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Journal of Archaeological Science 56 (2015) 168e176

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Journal of Archaeological Science

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The contributions of animal bones from archaeological sites: the past

and future of zooarchaeology
Teresa E. Steele a, b, *
Department of Anthropology, University of California, Davis, One Shields Ave., Davis, CA 95616-8522, USA
Max Planck Institute for Evolutionary Anthropology, Department of Human Evolution, Deutscher Platz 6, 04103 Leipzig, Germany

a b s t r a c t
Keywords: Humans and animals share a universal and intimate relationship that stretches from the earliest
Archaeozoology appearance of our lineage, through our prehistory, and into history and the modern era. Consequently,
Faunal analysis
zooarchaeology – the analysis of animal remains from archaeological sites – transcends all cultural,
temporal, and geographic boundaries, and therefore can be used to address questions spanning the
breadth of archaeology. Over the past decades, the field has grown tremendously, from a small group of
specialized practitioners to a large sub-discipline that is integrated with the larger field's research di-
rections. Zooarchaeological analyses have addressed questions ranging from forager ecology to the
processes of domestication to how animals function in the social realm. In each instance, methodological
developments have been crucial in allowing existing questions to be tackled and new questions to be
posed. One of the largest changes in zooarchaeology is a shift from considering how the environment has
shaped human societies to considering how humans have altered the environment, a process that began
deep in our prehistory. Applied zooarchaeology looks towards using zooarchaeological data to address
questions relevant to conservation biology, two fields that are becoming increasingly integrated. Stan-
dardization of data collection and presentation, open access to these data, and global access remain
challenges for the future.
© 2015 Elsevier Ltd. All rights reserved.

1. Introduction an important role in the growth and development of zooarch-

aeology. This perspective is in honor of his achievements.
While the importance of ancient animal bones for understand- The roots of faunal analysis are diverse, and this is reflected in the
ing human prehistory was recognized by some as early as the 1700s multiple names for the discipline. In North America, the discipline
– especially in establishing that humans even had a prehistory grew primarily out of archaeology and with a focus on anthropo-
(Grayson, 1983) – early examples of faunal analysis, the study of logical questions; it therefore became known as zooarchaeology,
ancient animal remains, are few and far between. As the discipline with an emphasis on human behavior. In Eurasia, the early discipline
of archaeology grew in the following centuries, the attention came from zoology and veterinary science, focused on domesticated
focused on the bones preserved within the archaeological sites also animals, with a close relationship to history and classics; it therefore
increased. While faunal analysis had its beginnings as marginal became known as archaeozoology, with an emphasis on the ani-
appendices to site reports, today the discipline, now commonly mals. Today, the largest organization supporting faunal analysts is
known as zooarchaeology, is a robust, well-integrated component the International Council of Archaeozoology (ICAZ), but both
of archaeology. This growth is reflected by the exponential increase zooarchaeology and archaeozoology are commonly-used terms.
in specialized practitioners and in papers covering zooarchaeo- Many zooarchaeologists also study faunal assemblages accumulated
logical topics published in major international journals. As a faunal by non-human agents as part of their work, so many prefer the more
analyst for the majority of his career and as an editor of the Journal encompassing terms of faunal analysis or paleozoology. Tradition-
of Archaeological Science for 30þ years, Dr. Richard G. Klein played ally, paleontology is more narrowly defined as studies of fossils that
are unrelated to humans, either because the assemblages predate
* Department of Anthropology, University of California, Davis, One Shields Ave.,
humans (broadly defined) or because the research questions are
Davis, CA 95616-8522, USA. Tel.: þ1 530 554 2804. focused on the taxonomy, evolution, adaptation, etc. of the animals,
E-mail address: independent of humans.
0305-4403/© 2015 Elsevier Ltd. All rights reserved.
T.E. Steele / Journal of Archaeological Science 56 (2015) 168e176 169

No matter the name, the field is focused on the study of animal in a regression analysis, instead of taking sex assignments as known
remains associated with archaeological materials to address ques- (which is what zooarchaeologists typically do, even if we are not
tions of humaneanimal-environmental relationships, often groun- certain about the designation), for each specimen a degree of cer-
ded by investigations of the biology and ecology of the animals tainty of the male or female designation can be included. In this
themselves. Zooarchaeological analyses have addressed topics way, Bayesian analyses do a good job of dealing with uncertainty at
ranging from diet, of both humans and animals, nutrition, resource all levels. Another major advantage to these approaches is that the
use, economy, prey choice, intensification and domestication pro- models are updated when new data are obtained; the results of one
cesses, herd management, breed development, economic and social model become the prior knowledge of the next (Litton and Buck,
exchange networks, technology and technological adaptations, 1995). Bayesian methods are part of a more general movement
human demography, urbanization, colonization, society, status and towards model-fitting statistical approaches in many scientific
wealth, gender, social identity, ideology, ritual, companionship, disciplines.
