Dietary reconstruction, mobility and the analysis of ancient skeletal tissues: Expanding the...

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Journal of Archaeological Science 56 (2015) 146e158

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Journal of Archaeological Science

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Dietary reconstruction, mobility, and the analysis of ancient skeletaltissues: Expanding the prospects of stable isotope research inarchaeology

Cheryl A. Makarewicz a, *, Judith Sealy b

a Institute for Prehistoric and Protohistoric Archaeology, Christian-Albrechts University, Kiel, Johanna-Mestorf Strasse 2-6, Kiel, D-24118, Germanyb Department of Archaeology, University of Cape Town, Private Bag X3, Rondebosch, 7701, South Africa

Keywords:Dietary reconstructionNitrogenOxygenHydrogenIsoscapesMixing-modelDietary routingAmino-acid

* Corresponding author.E-mail addresses: [email protected]

[email protected] (J. Sealy).

http://dx.doi.org/10.1016/j.jas.2015.02.0350305-4403/© 2015 Published by Elsevier Ltd.

a b s t r a c t

The use of stable isotope ratio analysis in archaeology has exploded over the past few decades to thepoint where it is now an established tool that is routinely used to investigate questions relating to dietand mobility. Early applications focused mostly on the analysis of human skeletal tissues as a way toreconstruct major shifts in human diet, but current stable isotopic approaches have expanded to includehigh resolution analyses of human, animal, and plant remains, which are helping to better define theresource exploitation and management strategies that underscore changes in the human diet. In addi-tion, stable isotopic data sets are now regularly filtered through interpretive archaeological theoreticalframeworks to explore socially mediated food acquisition and consumption choices, mortuary practices,and social identity. Much work remains to be done in documenting the biological and ecological variationin the distribution of stable isotopes in ancient food webs and the mechanisms responsible for theisotopic signals observed in archaeological plant and animal tissues. Here, we identify several areas instable isotope analysis where additional ‘first principles’ driven research would help to improve existingisotopic methods, or develop new ones, and consequently improve our ability to answer questions ofarchaeological significance. We consider the strengths and limitations of the application of stable isotopeanalysis to ancient skeletal material obtained from archaeological contexts. We also pay particularattention to nitrogen isotopic variation in ancient ecosystems, organic oxygen and hydrogen isotopesto;mixing models as a means of estimating source contributions in human diet, mobility, and isoscapes;and to how compound specific analyses may help detangle dietary routing. We conclude with a plea forgreater scientific rigour and more informed use of stable isotope analyses and call for a closer integrationof stable isotope analysis with other aspects of archaeological research programmes, in order to optimisethe information that isotopes can provide.

© 2015 Published by Elsevier Ltd.

1. Introduction

Over the past several decades, the role of stable isotope analysisin archaeology has been transformed from a highly specializedapproach, employed only occasionally, to a relatively conventionalapplication routinely incorporated into archaeological projectsbefore the first trowel is even planted in the ground. Indeed, stableisotope analysis in archaeology has exploded over the past decade

(C.A. Makarewicz), Judith.

to become one of the most frequently used methods in thearchaeological sciences (Canti and Huisman, 2015; Rehren, 2015;Szpak, 2014). In the early days, isotope approaches were applied toa relatively narrow range of questions focusing on major humandietary transitions, e.g., the development and spread of maizeagriculture and marine vs. terrestrial resource exploitation. Theseissues could readily be addressed using major isotopic dichotomies,such as that between carbon isotopes in C3 and C4 plants or ni-trogen isotopes in marine and terrestrial systems. Today, stableisotope analysis is used to investigate an incredibly diverse spec-trum of topics relating to dietary, subsistence, mobility, and socialpractices of the human past (Lee-Thorp, 2008). Isolating themechanisms involved in plant and animal management and

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C.A. Makarewicz, J. Sealy / Journal of Archaeological Science 56 (2015) 146e158 147

domestication processes, documenting human mobility and live-stock movement, establishing social organization and complexity,exploring identity construction and gender differences are but asampling of the topics now being pursued using stable isotopeanalysis (Ambrose et al., 2003; Balasse et al., 2001, 2002; Barrettand Richards, 2004; Gregoricka, 2013; Makarewicz and Tuross,2006, 2012; Müldner et al., 2014; Sealy, 2006; Somerville et al.,2013; Tung and Knudson, 2008). This expanded range of applica-tions to increasingly complex questions is extending the limits ofthe field. Unfortunately, it also leads to greater potential for ill-conceived or poorly executed studies.

