen, 1983; Clark,
1997c; Colenso, 1877). To understand how these processes affected
kurı̄-human relations, we initially examine the distribution and abun­
dance of dogs across the whole of Aotearoa.
3
Journal of Archaeological Science 139 (2022) 105556
2.2. Dental markers of diet and health
A second component of our analysis was a regionally focused study of
canine dental markers, aimed at understanding spatio-temporal vari­
ability in dog diet and health. This analysis was based on the premise
that dog developmental stages, such as birth and weaning, along with
environmental conditions, such as seasonality and disease, shape the life
history of individuals and are reflected in their teeth (Gawor, 2006;
Niemiec, 2008; Nomokonova et al., 2020; Williams and Evans, 1978;
Witzel, 2008). Additionally, stressors that impact individuals can be
accelerated or inhibited by husbandry management practices and
changes in regional or local environmental conditions (Collins, 2013;
Dobney and Ervynck, 2000:602; Littleton et al., 2020). In the present
analysis we used five proxies of kurı̄ dental health and diet: caries,
calculus, periodontal disease, tooth wear, and developmental defects,
building on the study of Pillay (2020), who also considered ante-mortem
tooth fracture and tooth loss. Our analyses also extend the prior studies
of Allo (1971) and Clark (1995, 1997b, 1997c) by including new dental
markers (i.e., enamel hypoplasia), incorporating more recently exca­
vated dog assemblages, and in making further statistical comparisons.
Before detailing our specific methods, we briefly review key factors that
affect the occurrence or character of the dental markers of interest.
Dental caries result from a combination of the frequent consumption
of fermentable carbohydrates, host susceptibility, and disturbances of
the balance between commensal and pathogenic oral bacteria (Hillson,
2005; Hale, 2009; Sheiham and James, 2015). The occurrence of caries
indicates that foods high in fermentable carbohydrates were consumed
(Sheiham and James, 2015). Although the veterinary literature suggests
that caries is relatively rare (Hale, 2009; Kyllar and Witter, 2005), the
condition has been observed in dog assemblages from other Polynesian
localities (Svihla, 1957)where sweet potato and taro were important
traditional carbohydrates.
The formation of dental calculus typically reflects an imbalance be­
tween foods producing an alkaline environment, facilitating plaque
mineralisation, and those inhibiting its formation or removal (e.g.,
gritty, acidic, or high-fibre foods) (Dobney and Brothwell, 1987; Hillson,
1979:150). Although the prior studies of Allo and Clark failed to identify
calculus in Māori kurı̄, it is commonly observed in contemporary dogs
(Harvey et al., 2015; Kyllar and Witter, 2005; Wallis et al., 2018).
Moreover, calculus has been observed in human populations of northern
Aotearoa (Campbell, 2008), suggesting that if kurı̄ were sharing food
resources with their human handlers, then they too might exhibit this
condition.
Periodontal disease is an infection of the soft tissues that ultimately
infects the alveolus. Periodontal disease in dogs has a complex aetiology
that is not directly analogous to that of humans (Page and Schroeder,
1981). This pathological condition can result from dental attrition,
irritation by calculus deposits, poor diet, or irritation from a fractured or
damaged tooth (Sauer et al., 2018). It also has been previously observed
in North Island kurı̄ (Allo, 1971).
Tooth wear is not a pathological process but is important for un­
derstanding other aspects of feeding regimes and local environments
(Greenfield and Arnold, 2008; Lemoine et al., 2014; Littleton et al.,
2015; Losey et al., 2014; Miller et al., 2017). Rates of dental wear and
tooth eruption sequences can be used to develop approximate age esti­
mates of an individual (Nomokonova et al., 2020). Importantly, given
that tooth wear increases with the age of an animal, it can also affect the
visibility of other pathologies over time. In earlier studies of kurı̄
dentition, Allo (1971) suggested tooth wear in Māori dogs varied
geographically, with North Island dogs exhibiting little wear. Clark
(1997c), in contrast, found little evidence of wear amongst early Māori
dogs, and proposed it was temporally variable, arising from a shift from
consumption of large meaty prey (e.g., moa, fur seals) to the that of
marine foods (fish and perhaps dried shellfish), and possibly processed
fern root (Aruhe, Pteridium esculentum), which was an important Māori
food, particularly in late prehistory. Thus, while tooth wear has been
P. Pillay et al. Journal of Archaeological Science 139 (2022) 105556