species biogeography, adaptation, environment, climate, and site Rogers was one of the first to apply model-fitting statistical ap-
formation, to name just a subset of the topics covered. This wide proaches to zooarchaeology (Rogers, 2000; Rogers and Broughton,
array of research questions reflects the universal and intimate re- 2001). He was trying to address what he referred to as “the statis-
lationships that animals have had with humans throughout pre- tical problem: to develop statistical methods that can use knowledge
history and history and reveals zooarchaeology's multidisciplinary about transport and attrition to draw inferences from bone counts”
and interdisciplinary nature and its diverse intellectual heritage. (Rogers, 2000). Rogers used prior knowledge about Hadza skeletal
Generally, these questions fall into three, not mutually exclusive, element transport decisions (including the relationship to animal
categories: methodological, anthropological, and biological (Reitz body size) and about the susceptibility of those elements to attrition
and Wing, 2008). One of the strengths of zooarchaeology is that it (based on bone density) to build a model that estimated the prob-
transcends all cultural, temporal, and geographic boundaries. Given abilities that an archaeological faunal assemblage reflects selectivity
the breadth of zooarchaeology, it is impossible to cover all aspects of in element transport. As elegant as this approach may have been, it
the discipline in this brief perspective, but I hope to provide a sense was not further employed, perhaps in part because it was not easy
of current, state-of-the-art research and where the field is headed in for most zooarcheologists to apply. However, more recently, as
the future. computers have become more powerful and the use of model-based
statistical approaches has become more widespread in other disci-
2. Methodological questions plines, general-use statistical packages have been implemented in
flexible statistical environments such as “R”, making these ap-
2.1. Quantitative approaches proaches more readily available to zooarchaeologists. For example,
Schoville and Ota rola-Castilla (2014) recently examined skeletal
Beginning with White's (White, 1952, 1953) landmark work on part transport with a model that incorporates the transport distance
quantifying the number of animals in a faunal assemblage, dis- from the kill to the camp and the number of carriers available. There
cussions of how best to quantify the many types of data collected is much space to apply Baysesian and other model-based statistical
during a zooarchaeological analysis are abundant (Grayson, 1984; approaches to our zooarchaeological data. However, a big challenge
Lyman, 1994, 2008). Zooarchaeologists count and measure things, will be appropriate quantitative training for all of us, especially our
all kinds of different things, and our sample sizes range from very students, who are the future of the discipline.
small to overwhelmingly large. All faunal analyses begin with a
basic count of the number of identified specimens (NISP). From 2.2. Zooarchaeology by mass spectrometry (ZooMS)
here, we may calculate the minimum number of elements (MNE),
minimum number of animal units (MAU), and/or minimum num- The application of wet-laboratory techniques, including the
ber of individuals (MNI). In addition, we may also quantify the ages analysis of stable isotopes and of ancient DNA preserved in faunal
and sexes of the animals and the seasons in which they died. remains, have lead to major advances in our ability to address a
Finally, we may then count modifications such as cut-marks, variety of zooarchaeological research questions. These topics form
impact-notches, burning stages, etc. These counts are ordinal or such large components of current archaeological science that in-
categorical rather than continuous variables, which present chal- dependent perspectives on stable isotopes (Sealy and Makarewicz,
lenges for analysis that have traditionally been addressed with non- in this volume) and on ancient DNA in zooarchaeology (Horsburg,
parametric approaches (Lyman, 2008), although model-fitting in this volume) are provided in this volume; I refer readers to
statistical approaches are starting to make inroads into the disci- those papers and therefore do not review the stable isotope and
pline. More challenging, however, is to consider: what is the pop- ancient DNA literature here. However, an additional method e
ulation that we are trying to reconstruct? How has that population Zooarchaeology by Mass Spectrometry (ZooMS) e is poised to also
been sampled? These are deeper statistical questions that impact zooarchaeology.
zooarchaeologists often do not fully consider or discuss when One of the biggest challenges many zooarchaeologists face is
choosing their analytical methods. also one of the most fundamental aspects of what we do e iden-
One approach that is becoming more commonly employed in tifying which animal species a bone came from. Our ability to
other disciplines is Bayesian statistics. The most widespread use of correctly identify the species depends on a number of factors e first
Bayesian methods in archaeology is for calibrating the results of and foremost is the preservation of the faunal remains. How
individual radiocarbon samples and for interrogating the spread of complete are they and how intact are their surfaces? Also impor-
radiocarbon results for a project. However, there is great potential tant is the experience of the analyst, the availability of suitable
for applying Bayesian approaches to questions in zooarchaeology. comparative material, including keys and atlases, the distinctions
Bayesian approaches combine prior knowledge, an analytical between the taxa (are the distinctions based on behavior or pelage,
model, and the likelihood of the observed data to produce (poste- and not reflected in skeletal morphology), the variability of the taxa
rior) probability distributions for different parameters of interest, involved, and the range of taxa that conceivably might be present in
which provides information about the uncertainty of the results the assemblage.