Below, we briefly detail how stable isotope analysis was initiallyincorporated into archaeology and later became one of the fastest-growing, most dominant fields in archaeological science. This rapidgrowth reflects how amenable stable isotope analysis is to inves-tigating questions about diet, dietary differentiation, life history,and other topics of interest to archaeologists. In addition, we arguethat a confluence of technical advances in mass spectrometry anddevelopments in archaeological method and theory was a keystimulant amplifying the use of stable isotope analyses in archae-ology to the level of near ubiquity we see today. We then discussseveral pitfalls that have contributed to some misinterpretations ofstable isotope values obtained from ancient human and animaltissue. These include an unsophisticated understanding of isotopicvariation at the floral base of the foodweb and an over estimation ofthe degree to which stable isotope mixing models can providequantitative information about diet. Finally, we identify severalgaps in current archaeological stable isotope research and offerrecommendations on where the field might head, with particularattention to the potential for isoscapes to generate more refinedinsights into human mobility and animal movement, how uncon-ventional analyses such as hydrogen and oxygen isotopic analysesof collagen could offer new insights into seasonality and diet, andhow compound specific work may shed new light on the dietaryand metabolic pathways responsible for driving the isotopiccomposition of bulk collagen. We believe that these subjects areparticularly topical at this juncture in the development of the field;different authors might well have chosen others. We do notattempt to provide an exhaustive review of the extensive literatureon isotope-based palaeodietary tracing.

2. Stable isotope analysis: from the domain ofbiogeochemistry to the domicile of archaeology

Stable isotopes record dietary and environmental inputs in thetissues of plants and animals and provide a means to investigatehuman diet and mobility, as well as explore the ways in whichpeople exploited wild and domesticated plant and animal re-sources in ancient societies. The strength of stable isotope analysisas a dietary tracer makes the approach well-suited to archaeology,where the material remains of certain categories of foods are oftenabsent or under represented in the depositional record. Much ofthe research that uses a stable isotopic approach in archaeologicalcontexts draws on the far larger, longer-established field of isotopegeochemistry in the earth and plant sciences (Craig, 1953, 1954;Park and Epstein, 1960). The field of dietary tracing emerged outof radiocarbon dating, after it was recognised that radiocarbondeterminations derived from plants using the C4 photosyntheticpathway yielded dates that were consistently ‘too young’ relativeto those fixing carbon through the C3 pathway (Bender, 1968).Subsequent research capitalized on this observation to use stableisotopes as a natural tracer of the flow of carbon isotopes throughfoodwebs (e.g., DeNiro and Epstein, 1976, 1978; Van der Merweand Vogel, 1978; Vogel and van der Merwe, 1977). The field soongrew to include studies of the distribution of nitrogen isotopes in

animal tissues (DeNiro and Epstein, 1981; Minagawa and Wada,1984; Schoeninger et al., 1983; Schoeninger and DeNiro, 1984),oxygen isotope fractionation effects between local meteoric wa-ters, mammalian body water and bone phosphate (Longinelli,1984; Luz et al., 1984), and geologically sourced strontium andthe expression of 87Sr/86Sr in human bones and teeth (Ericson,1985).

The field of stable isotope dietary tracing began with seminalstudies on the spread of maize agriculture in North America (Vander Merwe and Vogel, 1978; Vogel and van der Merwe, 1977), thetransition frommarine to terrestrial diets during the Neolithizationof northern Europe (Tauber, 1981), and subsequent work on similartopics but within different environmental contexts (Noe-Nygaard,1988; Schwarcz et al., 1985; Sealy and van der Merwe, 1985,1986; Walker and DeNiro, 1986). Studies focused on archaeolog-ical questions were paralleled by research that aimed to documentthe range of natural variation in stable isotopes and better under-stand the relationship between diet and carbon and nitrogenisotope ratios in consumer tissues (DeNiro and Epstein, 1981;Tieszen et al., 1983; Van der Merwe and Medina, 1989), some-times through analysis of best-case-scenario samples, whichhappened to be archaeological material.