evidenced in Māori kurı̄, the underlying causes and patterning remain 3. Materials and methods
unresolved.
Shifts in animal husbandry practices or the rise of novel environ­ 3.1. Spatio-temporal patterning in kurı̄ distributions and abundances
mental conditions can also act as stressors and may in turn affect tooth
development. Enamel hypoplasia, which includes several forms of In collating information on the initial dispersal of kurı̄, and changes
developmental defects, can be a non-specific stress indicator in humans in their distributions and abundances over time, we drew on the
and other animals (Hillson, 2005:168–169). Enamel hypoplasia in canid following resources: a) the earlier compilations of Allo (1971) and Clark
teeth can be caused by diseases such as canine distemper but may also (1995, 1997a, 1997b); b) two regionally delimited studies of Smith and
arise from trauma and dietary deficiencies (Losey et al., 2014). In James-Lee (2010); and c) recent site reports from the Heritage New
contrast to many mammals, dog developmental stages occur in rapid Zealand Archaeological Reports Digital Library (https://www.heritage.
succession. Thus, enamel hypoplasia in dogs is typically reflective of org.nz/protecting-heritage/archaeology/digital-library). Following
short periods of time (Dobney and Ervynck, 2000; Guatelli-Steinberg preliminary survey of these sources, we selected assemblages with
and Skinner, 2000). When enamel hypoplasia forms on teeth in utero it relatively detailed faunal, stratigraphic, and chronological records for
potentially informs on the mother’s health. In contrast, defects in the our analysis (see Supplementary Materials). Radiocarbon dates from the
post-weaning period may signify a period of significant caloric demand respective sites were re-calibrated and, given the variable number and
and are likely influenced by anthropogenic factors such as dog feeding quality of 14C results from any given site, were assigned to three broad
regimes and exposure to pathogens (Malm and Jensen, 1996; McMillan, time intervals: Early (AD 1250–1450), Middle (AD 1450–1650 AD), and
2017). Consequently, developmental defects like enamel hypoplasia Late (AD 1650–1800). Based on the reported Minimum Number of In­
provide an opportunity to explore the life history of both new-borns and dividuals (MNI), the kurı̄ remains were also assigned to one of three
their mothers. Enamel hypoplasia (EH) commonly takes the form of ordinal categories: abundant (>30), common (11–30), or few (1–10).
linear grooves (linear enamel hypoplasia; hereafter LEH) or pit defects
and may occur as individual or multiple lesions. These kinds of defects 3.2. Dental study assemblages
are common in archaeological faunal dentitions (Dobney and Ervynck,
2000; Losey et al., 2014) and have been observed in Polynesian dogs as The detailed dental analysis involved eight northern North Island
well (Littleton et al., 2015; Miller et al., 2017). archaeological assemblages, dating from the late 13th to late 17th
Table 1 summarises the predicted patterns in each of these dental century. The assemblages were chosen based on three criteria: the
markers if our two hypotheses are valid. The hypothesised changes quality and quantity of kurı̄ remains, the availability and quality of
might be intensified if dog husbandry methods involved restraints (e.g., associated chronometric data, and information on the nature of on-site
tethering or penning) or, alternatively, offset if self-foraging was activities (Table 2). Although attempts were made to increase the
common. number of Middle Period assemblages, none of the available assem­
blages met the foregoing criteria. Prior to analysis, consultation was
undertaken with Māori iwi (indigenous tribes) who affiliate with regions
Table 1
Expectations for temporal patterning in North Island kurı̄ dentition (see text for represented by these collections. They not only gave approval to proceed
details). but particularly appreciated the non-destructive nature of the analysis.
The conventional radiocarbon ages (CRA) associated with the study
Dental Markers Early Middle Late Period Confounding
Period Perioda Factors assemblages were recalibrated (see SI, Table S1) using OxCal version 4.4
(Bronk Ramsey, 2020), along with the Southern Hemisphere calibration
Caries Low rates →
Moderate to Differences in
high due to sample
curve (ShCal20) (Hogg et al., 2020) for terrestrial samples, and the
availability of composition Marine20 curve (Heaton et al., 2020) for marine samples, and using
starchy root (teeth delta R values from Petchey and Schmid (2020). The sites were assigned
crops represented, to the same three temporal categories as above based on the 14C
individual ages),
determinations.
preservation
Calculus Minimal → Moderate to Differences in
high due to sample 3.3. Dental analysis methods
availability of composition
starchy root (teeth Overall, a minimum number of 135 individuals were analysed.
crops represented,
individual ages),
Specimen preservation ranged from excellent to poor, with some as­
preservation semblages being quite variable (Table 3). Each tooth was identified as
Periodontal Absent to → Moderate to Difference in deciduous or permanent, by tooth type, maxillary or mandibular, and,
Disease low high due to sample where possible, sided. Complete specimens were assigned to an age
prevalence availability of composition
group, but only if mandible or cranial structures were sufficiently
starchy root (teeth
crops represented, complete to control for measurement error. Ageing protocols were
individual ages), developed using Clark’s (1995) methods, as well as information on
preservation contemporary domestic dog dental eruption and tooth wear stages
Tooth Wear Slight to → Moderate to Age of individual, (Table 4).
moderate high due preservation,
availability of taphonomic
The MNI were calculated based on guidelines provided in Lyman
root crops & processes (2019). In stratified sites, each layer was treated as a unique context and
shellfish within each context MNI values were based on the most common side of
Developmental Low → Moderate to Animal ecology, the most abundant element. When complete skulls and mandibles were
Defects prevalence high due to anthropogenic
closely associated, they were assigned a MNI of 1 if a single individual
(enamel increased niche expansion,
hypoplasia) diseases, preservation, was suggested by the specimen’s age and context. If the provenance of a
parasite loads taphonomic specimen was unknown, then the MNI was calculated using the most
or dietary processes common element for the assemblage at the overall site level.
deficiencies Dental wear was defined using a 10-point scale from Lemoine et al.
a
The middle period is expected to be a period of transition as indicated by the (2014), modified here to an ordinal dental attrition scoring protocol for
arrows. use on kurı̄ teeth, beginning at 0 (unerupted teeth) and ending at 10