(Buck, 1999). One advantage of these approaches is that they allow Recently, however, mass spectrometry of organic components of
uncertainty to be easily incorporated into the analysis. For example, faunal remains has contributed to our ability to attribute difficult
170 T.E. Steele / Journal of Archaeological Science 56 (2015) 168e176

pieces to taxon, and the term Zooarchaeology by Mass Spectrom- further, ZooMS will become more widely used. We have only seen
etry or ZooMS (Buckley et al., 2009) has been coined. The method the beginning of proteomics applied to archaeology (Cappellini
takes advantage of the fact that between taxa there are small dif- et al., 2014; Hofreiter et al., 2012), and the research potential is
ferences in the peptide sequences of bone (and dentine) collagen enormous.
type I (COL1) that result in differences in mass, which can be
detected through mass spectrometry, providing the “fingerprint” of 3. Anthropological questions
a taxon. Essentially, the method provides a minimally destructive
way (if a buffer-only extraction is used (van Doorn et al., 2011)) of Many advances in zooarchaeology have come not from tech-
making taxonomic identifications of specimens that would nor- nology but from how we formulate our questions. Theoretical de-
mally be considered non-identifiable using traditional methods. velopments that help frame research questions and guide
The ability of ZooMS to discriminate between groups depends on interpretations have allowed zooarchaeology to make meaningful
the degree of difference between their peptide sequences and the contributions to our understanding of humans and have moved
completeness of the libraries to which the samples can be zooarchaeology into the position of “driving” research.
compared. The advantage of employing ZooMS before or instead of
ancient DNA is that ZooMS is only semi-destructive (while a small 3.1. Forager ecology
piece is necessary, it is not destroyed during processing and
therefore can be further analyzed by a variety of methods, including During most of our prehistory, humans were foragers, living by
AMS radiocarbon dating and ancient DNA analyses); relies on a hunting and gathering wild foods. Because of this dependence on
persistent molecule, especially compared to fragile DNA; depends hunting, zooarchaeological analyses have always been poised to
on an abundant molecule, COL1, which minimizes the potential for contribute significantly to research into hunter-gatherer subsis-
contamination; is relatively inexpensive; and is applicable to a large tence and economy. In recent decades, the incorporation of models
number of small samples with a fast turn-around time (Buckley from evolutionary ecology and Human Behavioral Ecology (HBE)
et al., 2010; Hofreiter et al., 2012). While ZooMS is still a new and into archaeological research has had profound influences on
specialized technique, it can provide major contributions towards zooarchaeology and provided a context in which the discipline is
answering zooarchaeological questions, as the following examples flourishing. In this volume, Codding and Bird (in this volume) give
illustrate. their perspective on the influence and future of behavioral ecology
Surprising to any zooarchaeologist who has tried to consistently in archaeology. HBE uses principles of evolutionary ecology to
distinguish sheep from goat bones morphologically, these two investigate human behavior. Because subsistence is fundamental,
species can be readily separated using ZooMS because of a differ- until recently zooarchaeological analyses of hunting and gathering
ence at one peptide in collagen molecules of these two taxa have relied most heavily on Foraging Theory, which posits that
(Buckley and Kansa, 2011; Buckley et al., 2010). When ZooMS was people will try to acquire the most energy while expending the
applied to morphologically unidentifiable juvenile “ovicaprid” re- least. This framework helps us predict which foods should be
mains from a late Neolithic site in Turkey, the majority of these chosen if energetic return is the only goal; of course, humans often
young animals were goats. These identifications highlighted that have other goals, so deviations are also interesting (see 3.3). Within
the assemblage reflected distinct herd-management strategies for zooarchaeology, the incorporation of models from HBE has
sheep and goat, where goats were harvested at a young age while contributed a much better understanding of why humans make the
more sheep were maintained into adult-hood for wool production, subsistence decisions that they do (prey choice, dietary diversity,
providing data about the economy of the prehistoric herders. and patch choice, to name a few) and how these choices interact
Another successful application of ZooMS has been to the “un- with environmental context and human demography. For example,
identifiable” component of a Paleolithic faunal assemblage from HBE provides a framework within which archaeologists can
central France. Because faunal assemblages with a larger number of investigate processes of intensification – the processes in which
identified specimens typically have a larger number of identified low ranked resources (usually resources such as hares, birds, tor-
taxa (richness), in this instance, because the assemblage was small, toises, and mollusks that do not provide a high caloric return
increasing the sample size of the “identified” specimens increased because they are small and energetically expensive to capture or
the species richness from five to nine. The addition of “rare” taxa process) are incorporated into the diet because of increased in-
influenced the environmental reconstruction based on the assem- vestment of human labor, either directly or through technology.
blage (Welker et al., 2015). Because of the implications of their presence, these small resources
Finally, ZooMS has been used to identify challenging remains, have received an increasing amount of attention from zooarch-
such as fish bones (Richter et al., 2011), eggshell fragments (Stewart aeologists (for example, see Thomas, 2015).
et al., 2013), hair (Buckley et al., 2013), and intensively worked It is often challenging for archaeologists to test our explanations,
antler combs (von Holstein et al., 2014). In this last instance, and HBE provides a framework for developing more rigorous tests.