A key question here, one that emerged early on in the devel-opment of the discipline and that remains highly relevant, revolvesaround the extent to which the isotopic ratios we measure inancient bones and teeth are reliable: can we identify and deal withpost-depositional (diagenetic) contamination? The 1980s and1990s saw a series of breakthroughs in bone preservation researchthat examined how diagenesis might influence the isotopic integ-rity of the organic and inorganic components of ancient hard tis-sues (Krueger, 1991; Lee-Thorp and van der Merwe, 1987; Masters,1987; Nelson et al., 1986; Schoeninger and DeNiro, 1982, 1983;Sullivan and Krueger, 1981, 1983; Tuross et al., 1988). Many of thesestudies focused on Pleistocene materials, given the likelihood thatolder samples would exhibit diagenetic alteration to biogenicmatrices accompanied by shifts in isotope values. This workdeveloped sample pretreatment procedures, specified methods foridentifying diagenesis and, ultimately, broadcasted the promiseand limitations of stable isotope analyses conducted on ancientbones and teeth in both geochemical and archaeological journals,providing the necessary confidence that isotopic results obtainedfrom older and/or less well-preserved archaeological materialcould be reliable. A major push of stable isotope analyses intoarchaeology followed the slipstream of diagenesis research to focuson archaeological questions relating to paleoenvironmentalreconstruction and hominid dietary change in Pleistocene envi-ronments (Bocherens et al., 1991, 1994, 1995; Fizet et al., 1995;Iacumin et al., 1996, 1997; Lee-Thorp et al., 1994).

This trend was further promoted by a host of technical andtheoretical developments within mass spectrometry and anthro-pological archaeology. The development and widespread avail-ability of continuous-flow mass spectrometry during the early1990s increased dramatically the volume of sample through-putwhile simultaneously decreasing the cost of analysis (Brennaet al., 1997). This technical advance provided the means to pro-cess the larger numbers of analyses required in archaeological datasets to disentangle environmental from human inputs, better cap-ture the range of variability associated with human activities, andpinpoint the role of diagenesis in influencing the stable isotopevalues of sampled archaeosubstrates. In addition, the decreasedanalytical costs afforded by continuous-flow mass spectrometrywere more in tune with archaeological research budgets, whichhelped to turn archaeologists into consumers of stable isotopeanalysis who, ultimately, dictated how and when to deploy stableisotope analysis on archaeological materials.

C.A. Makarewicz, J. Sealy / Journal of Archaeological Science 56 (2015) 146e158148

Coincident with the increased accessibility of mass spectrom-etry was a shift within archaeological theoretical agendas thatfurther facilitated the uptake of stable isotopic approaches intoarchaeology. The processual approach championed by Binford,which conceptualized human behaviors largely as adaptations toenvironments and sought to evaluate evolutionary models of cul-tural process through hypothesis-testing, was under increasinglyheavy critique during the mid-1980s for its failure to address di-versity in human practice and social agency in ancient commu-nities (Hodder, 1986; Shanks and Tilley, 1987). While processualarchaeology tended to focus primarily on questions relating tohuman subsistence and economy, systems interaction, and taph-onomic processes (Binford, 1962, 1967; Clarke, 1968; Trigger, 2006),post-processualists explored the subjectivity of interpretation inarchaeology, the role of symbols and ideologies in shaping socialworlds and political structures, and how human manipulation ofsocial rules was a dynamic force of cultural change (Brumfiel, 1983;Earle and Preucel, 1987; Hodder, 1982; Miller and Tilley, 1984;Shanks and Hodder, 1995). The initial post-processual movementengendered the subsequent florescence during the mid-1990s andearly 2000s of theoretical frameworks that highlighted theimportance of the roles of acting individuals, social practice, andidentity in shaping human worlds. This body of interpretivearchaeological theories led archaeologists to ask fresh questionsabout the social dynamics of ancient communities that could beinvestigated using a variety of ‘archaeological scientific ap-proaches’, including stable isotope analysis. The theoretical trac-tion provided by interpretive archaeologies unlocked whole newarenas of stable isotope research, including the collective social andpolitical motivations for human mobility, socially-constructedproscriptions for food acquisition and distribution within com-munities, and the ritual circulation of animals (Ambrose et al.,2003; Henton et al., 2014; Kellner and Schoeninger, 2008;Laffoon et al., 2013; Müldner and Richards, 2005; Sj€ogren andPrice, 2013).