4
P. Pillay et al.
Table 2
Details of the North Island sites with kurı̄ remains used for dental analysis.
Site name (no.)a Kurı̄ MNI Region
valuesb
Early Period: AD 1250–1450
Hot Water Beach (HWB) 6 Coromandel Peninsula
(N44/69)
Whangamata Wharf 5 Coromandel Peninsula
(T12/2)
Houhora (N03/59) 40 Northland
Te Mataku (T1/358) 9 Ahuahu/Great Mercury Island
Middle Period: AD 1450–1650
Long Bay (R10/1374) 4 Auckland region
Late Period: AD 1650–1800
Airport NRD (R11/859) 12 Manukau Harbour area
Kohika (V15/80) 36 Bay of Plenty
Oruarangi Pā (N49/28) 23 Thames area, Coromandel
Peninsula
a
Site numbers from the New Zealand Archaeological Association site recording scheme.
b
MNI values are based on the material used in this study and may differ from values reported in previous studies.
(extreme wear) (Table 5). Teeth that could not be scored due to pre­
mortem fracturing or post-depositional factors were not considered.
Wear scores were averaged for each assemblage to determine if there
were differences in rate of wear across time periods. Teeth were scored
for the presence or absence of hypoplastic lesions by tooth type and
location on the crown (Boy et al., 2016; Littleton et al., 2015; Miller
et al., 2017).
To evaluate the occurrence of enamel hypoplasia across time periods,
non-parametric Kruskal-Wallis tests for statistical significance were
undertaken (Warren, 2004). Levene’s test for equal variances was used
to assess variability across each time period (Nordstokke and Zumbo,
2010). Statistical significance was accepted at a p-value of 0.05 for all
tests. All Kruskal-Wallis and Levene’s tests were performed using SPSS
(2017) version 25. Due to the small sizes of some assemblages and
non-normal data, Pearson’s Chi-Square test for independent samples and
Fisher’s exact non-parametric tests were performed using RStudio
(2015) version 3.5.1 to assess spatial and temporal patterning.
4. Results
4.1. Kuri distributions and abundances
The results (Fig. 2) show that the founding population of dogs rapidly
dispersed across the length and breadth of Aotearoa within the first two
hundred years of human arrival, presumably alongside their human
managers. Remarkably, they are also found in initial human occupations
of many larger offshore islands, as for example, Motutapu (15.1 km2),
Ahuahu/Great Mercury (18 km2), Rangitoto ki te Tonga/D’Urville (150
km2), and Rakiura/Stewart (1746 km2). Kurı̄ are also evidenced on the
southerly Auckland Islands (Anderson, 2005), on Whenua Hou/Codfish
Island (Smith and Anderson, 2009) at the southern end of the South
Island (Te Waipounamu), and on Raoul/Rangitāhua Island in the Ker­
medecs group to the north of mainland Aotearoa (Anderson, 1980),
although the chronological contexts are poorly defined for the latter
two. As a whole, these early sites encompass enormous environmental
variability, crossing-cutting multiple climatic, floristic, and geographic
zones. Early kurı̄ dispersal patterns speak to the remarkable behavioural
and physiological plasticity of Polynesian dogs, their ability for rapid
adaptive responses, and the intensity and resilience of the Māori-kurı̄
mutualism.
Dog abundances, and their areas of concentration across Aotearoa in
the Early Period, also inform on their changing place within the
anthropogenic niche. Kurı̄ are most abundant on the South Island and
specifically along the eastern seaboard, where human settlements map
Journal of Archaeological Science 139 (2022) 105556
Geographic setting Time period (AD) Reference
Open beach 14th century-early 15th century Leahy (1974); Gumbley et al.
(2018)
Mouth of estuary w/ Late 13th century Gumbley et al. (2014)
beach
River mouth w/beach 14th century Furey (2002); Petchey (2000)
Open beach 14th century - early 15th Phillipps et al. (2014)
century
Open beach Late 15th century- early 16th Campbell et al. (2019)
century
Harbour bluff Late 15th century-17th century Campbell et al. (2008)
Inland swamp 17th century Irwin (2004); Taylor and Irwin
(2008)
Inland swamp Late 15th - 17th century Furey (1996)
onto areas rich in large game (moa, seals), or which provided easy access
into the interior (Anderson, 1981). These patterns suggest South Island
kurı̄ played an important role in Māori hunting. However, kurı̄ are also
well represented in the subtropical north, albeit usually in reduced
numbers relative to South Island assemblages (Clark, 1995). They were
particularly abundant at the 14th century village of Houhora (Fig. 2),
where moa hunting was an important activity, but dog consumption is
also suggested (Allo Bay-Petersen, 1983; Furey, 2002; Shawcross, 1972).
Sometime after AD 1450 the situation begins to change. Between AD
1450 and 1650, kurı̄ are poorly represented everywhere (Fig. 2). This
follows the extinction of all species of moa, the loss of other bird re­
sources, and the collapse of seal rookeries—in short, the demise of large
game hunting (Anderson, 2016; Holdaway and Jacomb, 2000; Smith
and James-Lee, 2010; Smith, 2013; Worthy and Holdaway, 2002). This
decline in dogs appears to reflect a situation of dog-human competition
as subsistence economies were reorganised. We suggest the possibility of
a genetic bottle neck during the mid-sequence decline in Aotearoa based
on the spatial distribution of kurı̄ in archaeological contexts. The pos­
sibility that the decline in kurı̄ during the Middle Period pattern is an
artefact of archaeological sampling also cannot be altogether discounted
on current evidence.
Perhaps the most unexpected pattern is the marked increase of dogs
after AD 1650, with kurı̄ again well represented in both North and South
Island locations, in some cases rivalling abundances of the Early Period
(e.g., the Kohika lakeside settlement). This late pattern is consistent with
observations of the earliest European visitors who describe instances of
kurı̄ accompanying Māori in varied contexts. At this time South Island
Māori continued to undertake a range of foraging activities, with long-
distance trade and exchange playing crucial roles in the management
and distribution of critical food resources (Anderson and Smith 1996). In
the north, kurı̄ are well represented as well, often in the context of
relatively sedentary communities, semi-anchored by sweet potato
cultivation. These agricultural pursuits were typically combined with
intensive, often seasonal, marine, and freshwater harvesting activities,
as well as birding, where kurı̄ may have played a role. Although Māori
themselves were increasingly navigating inter-tribal rivalries, territorial
intrusions, and outright warfare (Allen, 1996; Anderson, 2016; McCoy
and Ladefoged, 2019), especially in the agricultural north, kurı̄
remained abundant. Notably, they were also valued as raw materials for
prestigious cloaks (skins and hair) and as a ceremonial food (Colenso
1877: 151), but the relative importance of these differing uses (e.g.,
McKechnie et al., 2020) have not been systematically investigated in the
Aotearoa setting.
To better understand these distribution and abundance patterns, and
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P. Pillay et al. Journal of Archaeological Science 139 (2022) 105556

Table 3 Table 4
Kurı̄ remains used in dental analysis by site and preservation conditions. Age categories and associated cranial and mandibular features.a
Time Site name Approx. Assemblage NISP Assemblage Relative Age Distinctive Features in the Distinctive Features in the
period Site area composition preservation Cranium Mandible
(m2)
Juvenile (less 1. Deciduous teeth (if 1. Deciduous teeth present (if
Early Hot Water 26.5 Crania 9 Variable, than 7 permanent dentition then use mixed dentition then use
Beach Mandibles 4 fragmented months) further criteria). further criteria).
Loose 6 2. Cranial sutures are irregular, 2. Condyloid, angular and
Incisors 2 and edges feathered; coronoid processes show no
Loose 7 mineralisation of bone indication of muscle
Canines 7 incomplete making bone attachment.
Loose porous. 3. Overall size in relation to
Premolars 3. Weakly developed other individuals in
Loose Molars zygomatic processes and assemblage. Specimens
Whangamata 9.24a Crania 6 Good narrow may vary geographically.
Mandibles 9 condition frontal bone.
Loose 1 4. Overall size in relation to
Incisors 3 other individuals in
Loose 1 assemblage. Specimens may
Canines 1 vary
Loose geographically.
Premolars Sub-Adult 1. Permanent dentition only, 1 Permanent dentition only.
Loose Molars (7–18 sagittal crest and nuchal crests 2. Condyloid, angular and
Houhora 245 Crania 15 Variable, months) not pronounced coronoid processes show some
Mandibles 29 weathered, 2. Feathered edges still present signs of muscle attachments.
Loose 24 fragmented in places, sutures not 3.Overall size in relation to
Incisors 13 completely fused, but bone other individuals in
Loose 26 mineralisation is almost assemblage. Specimens may
Canines 26 complete vary geographically.
Loose 3. Zygomatic processes of the
Premolars frontal bone not sharp and
Loose Molars pointed, but still blunt.
Te Mataku 24 Maxilla 9 Weathered, 4.Overall size in relative to
Mandibles 14 fragmented other individuals in
Middle Long Bay 6000 Maxilla 4 Weathered, assemblage. Size may vary
Mandibles 3 fragmented geographically.
Late Airport NRD 2,225b Crania 5 Variable, Adult 1. Permanent dentition. 1. Permanent dentition.
Mandibles 32 weathered, 2. Cranial sutures have fused in 2. Muscle markings are
Loose 9 fragmented places, but suture lines are still observable on the posterior
Incisors 13 visible. 3. Prominent sagittal processes of the mandible and
Loose 9 and dorsal nuchal crests; ascending ramus.
Canines 11 overall 3. Mineralisation of bone is
Loose size of cranium. complete, robust, and no
Premolars 4. The parietal and temporal longer porous.
Loose Molars bones have robust muscle 4. Overall size in relation to
Kohika 220 Crania 26 Good attachment markings. other individuals in
Mandibles 75 condition 5. Overall size in relation to assemblage. Specimens may
Loose 1 other individuals in vary geographically.
Incisors 1 assemblage. Specimens may
Loose 8 vary geographically.
Canines Mature Adult 1. All elements of adults plus 1.All elements of adults plus
Loose Molars cranial sutures fused stage 9 or stage 10 tooth wear.
Oruarangi 20,000c Crania 4 Good completely. Note: this category requires
Mandibles 40 condition 2.Very pronounced and robust crania and post cranial
Loose 6 sagittal and nuchal crests. elements for confident age
Canines 1 3. Overall size in relation to assignments and tooth wear
Loose Molars other individuals in can relate to diet.
a assemblage. Specimens may 2. Overall size in relative to
This estimate represents only the Wharf Site excavations (Jolly 1978:136). vary geographically. other individuals in
b
Approximate site area calculations for the Airport NRD Area A and Area B assemblage. Size may vary
excavations. geographically.
c
This estimate represents the entirety of the Oruarangi excavations. a
Modified from Clark (1995:62–65). The sub-adult and mature adult cate­
gories were devised specifically for this study. Developmental features are listed
in particular the possibility of interspecific competition, we analysed from most to least accurate.
kurı̄ dentition for diet-related markers and developmental defects
relating to nutritional and physiological stressors from eight north island assemblages are dominated by younger rather than older individuals
archaeological sites with available kurı̄ dentitions. We anticipated that: and this does not vary by time period. A Pearson’s Chi-Square test in­
a) top-down processes (human population growth, climate change) dicates no statistically significant differences in demographic profiles
would result in a coherent regional pattern and a clear temporal trend; between early and late sites (X2 = 6.1643, df = 4, Exact sig. = 0.1872)
or b) if these factors were inconsequential, then variability might be supported by the Fisher’s exact test (X2 = 6.023, df = 4, Exact sig. =
localised. 0.164).
For the eight North Island assemblages used in the dental analysis,
most individuals could be assigned to a relative age category. Sub-adults
and adults were the most common groups; very old individuals were 4.2. Caries, calculus, periodontal disease and tooth wear
uncommon with only one mature adult in the Kohika assemblage. The
No caries or calculus was observed in the analysed kurı̄ assemblages.