because of the significance of the implication, the results were Broughton (1994, 2002) conducted early faunal analyses guided by
confirmed by ancient DNA. The application ZooMS had a significant Foraging Theory on a large assemblage from the edge of San
impact, because the original incorrect morphological identifica- Francisco Bay (California, USA). Using prey indices that reflect
tions had lead to erroneous interpretations of Scottish history (von relative prey abundance, he documented declines in the highest
Holstein et al., 2014). ranked prey (prey that provide the highest caloric return. In this
Zooarchaeologists should not be afraid of ZooMS taking over an case, elk, sturgeon, and goose) though time, while deer abundance
important aspect of our discipline, the identification of bones to increased in more recent times; these changes appear independent
taxa; instead they should see the method as an opportunity to open of climatic changes, and therefore environmental change does not
up new research questions. While the possibilities are great, ZooMS provide a likely explanation for changing prey choice. Broughton
only provides taxonomic identification; the method cannot provide argued that the shifts he identified reflect resource depression,
information about skeletal element, age, sex, surface modifications, where human activities e regular hunting of high-ranked animals
pathology or any of the other pieces of data that zooarchaeologists e created declines in foraging efficiency because the higher-ranked
routinely collect. As comparative reference libraries expand, high- prey declined in abundance and therefore lower-ranked prey were
throughput techniques improve, and expenses decline even taken more often. Broughton's conclusions were consistent with
T.E. Steele / Journal of Archaeological Science 56 (2015) 168e176 171

predictions from Foraging Theory, but additional testing was not determined using available (often limited) comparative material or
possible until recently when segments of ancient DNA were “known” archaeological samples. Recognizing the large datasets
extracted from archaeological elk samples. These sequences reveal available of traditionally collected linear measurements of molar
that there was a decline in elk genetic diversity and a population length and width, Evin et al. (2014) established cut-off values for
bottleneck consistent with when elk decline in abundance in the molar teeth, but these values were more statistically informed than
archaeological assemblage (Broughton et al., 2013), supporting the previous work and the authors provided an assessment of the risk
conclusion of resource depression over a change in prey preference. of error, increasing their utility. For a more nuanced investigation of
The successful testing of these predictions formulated through shape, Evin et al. (2013) used two-dimensional (2D) Geometric
Foraging Theory provide support for the validity of these ap- Morphometrics (GM) to investigate the shape of pig and boar
proaches and suggest ways that the models may be further tested molars, to compare with two measures of size (length and centroid
in the future. size, which is an overall measure of size typically used in GM
studies). GM examines shape independent of size in a way that
3.2. Reconstructing human demography preserves the geometry of the object (2D or 3D coordinates instead
of linear measurements) and allows for the visualization of shape
Archaeologists have long recognized the challenges of recon- changes along any axis of variation (Adams et al., 2004; Rohlf and
structing changes in human population sizes and densities from the Marcus, 1993). Using standardized digital photographs of occlusal
archaeological record. However, the small resources studied by surface of the pig and boar molars, Evin et al. (2013) digitally
zooarchaeologists provide some of the best evidence of human measured landmarks and sliding semi-landmarks (along curves or
demographic changes (Stiner et al., 2000, 1999). Mollusks in outlines), and they used Generalized Procrustes Analysis (GPA) to
particular have the potential to be quite informative, when it is obtain shape data (alternatively, elliptical Fourier analysis has been
possible to account for environmental variation. Ethnographic used (Larson et al., 2007)). These shape variables were then
work indicates that humans collect the largest individuals first, and analyzed and visualized with Principal Components Analysis (PCA).
some taxa of mollusks are slow but continuously growing and have Classifications as either domestic or wild pig were predicted using
long life histories, which means that there are not that many large leave-one-out cross-validation linear discriminant analyses, which
individuals and the largest individuals have the highest reproduc- also provides information on classification accuracy. Using predic-
tive potential. With increasing human predation pressure, the large tive linear discriminant analyses, unknown specimens are assigned
individuals are all removed and humans must then take smaller to either domestic or wild pig, and a posterior probability, which
ones, so with high human predation pressure we find a smaller depends on how close the unknown specimen is to the group mean,
average size in an archaeological assemblage (Steele and Klein, provides information about the degree of confidence in the classi-
2008). Increasing human population densities is now seen as the fication. Evin et al. (2013) also systematically investigated the in-
most likely explanation for the declining mollusk sizes in a number fluences of sample size. In sum, their analysis shows that for pig and
of case studies, with the interpretation being most convincing boar molars, shape provides the most robust indicator of domes-
when influences of environmental factors on growth rates can be tication status. Owen et al. (2014) take the investigations further by
discounted as the explanation (Erlandson et al., 2011, 2008; comparing shape variation in the entire crania of pigs, boar, and
Mannino and Thomas, 2002; McCoy, 2008). This approach has hybrids, again using 3D GM. 3D landmark coordinates were taken
been used with samples from the coasts of South Africa to inves- with a digitizer and transformed through GPA and investigated
tigate the relationship between demographic changes and tech- through PCA, Analysis of Variance, and Canonical Variates Analysis,
nological changes and models of modern human origins (Klein and and discriminant functions were produced. Shape variation was
Steele, 2013; Steele and Klein, 2005/06). visualized by morphing a 3D surface scan of a boar skull. To reflect
the realities of heavily fragmented zooarchaeological records, they
3.3. Domestication: humans as producers also successfully conducted these analyses on subsets of landmarks
that are usually preserved together (nasal, orbit, basicranium, etc.).