6
P. Pillay et al.
Table 5
Tooth wear scoring system used in this study based on Lemoine et al. (2014) but
modified for kurı̄ dentition based on wear observations made by Allo (1971) and
Clark (1995).
Description
Score
0 Unerupted.
1 Tooth visible in crypt. Crown beginning to mineralise.
2 Mineralisation of crown complete.
3 Roots forming.
4 Roots completed and tooth erupted but no visible wear.
5 Little enamel wear. Some abrasion on the inner surfaces
of carnassial teeth; incisors have retained the “Fleur-de-
lys” shape and are slightly worn or mostly unworn;
canines showing signs of abrasion on the crown.
6 Cusps on carnassial teeth (first and second molars)
showing signs of wear; “fleur-de-lys” shape of incisor
begins to disappear. The buccal and lingual cusps on P4
show pinprick signs of wear, while wear centres in the
molars have started showing signs of wear on the major
cusps.
7 Carnassials are very worn with partial disappearance of
lingual cusps on upper first molar; canines display
further signs of wear, loosing shape of crown; “fleur-de-
lys’ shape of incisors has disappeared.
8 Carnassial cusp wear has expanded to the extent that
there is a continuous band of exposed dentine along the
length of the tooth. Wear centres of the upper M1 and
M2 continue to expand until wear on the buccal cups
have joined and lingual cusps have joined respectively.
Posterior cusps in lower carnassial are worn flat.
Extensive wear on crowns of incisors and premolars.
Canines crown shape now reduced in places.
9 Incisors worn almost flat to original gum surface; wear
on canines gives spatulate flat top crown. The molars
worn to a smooth concave surface; upper M1 and M2
buccal and lingual cusps have coalesced. Large anterior
cusps of premolars completely worn away; pulp cavities
occasionally visible.
10 Pulp cavity is now visible in canines and carnassial teeth;
P4 showing exposed dentin and enamel completely worn
away and surface almost smooth. All teeth have lost
original shape. Destruction of the upper M1 and M2
cusps are complete. Tooth roots in canine and carnassial
teeth exposed.
With respect to periodontal disease, only one individual from Kohika
displayed early signs of possible periodontal disease. No cases of
abscessing were observed in any of the assemblages. These results sug­
gest that kurı̄ oral health was relatively high and remained so
throughout the Aotearoa sequence.
Only permanent teeth were used for the tooth wear analysis given a
lack of wear on deciduous teeth. Light to moderate wear (stages 4 and 5)
was typically observed on the molars and upper fourth premolar teeth
across all sites (Fig. 3). There was less wear observed on teeth from Early
Period sites relative to the Late Period (Table 6). This was confirmed by
the non-parametric Levene’s Test, which showed that variances in wear
scores were statistically significant between early sites (p = 0.000) and
between late sites (p = 0.000). Therefore, differences in the sample
variances were unlikely to be a result of random sampling from a pop­
ulation of equal variances. The Kruskal-Wallis test demonstrated sta­
tistically significant differences in wear between early and late time
periods held, even when the Kohika sample was excluded (Kruskal-
Wallis H = 602.45, df = 9, p-value < 2.2e-16). Within the late period no
differences were found between the wear observed between the Airport
NRD and Oruarangi assemblages (Kruskal-Wallis chi-squared = 7.9581,
df = 6, p-value = 0.2412). However, when age groups were considered,
a temporal change in wear scores was observed between samples
(Table 7 and Table 8). In short, a temporal change is indicated, although
it is less marked when the Kohika sample is excluded.
For the Kohika sample, the lack of difference in wear by age
Journal of Archaeological Science 139 (2022) 105556
demonstrates rapid rates of wear earlier in life (i.e., juveniles and sub-
adults) in comparison to elsewhere. At Kohika, tooth wear was the
most severe, with the average wear score for individuals being stage 7.
In this case, tooth wear was observed on incisors, canines, upper and
Wear Type lower molar teeth, and the maxillary fourth premolar. This holds when
controlling for age (Table 8), albeit the considerable variation observed
Pre-Wear between sites from the same time period appears to be driven by dietary
formation abrasiveness. The Kohika specimens were the most distinctive of all
assemblages analysed, with extremely high tooth wear scores even in
Pre-Wear
juvenile remains (stages 7 and higher), suggesting quite different dietary
eruption or husbandry conditions.
Light wear
4.3. Developmental defects
Moderate wear Hypoplastic lesions were observed mostly on permanent teeth, with
only one case on a deciduous premolar (Fig. 4, Table 9). Both pits and
lines were frequently observed on maxillary dentition. All incidences of
hypoplasia occupied less than one-third of the overall tooth. The per­
manent canines had the highest incidence of hypoplasia (n = 4). No
Heavy wear hypoplastic defects were observed on incisor teeth but permanent pre­
molars and the mandibular first molar had hypoplastic defects, sug­
gesting that hypoplasia is more likely associated with specific tooth
types (Table 9). This was expected as these teeth incorporate the longest
period of development (Arnall, 1960).
While the overall occurrence of hypoplastic defects was low across
all assemblages (Table 9), defect frequency varied across the three time
periods (Table 10). Out of the total MNI identified in each assemblage, at
least 11.7% from Early Period sites had hypoplastic defects and 7% from
Late Period sites (Table 10). No defects were observed in the Middle
Extreme wear Period.
The timing of hypoplastic defects tended to correlate with the onset
of weaning at approximately three to four months (Fig. 5). This pattern
reflects the transition from suckling to self-feeding, rather than hus­
bandry practices. The timing of defects can also inform on the health of
the mothers of these individuals. However, no hypoplastic defects were
observed on teeth that developed in utero and only two defects were
observed prior to three months of age, both suggesting good maternal
health.
5. Discussion
5.1. Aotearoa-wide distribution and abundance over time
The islands of Aotearoa constitute a considerable geographic area
and a significant contrast to the smaller land masses of central East
Polynesia. Extending the length of both the North and South Islands, and
onto several of the larger offshore islands, the early dispersal of dogs in
Aotearoa (AD 1250–1450) speaks to their remarkable behavioural and
physiological plasticity, particularly their ability to rapidly adapt to a
diversity of environments and novel food sources. Their distributions
also reflect the intensity, resiliency, and success of their commensal
relationship with their human managers, who not only translocated
them to Aotearoa, but carried them between the many islands within
this archipelago. Overall, it would seem that the Aotearoa environment,
in combination with the adaptability of Polynesian dogs, allowed for
their successful dispersal and rapid increase, with little to no competi­
tion with their Māori managers.
These early outcomes contrast markedly with observations from the
middle of the Māori sequence (AD 1450–1650). During this interval kurı̄
occur in a small number of sites and in reduced numbers. They essen­
tially disappear from all areas other than the Auckland region of the
North Island and the Otago region of the South Island (see also Allo
Bay-Petersen 1983: 126). As discussed earlier, the socio-economic
re-orientations of this time interval were considerable. As Little Ice
Age conditions intensified, many Māori populations shifted northward
from the South Island, and some southern areas of the North Island were
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P. Pillay et al. Journal of Archaeological Science 139 (2022) 105556

Fig. 2. Distribution of kurı̄ (MNI) in Māori archaeological assemblages across three time periods (see text for details).

Fig. 3. Examples of tooth wear. Specimen AIM 19595 from Oruarangi (left), courtesy of Auckland War Memorial Museum Archaeology collections, had very light to
mild enamel wear and an average wear score of 5.13. In contrast, individuals from Kohika exhibited severe wear; Specimen 1B-0070 (right) illustrates completely
worn cusps and was assigned an average wear score of 9. (Photograph by Tim Mackrell).

abandoned as conditions became marginal for traditional gardening. A
recent analysis of Māori population dynamics, based on 14C summed
probability distributions, identifies a “flat” growth period in the central
region of the North Island during this Middle Period (Brown and Crema,
2019). The authors suggest “some form of carrying capacity was met,
which may have elicited a similarly competitive social arena and the
development of pā [fortified sites]” (Brown and Crema 2019:16). For­
tified sites do indeed appear from around the 16th century, particularly
in areas suitable for cultivation. Today there are more than 7000
archaeologically recorded fortified sites, some with long histories and
multiple periods of use, and all post-16th century AD in age. As a whole,
they highlight the intensity of inter-tribal competition from the 16th
century onward (Anderson, 2016). The poor representation of kurı̄
across the country in this time interval could well be signalling inter­
specific competition (between Māori and kurı̄).
Perhaps the most unexpected finding from the distributional analysis
was the apparent increases of kurı̄ in the Late Period (AD 1650–1800).
Although the geographic extent of kurı̄ observed in the Early Period is

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P. Pillay et al. Journal of Archaeological Science 139 (2022) 105556

Table 6 Table 8
Dental wear scores by site and time period based on the 10-point scale (see text). Kruskal-Wallis statistical dental wear scores for kurı̄ age group by time period.
Only wear scores have been included in these calculations and only permanent Table also includes degrees of freedom (df).
teeth are included, as all deciduous teeth had no observable tooth wear. Group Variables Kruskal- N df Sig.
Site NISP MNI Mean Wear Score Std. Deviation Wallis H

EARLY PERIOD Comparisons with

Hot Water Beach 40 6 4.92 1.05 Kohika sample
Whangamata 28 5 4.82 0.98 Between Early and Late
Houhora 111 40 5.51 1.16 Periods
Te Mataku 38 9 5.34 1.19 All Age 34.832 333 6 <0.001
TOTAL EARLY PERIOD 217 60 5.29 1.15 Groups
MIDDLE PERIOD Adults 46.985 104 6 <0.001
Long Bay 10 4 4.8 0.789 Sub-Adults 23.858 144 6 0.001
LATE PERIOD Adults vs Sub- 36.286 181 6 <0.001
Airport 76 12 5.24 1.78 Adults
Kohika 170 36 7.01 1.63 Juveniles 9.0267 25 6 0.721
Oruarangi 63 23 5.13 1.41 Early Period only
TOTAL LATE PERIOD 319 71 5.79 1.86 All Age 10.546 77 4 0.032
Groups
Adults 4.9614 35 4 0.291
Sub-Adults 5.7361 15 3 0.125
Table 7 Adults vs Sub- 6.7714 64 4 0.149
Wear scores by age categories from the eight North Island archaeological sites. Adults
Note that MNI refers to specimens that could be aged not total MNI. Juveniles 1.4 8 1 0.237
Late Period only
Site Age MNI N Mean wear Std. All Age 60.988 253 6 <0.001
(teeth) score deviation Groups
Hot Water Adults 4.8814 69 4 0.300
Beach Sub-Adults 5.7766 129 6 0.449
Adult 5 5 5.2 0.98 Adults vs Sub- 56.71 225 6 <0.001
Sub-Adult 1 3 5.3 1.25 Adults
Juvenile 0 – – – Comparisons excluding
Whangamata Kohika sample
Adult 1 3 5 0.82 Between Early and Late
Sub-Adult 1 – – – Periods
Juvenile 2 5 5.2 0.98 All Age 11.298 185 6 0.080
Houhora Groups
Adult 10 13 6 1.11 Adults 12.239 43 4 0.016
Sub-Adult 9 10 6.1 0.83 Sub-Adults 14.309 47 6 0.026
Juvenile 11 3 6 – Adults vs Sub- 11.035 89 6 0.087
Te Mataku Adults
Adult 9 14 5.43 0.73 Late Period only
Sub-Adult 0 – – – All Age 11.927 98 6 0.064
Juvenile 0 – – – Groups
Long Bay Sub-Adults 7.2711 92 6 0.297
Adult 0 – – – Adults vs Sub- 56.71 225 6 <0.001
Sub-Adult 2 3 5 0.75 Adults
Juvenile 1 – – – Kohika sample only
Airport NRD – – – All Age 9.7566 42 6 0.135
Adult 0 – – – Groups
Sub-Adult 2 15 6.13 2.33 Adults vs Sub- 9.4345 34 5 0.093
Juvenile 3 – – – Adults
Kohika
Mature 1 9 9 0.58
Adult and our assignment of the Masonic Lodge assemblage to the Early Period
Adult 29 64 7.25 1.19 based on 14C data, the presence of moa and other typically early fauna,
Sub-Adult 3 23 7.26 1.73 and early style artefacts (see Wood et al. 2016: 503). Nonetheless, dogs
Juvenile 3 20 7.3 1.18
were clearly important in the late part of the traditional Māori sequence
Oruarangi
Adult 3 5 7.4 0.49 and their uses may have changed. Dogs continued to be a source of food
Sub-Adult 5 20 5.85 1.31 for Māori but may have become largely restricted to elites and occasions
Juvenile 1 – – – of feasting. Also, at western contact dog-skin cloaks were highly valued,
typically reserved for elites, sometimes named, and oftentimes were
passed from one generation to the next and further analysis might reveal
not achieved, the number of sites with dogs and dog abundances within
associated changes in kurı̄ management.
these sites suggest a population rebound. In this respect our review of
kurı̄ occurrences from published accounts and the Heritage New Zealand
Archaeological Reports Digital Library differ to the recent review of 5.2. Dental conditions and diet
Greig and Walter (2021), published while our manuscript was in revi­
sion. They argue that although the cited studies “seem to show dog To better understand kurı̄ temporal patterns, and assess dog diet and
abundance levels decreasing with time, the evidence for a decline is health more directly, our dental analysis focused on assemblages from
equivocal” and they attribute variability in dog abundances to differing the populous northern region of the North Island. This was an area
site functions. Although we are not in a position to systematic evaluate where traditional agricultural practices intensified over time (Anderson
site function here, a cross-section of habitation sites is represented in our and Petchey, 2020; Prebble et al., 2019), and where fortified sites were
data sets. Our observations may also differ from theirs because of our concentrated after AD 1650. With one exception, the analysed assem­
wider geographic coverage, a somewhat larger number of assemblages, blages indicated a high level of dental health and only modest dietary