Few processes in human prehistory have been as profoundly In conclusion, pig and boar crania, whole and in parts, are suffi-
influential as the domestication of wild animals and plants, and this ciently distinct to be consistently separated morphologically. GM
shift towards food production has a long history of research within methods have been widely applied in biological anthropology to
zooarchaeology. Using faunal remains, the processes of domesti- human and primate skeletal morphology, but have not been widely
cation have been investigated typically through the examination of adopted in zooarchaeology. There is great potential for their
body size changes (with the expectation that body size decreases as employment to document morphological variation in faunal
animals evolve from wild to domesticated varieties) and mortality samples.
profiles (with the expectation that culling practices will target
optimal meat or milk production, which will likely result in a 3.3.2. Genetic changes indicative of processes of domestication
different signature than hunting). In addition, researchers look for Recent molecular advances have contributed significantly to
shifts in species composition, special treatment of animals such as investigations into the processes of animal domestication, and the
burials, and pathologies indicative of an animal's use for labor. most influential studies combine genetic analyses of modern and
While zooarchaeologists still research all of these areas, some ancient animals (see also Horsburg, in this volume). Most of these
recent studies highlight the directions the field is moving. studies have revealed complex patterns of migration, replacement,
and introgression (back-crossing; inter-breeding). This combined
3.3.1. Morphological changes indicative of processes of approach has contributed to a more detailed understanding of dog
domestication (Larson et al., 2012) and pig (Ottoni et al., 2013) domestication and
A series of studies investigating the distinction of wild boar from dispersal, as well as the domestication of other taxa. Both dogs and
domestic pigs in Eurasia illustrates the variety of methods available pigs appear to have undergone domestication in multiple locations,
to investigate the morphological changes indicative of the pro- dispersed widely, interbred with wild counterparts (wolves and
cesses of domestication. Traditionally, domestic pigs have been boars), and been frequently replaced by more recent migrants in
distinguished from their wild counter-parts on the basis of being many locations, processes which continued into the modern era,
smaller (Rowley-Conwy et al., 2012), and size cut-offs have been especially with the development of modern breeds. Future work
172 T.E. Steele / Journal of Archaeological Science 56 (2015) 168e176

will reveal the unique complexities for other taxa as they also un- of the species preserved their assemblages, although these bio-
derwent the processes of domestication. However, it is important logical analyses are often informed by questions about human
to note that genetic analyses are not the end-point of research into behavior. For example, my early work focused on reconstructing
domestication. Rather, the influence of molecular approaches is mortality profiles of red deer found in Paleolithic faunal assem-
similar to the influence of radiocarbon dating where methodolog- blages. Following earlier work by others (Klein, 1978; Klein et al.,
ical advances and applications provide clearer structure to the data, 1981), this required investigation into the relationship between
which then allows researchers to ask different and more nuanced tooth crown height and age as a way of estimating age-at-death in
questions of the data. the specimens as well as examining the mortality profiles created
by modern wolves and human hunters (Steele, 2003, 2006; Steele
3.3.3. Integrating morphological and genetic data and Weaver, 2012). While this research was driven by questions
The most detailed reconstruction of pig dispersal into Eurasia about human behavior, the work also provided insights into red
innovatively combined molecular, morphometric, and chrono- deer, elk, and wolf biology and ecology.
metric approaches, in what represents the cutting-edge of these Because of zooarchaeology's focus on skeletal remains, some of
types of analyses. Ottoni et al. (2013) analyzed ancient mitochon- our work into the biology of the animals that we study considers
drial DNA from archaeological pig and boar specimens from why bones are shaped the way they are. The study of ungulate
throughout western Eurasia. Uniquely, they conducted linear size ecomorphology, how skeletal shape changes with ecology, provides
and 2D geometric morphometric shape analyses on some of the an example. The goal of ecomorphology is to use the shape of
same molars that were sampled for DNA. In doing so, they were bones, usually the femora, metapodials, and astragali of bovids, to
able to reconstruct how size and shape varies with genetic clade reconstruct habitats independent of their taxonomic identification
and geography, providing a detailed picture of the pattern of pig (especially important when dealing with extinct species). The
dispersal across the region. theoretical justification for this approach is that locomotion should
Despite the significant contributions these genetic and be specific to habitat type (for example, bovids living in open
morphological studies have made toward our understanding of the habitats should have adaptations, visible on the skeleton, for fast
geography and chronology of domestication, these analyses cannot running). Most work has been based on traditional linear mea-
address the anthropological side of the research: questions about surements and used multivariate statistical techniques to generate
why did domestication happen in those locations at those times? discriminant functions to assign a habitat category to each spec-
And how did it happen in relation to changes to the human's society imen (DeGusta and Vrba, 2003; Plummer and Bishop, 1994;
and the animal's biology (morphology)? For example, why was pig Plummer et al., 2008), although sometimes shapes and ratios
molar size decreasing in size? As a result of changes in diet or in have been used (Kappelman et al., 1997). While issues of size and
disposition? To understand domestication, we need full integration phylogeny were discussed, they usually were not quantitatively
of genetic data with archaeological, anthropological, and environ- controlled, and for these reasons, Klein et al. (2010) have ques-
mental data (Larson and Burger, 2013). Although HBE approaches tioned the validity of traditional ecomorphological approaches.