9
P. Pillay et al. Journal of Archaeological Science 139 (2022) 105556

Fig. 4. Examples of LEH and pit type defects: (A)

Arrows point to pit type defects observed on the mid-
crown region of deciduous fourth premolar from the
Airport NRD site; (B) Arrow points to linear enamel
hypoplasia defect on maxillary canine from specimen
1B0016 from Kohika assemblage; (C) Arrow points to
pit type defect on maxillary canine from HWB
(specimen AIM-AR3516 courtesy of Auckland War
Memorial Museum Archaeology collections). (Photo­
graphs by Tim Mackrell).

evidence a near absence of dental caries, calculus, and periodontal dis­

Table 9
ease. With the exception of the Kohika dogs, where wear was severe
Percentage (%) of teeth with hypoplastic defects by tooth type and defect type,
even in young animals, tooth wear was moderate. However, slightly
linear enamel hypoplasia (LEH) or pits; number of crowns with defects in pa­
rentheses. Note that crowns with both LEH and pit type defects present were
greater wear was documented in the Late Period dogs, even when the
counted once. anomalous Kohika assemblage was excluded, suggesting some dietary
change (Pillay, 2020).
N % of all hypoplasia % of LEH % of pits
Overall, our findings are consistent with those of (Allo Bay-Petersen,
Permanent 1983: 39), who identified calculus in low frequencies and at only two
Incisor 74 0 0 0
sites: Wairau Bar on the South Island and Houhora on the North
Canine 50 1.34 (7) 0.96 (5) 0.96 (5)
Premolar 92 0.76 (4) 0.38 (2) 0.38 (2) Island—localities where large samples were available. Similarly peri­
Molar 307 0.19 (1) 0.19 (1) 0 odontal disease was infrequent in her study and only observed in South
Total 523 2.29 (12) 1.53 (8) 1.34 (7) Island assemblages where it was uncommon and ranged from consid­
Deciduous erable (in two Early Period individuals) to slight (in five Late Period
Incisor 0 0 0 0
Canine 1 0 0 0
individuals) (Allo 1971:37–9). Extreme wear was only observed in
Premolar 12 0 0 7.69 (1) southern kurı̄ assemblages as well, a pattern Allo attributed to harder
Total 13 7.69 (1) 0 7.69 (1) diets.
The lack of caries in the Aotearoa assemblages is notable given that
Combined Totals 536 2.43 (13) 1.53 (8) 1.49 (8)
this condition was common in Hawaiian dogs, where it was attributed to
starchy carbohydrates and hand-feeding practices. Our initial inference
was that Aotearoa kurı̄ lacked caries because they had limited access to
Table 10 sweet potato (kumara, Ipomoea batatas), the most widely cultivated
MNI with hypoplasia by time period. starchy carbohydrate of Māori. However, when considered alongside the
Early Middle Late moderate wear recorded in the present study, we cannot exclude the
Total MNI 60 4 71
possibility that abrasives in the diet discouraged both conditions
MNI with hypoplasia 7 0 5 through “caries-attrition competition” (Maat and Van der Velde, 1987).
% of individuals with hypoplasia 11.67 0 7.04 Consumption of “fern root” (rhizomes of bracken fern, Pteridium escu­
lenta), an important wild resource that Māori processed into a fibrous
food, could also have reduced the build-up of dental calculus in kurı̄. In
change across the three time periods. More specifically, the results

10
P. Pillay et al. Journal of Archaeological Science 139 (2022) 105556

Fig. 5. Summary of the location and timing of hypoplastic defects in North Island kurı̄ tooth assemblages. Top section: Timing of dental formation in relevant
molariform crowns of modern domestic dogs, based on Arnall (1960), Silver (1969), and Williams and Evans (1978). Bottom section: archaeological individuals with
hypoplastic defects shown by tooth type and time period.

Table 11
Summary of published kurı̄ coprolite results from North Island archaeological sites. All identified materials are macrofossils unless otherwise indicated. Table updated
from Clark (1995:18).
Archaeological Site Period N analysed Contents (N of samples) Method Source

Cabana Lodgea Early unspecified sweet potato dry Gumbley et al. (2014)
taro
Houhorab Early 6 fish bone (6) wet Byrne (1973):24 in Clark (1995):18
fish scales (3)
fish teeth (5)
charcoal (5)
unid. plant (3)
fiber (3)
Hot Water Beachb Early 4 fish bone (4) wet Byrne (1973):24 in Clark (1995):18
fish scales (1)
fish teeth (3)
Masonic Tavernc Early 5 fish bone wet Wood et al. (2016)
marine shell
Macropiper sp.
Carex sp. (sedge)
Juncus sp. (rush)
Potamogeton sp. gourd
paper mulberry Plus several other taxa
Sarah’s Gully Early unspecified “triturated bone” dry Scarlett (1972):23 in Clark (1995):18
Sunde site Early 4 fish bone (4) wet Byrne (1973):24 in Clark (1995):18
fish scales (4)
fish teeth (4)
charcoal (4)
spores (2)
Long Bayb Middle 57 fish bone dry Campbell et al. (2019)
Kohikab,c Late 25 (out of 300) fish bone wet Horrocks et al. (2003); Irwin et al. (2004)
charcoal
marine shell
Cyperaceae
Solanum sp.
Unid plant parasites
Oruarangib Late 2 mixed bone dry Best (1980):78 in Clark (1995):18
marine shell
a
Based on microfossil analyses carried out by M. Horrocks. See Appendix Four in Gumbley et al. (2014).
b
Archaeological sites containing kurı̄ teeth assemblages that were used in present study.
c
Partial list of taxa represented by macroscopic materials, pollen, and aDNA analyses.