have gained the most traction in hunter-gatherer studies, they are Using more sophisticated quantitative approaches, including
applicable to the transition to economies of production as well. Phylogenetic Generalized Least Squares analyses, recent studies
However, most applications have focused on the transition to have successfully separated morphological and taxonomic signals
cultivated plants with only a few that considered animal domes- and supported the ecomorphological approach (Barr and Scott,
tication (for example, see chapters in Kennett and Winterhalder, 2014; Scott and Barr, 2014). Highlighting the development of new
2006), leaving abundant avenues for the future growth of these approaches, Barr (2014) offers a detailed functional analysis of the
approaches in zooarchaeology. bovid astragalus to support the interpretation that morphology can
reflect habitat-specific locomotion. Unlike the earlier analyses,
3.4. Animals in human society which relied on linear measurements and ratios or indices derived
from them, Barr (2014) used a laser scanner to produce a 3D point
Of course, the humaneanimal relationship goes well beyond cloud of the specimen's surface. From this, Barr took linear mea-
subsistence and economy. The recent rise in social zooarchaeology surements but also precise areas of the articular surfaces. In this
provides evidence that, increasingly, the discipline is directing its way, advances in imaging technology, which have been more
attention to other aspects of human existence. For example, faunal widely applied in human and primate skeletal studies, are also
remains are being used to address questions about the rise of social contributing to zooarchaeology.
complexity, ritual, ideology, agency, identity, gender, inequality,
and patterns of consumption (Campana, 2010; Ervynck et al., 2003;
Russell, 2011). The consistency and orientation of cut-marks and 5. The future
patterns of bone breakage, typical components of zooarchaeo-
logical analyses, have been used to construct arguments about food 5.1. Research questions
sharing and cultural learning, important aspects of ancient societies
(Blasco et al., 2013; Stiner et al., 2009, 2011). Models from HBE have Zooarchaeology is united by methods of faunal analysis and
also considered that resources might be exploited for reasons other common interests in the relationship between humans, animals,
than their direct caloric return e because they may provide costly and the environment, which can contribute to our understanding of
signals of status, wealth, and/or fitness (Hildebrandt and McGuire, human prehistory and history in a variety of ways. Modern
2002; Hildebrandt et al., 2009). zooarchaeology is not conducted in isolation; it is one of many ap-
proaches that researchers can employ to address a variety of ques-
4. Biological questions tions. In this way, it is important for all sub-disciplines to integrate
results from all classes of data (for example, VanDerwarker and
Making informed interpretations of faunal assemblages requires Peres, 2010). As noted previously, the universal and intimate re-
an analyst to have a solid foundation in the biology and ecology of lationships that animals have had with humans throughout pre-
the species under study. For this reason, most zooarchaeologists history and history means that faunal analysis can contribute to
also conduct some basic research into these more biological aspects most “bigger-picture” questions archaeologists want to ask.
T.E. Steele / Journal of Archaeological Science 56 (2015) 168e176 173

Recently, a prominent group of mostly archaeologists pub- editing the data, and challenges of consistent data presentation,
lished a list of the “grand challenges for archaeology” (Kintigh analysis, and interpretation remain (Atici et al., 2013; Kansa et al.,
et al., 2014): twenty-five questions that represent archaeology's 2014).
most important scientific challenges. They reflect the discipline's
contributions to understanding the human condition with a view 5.3. Global balance
towards keeping the questions relevant to the modern world.
Zooarchaeology can provide the primary data for many of the One of the major challenges to the future of zooarchaeology is
questions, especially those concerning humaneenvironmental global balance in practitioners and access to resources. As of
interactions and human movement and mobility. For example, December 2014, ICAZ recorded 550 members from 51 countries.
some of the most meaningful contributions of zooarchaeological Africa has the lowest number of ICAZ members for any geographic
work have been on how humans have shaped the natural world, region, especially given the size of the overall populations e nine
on the processes, spread, and economics of animal management, are from South Africa, two from Egypt, and one from Botswana
and on the occupations of extreme environments. With each of (ICAZ, 2014; see also Pilaar Birch, 2014). In addition, practitioners
the other questions on the list, it is possible to see how faunal from under-represented countries face inequality in access to re-
analysis will contribute data to further the investigations. For sources. In this volume, Killick (2015) provides a perspective on
example, animals provide a means of measuring wealth and sta- these issues and how they relate to archaeological science. For
tus that can be used to investigate growing inequality and zooarchaeology, the implications are profound for issues ranging
emerging market systems. The authors of the list highlight that from the lack of: heritage management, adequately trained stu-
addressing most of these questions will require sophisticated dents and colleagues to take advantage of any faunal collections
modeling and large-scale synthetic research of large, systemati- that may exist, access to suitable comparative material, facilities to
cally collected datasets, themes that I also think describe the conduct analyses that require laboratory equipment, the ability to
future of zooarchaeology. export materials to places that may have these facilities, and op-
portunities to interaction with international colleagues through
5.2. Open-access sharing of standardized data travel to conferences and seminars. The result is the lack of rep-
resentation of many geographic areas in global discussions of
As noted in 4.1, understanding human prehistory and history archaeological research. In zooarchaeology, we are in a position to
depends on the compilation of large, consistent datasets that allows help in small ways (see also Plug, 2014), particularly by providing
multiple assemblages to be compared (for recent examples, training opportunities for students. Unfortunately, a career in
Arbuckle et al., 2014; Conolly et al., 2011). This can be facilitated by archaeology is often not lucrative or even possible for many of these
two objectives, the employment of standard protocols for data students. In many of these regions, heritage protection laws are
collection and presentation and the open-access sharing of data. weak or insufficiently enforced, which creates a cycle of missed
If data are to be shared, then they must be collected using the opportunities.