11
P. Pillay et al.
support of these suggestions, calculus is also uncommon in Māori skel­
etal assemblages (Aranui, 2018; Campbell et al., 2019).
Further insights are provided by kurı̄ coprolite studies. These are
available for nine northern Aotearoa sites, including five that provided
dentition for the current study (Table 11). At the early 14th century site
of Cabana Lodge (T12/3), adjacent to our Whangamata Wharf (T12/2)
site, microfossil evidence indicates that dogs consumed sweet potato and
taro (Colocasia esculenta) (James-Lee and Gumbley, 2012)—the latter a
cold-intolerant tropical root crop that was valued but not widely culti­
vated. aDNA analysis of kurı̄ coprolites from the Masonic Lodge pro­
duced evidence of gourd (Lagenaria siceraria), paper mulberry
(Broussonetia papyrifera), and possibly yam (Dioscorea alata) (all culti­
vated species), along with several wild plants including bracken fern
(Wood et al., 2016). Although paper mulberry was not a Māori food, it
was cultivated for the production of bark-cloth, and thus highly valued.
These two Early Period coprolite assemblages demonstrate that kurı̄ at
least occasionally fed on culturally important plant resources, which
might indicate such food resources were readily available.
More commonly, fish bones, scales, and teeth are well represented in
kurı̄ coprolites, and from all time periods. This is consistent with early
historical accounts, which frequently mention that dogs were fed fish (e.
g., Forster, 1778: 189). Dogs will often self-forage for fish as well (per­
sonal observations). The handful of published stable isotopic analyses
are supportive of the foregoing, with marine dominated diets identified
at Te Mataku (Pillay 2020; see also SI) and Long Bay (Campbell et al.,
2019: 29), and mixed marine and terrestrial diets indicated at Wairau
Bar (Kinaston et al., 2013). Other coprolite components help explain the
moderate tooth wear patterns observed in our assemblages. The extreme
wear observed on the Kohika dog teeth, along with considerable tooth
fracture (Pillay, 2020), may relate to the consumption of shellfish, along
with ingestion of sandy sediments and possibly silica-rich grasses
(Horrocks et al., 2003).
These coprolite results demonstrate is that kurı̄ and human diets
overlapped but only partially. Results from Masonic Lodge and Kohika
suggest that dogs were probably self-foraging and possibly free roaming,
as indicated by evidence for numerous wild plants, grit, and charcoal.
These findings are consistent with suggestions of Greig and Walter
(2021), but this style of husbandry may have varied by location.
Free-roaming practices, for example, might be impractical when dogs
were raised specifically for feasting or for their skins.
Only a small number of dog dentition studies are available from
elsewhere in East Polynesia for comparison. A sizable sample (c. 30
MNI) of dogs from Hawaiian archaeological contexts contrast with the
Aotearoa results. Svihla (1957) found high rates of caries in the Ha­
waiian dogs (c. 30%), which he attributed to diets rich in starch and
sugars, and possibly the consumption of taro (Colocasia esculenta) and
breadfruit (Artocarpus altilis), two traditional Hawaiian staples. On the
small almost-atoll of Aitutaki (18 km2) (southern Cook Islands), both
caries (n = 1) and calculus (n = 1) were observed despite small sample
sizes spread out across three sites (MNI = 9) (Miller, 2013). In contrast,
on the large island of Nuku Hiva (341 km2) (Marquesas Islands), where
historic accounts (Ferdon, 1993) and wood charcoal studies (Huebert
and Allen, 2020) indicate breadfruit was a readily available staple, both
caries and calculus were lacking (MNI = 6). These results highlight both
regional variability in the incidence of caries and calculus amongst
Polynesian dogs and the challenges of interpreting the differential
occurrence/persistence of the latter. Starchy foods may promote dental
calculus, but other dietary components may remove them. Hand-feeding
(suggested in the Hawaiian case) or restraints on mobility such as
tethering, and enclosures may also be important in promoting or pre­
venting these conditions.
While the foregoing evidence identifies some occasions of food
sharing, stable isotope analyses can assist in evaluating dietary overlap
between dogs and their human managers over longer time intervals. At
the early Wairau Bar site (South Island) (Kinaston et al., 2013) kurı̄ diets
are most like those of from well ornamented and possibly elite human
12
Journal of Archaeological Science 139 (2022) 105556
burials but differ from most of the human individuals in having less
variability in δ13C and lower δ15N values. On Tahuata (Marquesas
Islands) dog and human diets overlap little, with dogs consuming more
marine foods, particularly early in time, but also feeding at lower trophic
levels relative to their human companions and showing considerable
variability in their δ13C values (Richards et al., 2009). On the very small
island of Aitutaki (Cook Islands), stable isotope analysis demonstrates
partial dietary overlap, with dogs having less negative δ13C values
relative to humans, suggesting less access to cultivated carbohydrates,
and higher δ15N values relative to those of humans, which may reflect
feeding off remains of large, carnivorous fish such as those identified in
human occupation sites (Allen and Craig, 2009; Craig, 2009).
5.3. Dental indicators of physiological and/or nutritional stress
A more direct indicator of physiological and nutritional stresses, in
both dog and human teeth, is enamel hypoplasia. In the sizable northern
Aotearoa dog assemblages, it occurs in low frequencies, being found at
one early site (Hot Water Beach, MNI = 6) and two late sites (Kohika,
MNI = 36; Airport NRD, MNI = 12), in frequencies of 11.7 and 7%
respectively. Although these temporal differences are not statistically
significant, a consideration of teeth by age groups shows the incidences
of enamel hypoplasia were concentrated around the timing of weaning.
Notably, weaning is widely recognised as a period when the risk of
malnutrition is high in mammals. Enamel hypoplasia was also recorded
in the Marquesan dog assemblage, but the sample size was small (MNI =
6) and hypoplastic pits were only observed on two teeth that were
probably from the same mature individual (Littleton et al., 2015: 290).
In sum, the Aotearoa dog dentition analysis provides little indication of
physiological or nutritional stressors, outside of those associated with
weaning—suggesting little competition with humans.
6. Conclusions
Recent archaeological studies have focused on mutualistic outcomes
of the human-dog symbiosis. Here we have considered another dimen­
sion of these relations—the potential for interspecific competition and
attendant effects on dog diets, health, and husbandry practices. Our
results demonstrate the conundrum that dogs presented to Polynesian
communities of the past. Their translocation across vast distances under
the challenging conditions of open ocean voyaging indicate the high
value placed on this domestic by oceanic colonists. Nevertheless,
although they were successfully established on many islands, their oc­
casional decline and extirpation are notable (Cramb, 2021). Ethnohis­
torical and archaeological evidence suggests that dogs could be costly to
maintain, especially when opportunities for dog self-foraging were
limited, as on small islands, and interspecific competition occasionally
emerged. However, previous attempts to explore these relationships
have been challenging as dog remains are scarce in many island settings
(Allo Bay-Petersen, 1983; Littleton et al., 2015; Miller, 2017).
Here we have leveraged the large Aotearoa New Zealand dog as­
semblages to better understand both local and wider Pacific Island
patterns. Country-wide distributional evidence was paired with a
regionally focused analysis of dental indicators of diet and health
(caries, calculus, periodontal disease, tooth wear, and enamel hypo­
plasia). Working from the premise that dog and human diets would not
necessarily overlap, we explore how they may have aligned or diverged
across different geographic settings and over time. Evidence from the
early settlement period (AD 1250–1450) is consistent with competitive
release on arrival in Aotearoa. Dogs were quickly dispersed across a
diversity of environmental and climatic gradients, on the two main
Aotearoa islands and on numerous offshore islands.
Subsequent declines in dog numbers, and contractions in their
regional distributions, suggest a phase of dog-human competition be­
tween AD 1450 and 1650, a period of significant economic reorientation
and population mobility on the part of Māori populations. However,
P. Pillay et al.
archaeological sampling biases could play a role in this patterning and
our solitary northern assemblage from the Middle Period is unremark­
able in terms of diagnostic dental features. In the Late Period (AD
1650–1800) dogs are once again well represented at individual sites, but
less widely distributed relative to the Early Period, with dental markers
in this period indistinguishable from those observed in earlier samples,
with one exception.
So how do these findings relate to our two initial hypotheses? A
reasonable case can be made that Polynesian dogs underwent compet­
itive release on arrival in Aotearoa. With respect to the proposal that
competition arose in the centuries after initial settlement, the distribu­
tional study identified the Middle Period as a time of possible inter­
specific competition but dental assemblages that might have permitted
further assessment were lacking. Hypothesis two, of increasing compe­
tition over time, only partially holds based on dog distributions, abun­
dances, and even kurı̄ dental assemblages from the region of most
intense intertribal rivalries. Rather, evidence from the Late Period sug­
gests dog populations rebounded and at most some modest dietary
change may have ensued based on moderate increases in dog dental
wear.
Among the more interesting findings from our study is demonstra­
tion of regional variability in dog dental markers across East Polynesia.
Caries, while rare in contemporary dogs, was well represented in the
Hawaiian assemblages. Calculus, a common problem in modern dogs,
was lacking in those of pre-western Aotearoa. Comparing the northern
North Island assemblages with those previously analysed from South
Island localities demonstrated regional variation in periodontal disease
in the New Zealand context. These findings suggest we might profitably
consider this measure in future dog dentition studies elsewhere in the
Pacific Islands. Enamel hypoplasia was rare in the large Aotearoa as­
semblages, where it was unambiguously linked to weaning, a recognised
life history event, rather than more generalised physiological or nutri­
tional stressors that might indicate poor health. Perhaps most impor­
tantly, the observations in these different island settings speak to
variability in human-dog relationships, husbandry practices, and the
place of dogs within the anthropogenic niche—relationships we are only
beginning to understand.
There are several ways the present analyses might be extended to
answer outstanding questions. Clearly additional samples from the
Middle Period would be useful. Additional stable isotopic and coprolite
studies would also be beneficial, allowing for more nuanced un­
derstandings of human-dog interactions over the roughly 600-year
sequence (e.g., Witt et al., 2021). Stable isotope analyses would not
only inform on major dietary components, but also provide a firmer
basis for evaluating dietary overlap between kurı̄ and their managers,
and changes that perhaps accompanied shifts from dog use as hunting
aids to status foods and use of their skins for status regalia. Other
questions could potentially be addressed through genetic study. Nuclear
DNA markers in particular (Greig et al., 2016, 2018) might prove useful
in assessing the possibility of a mid-sequence kurı̄ population decline
and/or the purposeful selection for new traits later in time, as for
example those related to coat colour or texture (e.g., McKechnie et al.,
2020).
Our findings contribute to emerging global discussions regarding the
place of one of humanity’s oldest and perhaps “best friends”. Pacific
Island dog assemblages underscore the dynamic nature of the human-
dog symbiosis, illustrating how these domestics may be highly valued
by, and tightly integrated with, human communities, yet sometimes
were pushed to the margins as their human companions navigated the
challenges of island living, both large and small.
Funding
MSA received support funding from Te Pūnaha Matatini (https
://www.tepunahamatatini.ac.nz) Grant no. UOA 9167-3705716.
13
Journal of Archaeological Science 139 (2022) 105556
Credit author statement
Patricia Pillay: Investigation and analysis, conceptualisation, orig­
inal draft preparation and writing, statistical analysis. Melinda Allen:
MA thesis supervision, conceptualisation, writing, visualisation. Judith
Littleton: MA thesis supervision, conceptualisation, dental analysis
methodology, review, and editing.
Declaration of competing interest
None.
Acknowledgements
The analysis of dental markers in northern Aotearoa New Zealand of
kurı̄ was undertaken by Patricia Pillay for her MA thesis supported by
the University of Auckland Kupe Leadership Research Masters Scholar­
ship and Royal Society Te Apārangi 2018 Skinner Fund. We thank Ngāti
Awa and Geoffrey Irwin for permissions to use the Kohika assemblages.
We also thank Louise Furey (Curator), Emma Ash (Associate Curator)
and Deirdre Harrison (Collections Manager) for assistance with collec­
tion loans from The Auckland War Memorial Museum Tāmaki Paenga
Hira. The support of Ngāti Hei and the co-directors of the Ahuahu Great
Mercury Island Archaeological Project (Rebecca Phillipps, Louise Furey,
Simon Holdaway, Thegn Ladefoged), along with support staff Alex
Jorgensen and Joshua Emmitt, is also much appreciated. We thank
Matthew Campbell for access to the Long Bay and NRD assemblages.We
are grateful to Ngāti Manuhiri, Ngāti Whātua o Ōrākei, Ngāti Whātua o
Kaipara, Ngāti Maru, Ngāi Tai ki Tāmaki, Te Kawerau ā Maki, and Ngāti
Pāoa for permissions to access the Long Bay materials. We express
gratitude to Te Ākitai Waiohua for supporting the Airport NRD analysis.
A special thanks to Seline McNamee for preparation of the distribution
map and illustration expertise, and to Timothy Mackrell for assistance
with the photographs. We also thank two anonymous reviewers for their
feedback.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.jas.2022.105556.
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