consistent methods, as much as possible. Grigson (2014: 6) states
that one of the first aims of the fledgling ICAZ in the early 1970s was 5.4. Human impacts and applied zooarchaeology
“to standardize measurements and data in publications”. ICAZ has
called for more standardization, and the current ICAZ protocols One of the major contributions of recent zooarchaeological an-
indicate that “publications should contain enough basic informa- alyses is our growing understanding of humans' impact on the
tion on analytical procedures and results to enable future scholars environment, not just recently but extending into prehistory. The
to evaluate the soundness of conclusions drawn and to replicate, to implications of these contributions are two-fold. First, as our depth
the extent possible, results presented, as is characteristic of any of understanding about the past impacts grows, we can no longer
competent scientific report” (Reitz, 2009). However, much of view humans as just influenced by the environment but that
zooarchaeological data collection, analysis, and presentation re- humans are also active participants in influencing our environments
mains opaque and unstandardized (Driver, 1992; Lyman, 1994; and our ecology, which includes the faunal resources that we
Wolverton, 2013). depend upon (Rick and Erlandson, 2009). This reflects a shift in view
In the least, publishing primary datasets should help make points about prehistoric humans' role in the natural world. The
methods more clear and replicable by other researchers, which examples of resource depression and reconstructing human
leads to the topics of data sharing, data publishing, and open- demography reflect this shift in perspective. Second, increasingly
access. ICAZ protocols encourage data sharing (Reitz, 2009). The zooarchaeologists are championing their data's ability to contribute
US National Science Foundation (2010) guidelines require the to conservation biology, in what has become known as applied
open-access distribution of data. Fortunately, zooarchaeology, and zooarchaeology, a term which reflects a more practical application
archaeology as a whole, is taking steps in the right direction (Kansa, for our zooarchaeological work (Braje and Rick, 2013; Lyman, 1996,
2010). More and more researchers are sharing and publishing their 2006; Lyman and Cannon, 2004; Makowiecki, 2003; Rick and
primary data. The ability for most journal articles to have associated Lockwood, 2013; Wolverton and Lyman, 2012). Learning about
on-line supplementary information, with no space limitations, fa- how humans have modified species distributions can be best
cilitates this. Open-access platforms designed for archiving accomplished through the examination of past distributions, data
archaeological data are becoming more common and frequently that can be collected from faunal remains. For example, studies of
used; such as Open Context, tDAR: The Digital Archaeological Re- past distributions of northern fur seals (Newsome et al., 2007) and
cord, Archaeology Data Service, and Digital Antiquity, and some are herrings (McKechnie et al., 2014) highlight the importance of the
available specifically for zooarchaeological data, through Bone- archaeological record for reconstructing past species distributions
Commons and more (Kansa and Kansa, 2011). ICAZ is supporting and abundances with implications for the future of the species. As
open-access efforts with their Junior Researcher Open Zooarch- indicated from the zooarchaeological record (Newsome et al., 2007),
aeology Prize (Kansa, 2014). Despite agreement that open-access northern fur seals once ranged throughout the western coast of
data sharing is generally a good idea, sufficient institutional sup- North America and likely had on-shore rookeries; more recently,
port and data standardization remain challenging. Integrating human exploitation severely limited the population to restricted
many datasets can require a large commitment to reviewing and off-shore breeding communities. In addition to this work providing
174 T.E. Steele / Journal of Archaeological Science 56 (2015) 168e176

previously inaccessible ecological information, as the fur seal pop- reveals complexity in the westward spread of domestic animals across neolithic
Turkey. PLoS One 9, e99845.
ulation rebounds due to protection, the zooarchaeological record
Atici, L., Kansa, S.W., Lev-Tov, J., Kansa, E.C., 2013. Other people's data: a demon-
indicates where new breeding colonies may form, and these sites stration of the imperative of publishing primary data. J. Archaeol. Method
can then be protected. Early examples of applied zooarchaeology Theory 20, 663e681.
struggled to gain the attention of conservation biologists. Fortu- Barr, W.A., 2014. Functional morphology of the bovid astragalus in relation to
habitat: controlling phylogenetic signal in ecomorphology. Am. J. Phys.
nately, more recently the two fields are becoming more integrated Anthropol. 275, 1201e1216.
and the terms historical ecology or conservation archeobiology are Barr, W.A., Scott, R.S., 2014. Phylogenetic comparative methods complement
sometimes used to refer to this fledgling field (Braje and Rick, 2013; discriminant function analysis in ecomorphology. Am. J. Phys. Anthropol. 153,
Szabo , in press). The need for insights gained from Applied Blasco, R., Rosell, J., Domínguez-Rodrigo, M., Lozano, S., Pasto , I., Riba, D.,
Zooarchaeology is becoming more urgent as biologists must make Vaquero, M., Peris, J.F., Arsuaga, J.L., Castro, J.M.B.d., Carbonell, E., 2013. Learning
management decisions for species that already face challenging by heart: cultural patterns in the faunal processing sequence during the Middle
Pleistocene. PLoS One 8, e55863.
circumstances due to habitat loss and climate change. Braje, T.J., Rick, T.C., 2013. From forest fires to fisheries management: anthropology,
conservation biology, and historical ecology. Evol. Anthropol. 22, 303e311.
6. Conclusions Broughton, J.M., 1994. Declines in mammalian foraging efficiency during the late
Holocene, San Francisco Bay, California. J. Anthropol. Archaeol. 13, 371e401.
Broughton, J.M., 2002. Prey spatial structure and behavior affect archaeological tests
In conclusion, zooarchaeology has made incredible strides, of optimal foraging models: examples from the Emeryville Shellmound verte-
moving from a marginalized small group of practitioners to a brate fauna. World Archaeol. 34, 60e83.
Broughton, J.M., Beck, R.K., Coltrain, J.B., O'Rourke, D.H., Rogers, A.R., 2013. A late
vibrant community of researchers who are having a major influ- Holocene population bottleneck in California tule elk (Cervus elaphus nann-
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Buck, C.E., 1999. Bayesian statistics for archaeology. In: Gaul, W., Locarek-Junge, H.
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creatively about their data, promote the sharing of mutually useful pp. 547e554.
datasets, and strive for global access. By following these principles Buckley, M., Collins, M., Thomas-Oates, J., Wilson, J.C., 2009. Species identification
by analysis of bone collagen using matrix-assisted laser desorption/ionisation
the discipline will thrive through the 21st century and beyond.
time-of-flight mass spectrometry. Rapid Commun. Mass Spect. 23,
Websites for organizations mentioned in the text Buckley, M., Kansa, S.W., 2011. Collagen fingerprinting of archaeological bone and
teeth remains from Domuztepe, south eastern Turkey. Archaeol. Anthr. Sci. 3,
Archaeology Data Service: Buckley, M., Kansa, S.W., Howard, S., Campbell, S., Thomas-Oates, J., Collins, M.,
BoneCommons: 2010. Distinguishing between archaeological sheep and goat bones using a
single collagen peptide. J. Arch. Sci. 37, 13e20.
Digital Antiquity: Buckley, M., Melton, N.D., Montgomery, J., 2013. Proteomics analysis of ancient food
Guides to Good Practice: http://guides.archaeologydataservice. vessel stitching reveals >4000-year-old milk protein. Rapid Commun. Mass Spect. 27, 531e538.
Campana, D.V., 2010. Anthropological Approaches to Zooarchaeology: Complexity,
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Shennan, S., 2011. Meta-analysis of zooarchaeological data from SW Asia and SE
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DeGusta, D., Vrba, E., 2003. A method for inferring paleohabitats from the func-
This paper is dedicated to Richard G. Klein, who has inspired tional morphology of bovid astragali. J. Arch. Sci. 30, 1009e1022.
Driver, J.C., 1992. Identification, classification and zooarchaeology. Ciraea 9, 35e47.
many, including me. I thank the editors for inviting me to submit
Erlandson, J.M., Braje, T.J., Rick, T.C., Jew, N.P., Kennett, D.J., Dwyer, N., Ainis, A.F.,
this perspective. I wrote this manuscript with financial support Vellanoweth, R.L., Watts, J., 2011. 10,000 years of human predation and size
from the University of California, Davis (USA) and while on sab- changes in the owl limpet (Lottia gigantea) on San Miguel Island, California.
batical at the Max Planck Institute for Evolutionary Anthropology J. Arch. Sci. 38, 1127e1134.
Erlandson, J.M., Rick, T.C., Braje, T.J., Steinberg, A., Vellanoweth, R.L., 2008. Human
(Leipzig, Germany) and the University of Cape Town (South Africa). impacts on ancient shellfish: a 10,000 year record from San Miguel Island,
Given the vast amount of literature that I could have cited in this California. J. Arch. Sci. 35, 2144e2152.
perspective, my apologies to anyone who feels that I overlooked Ervynck, A., O'Day, S.J., Van Neer, W., 2003. Behaviour Behind Bones: the
Zooarchaeology of Ritual, Religion, Status and Identity. Oxbow, Oxford, United
their work. The editors, three anonymous reviewers, Susan Lagle, Kingdom.
Naomi Martisius, Tim Weaver, and Frido Welker provided com- Evin, A., Cucchi, T., Cardini, A., Vidarsdottir, U.S., Larson, G., Dobney, K., 2013. The
ments on earlier versions of this manuscript. long and winding road: identifying pig domestication through molar size and
shape. J. Arch. Sci. 40, 735e743.
The International Council for Archaeozoology (ICAZ) provides an Evin, A., Cucchi, T., Escarguel, G., Owen, J., Larson, G., Vidarsdottir, U.S., Dobney, K.,
important forum for supporting the zooarchaeology community, as 2014. Using traditional biometrical data to distinguish west palearctic wild boar
do their numerous Working Groups, which reflect many of the and domestic pigs in the archaeological record: new methods and standards.
J. Arch. Sci. 43, 1e8.
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