Stable isotope analysis in archaeology has evolved into anintegrative discipline that considers both the subsistence and socialreasons that underscore human dietary andmobility decisions. Theapproach nevertheless requires appropriate research designs thattake account of the biological and ecological mechanisms respon-sible for the isotopic signals observed in archaeological plant andanimal tissues. Below, we discuss some of the pitfalls and promisesof stable isotopic approaches as we see them today and criticallyexamine examples relevant to the points we wish to make.

3. Nitrogen isotopic variation in foodwebs and consumers

Nitrogen isotopes offer a window into dietary behaviors relatedto protein intake, including terrestrial and aquatic protein-containing plants and animals. The stepwise enrichment of 15Nwith each upward shift in trophic level, as first demonstrated byMinagawa and Wada (1984) and Schoeninger and DeNiro (1984),fostered a series of studies using d15N of consumer tissues toinvestigate the role of meat in the human diet (reviewed inSchoeninger and Moore, 1992). Although such work provided, andcontinues to provide, important insights into human dietary di-versity, much of it also underestimates the complexity of the dis-tribution of nitrogen isotopes in ecosystems and how dietary 15N isincorporated into body tissues and then subsequently expressed incollagen d15N values.

Typically, the trophic level of consumers and the relative con-tributions of plant and animal foods in omnivores are establishedby comparing the d15N values of the omnivorous species of interest,usually a human or hominid, with the values of herbivores andcarnivores occupying the same ecosystem (e.g. Bocherens et al.,

1991, 2005; Richards and Trinkaus, 2009). Although this frame-work provides a starting point for subsequent interpretation, it isoverly simplistic in its modelling of both the distribution of nitro-gen isotopes in the plant-based portion of the food web andincorporation of dietary 15N into body tissues.

Interpreting the d15N values of higher trophic level consumersdepends in large part on a strong understanding of the nitrogenisotopic structuring in floral and faunal communities. There is nowan extensive body of research documenting the relationship be-tween macro-environmental inputs (i.e. temperature, precipitationand humidity levels) and microhabitat (i.e., local source N) indetermining floral and faunal d15N values (see review by Szpak,2014). Most archaeologists' knowledge of the distribution of ni-trogen isotopes at the base of ancient food webs is based largely onarchaeological studies of northern European terrestrial biomes,which are often interpreted as exhibiting little or no nitrogen iso-topic variability, despite evidence for moderate variability(Bocherens et al., 1997, 2015; Drucker et al., 2011; Noe-Nygaardet al., 2005). This misconception is the foundation of many inter-pretative frameworks employed in such environments, and liesbehind the practice of using herbivore d15N as a measure of climatechange (particularly aridity; see Drucker et al., 2011; Stevens et al.,2008) in prehistory, when other isotopes, such as oxygen, providemuch more direct evidence for environmental shifts. Given theimportance of soil nitrogen sources in defining plant d15N values,which reflect microbially mediated nitrogen cycling and denitrifi-cation processes occurring at local scales in addtion to broaderclimatic factors (see Szpak, 2014), too much weight is placed ontemperature and aridity as the primary factors driving nitrogenisotopic change in herbivores, calling into question the practice ofrelying on herbivore d15N values as away of tracking environmentalchange.

The relatively recent recognition that there are nitrogen isotopicdistinctions between herbivorous taxa corrects earlier mis-conceptions that assigned nitrogen isotopic homogeneity to planteaters, but some fundamental misunderstandings about the dis-tribution of nitrogen isotopes in herbivores remain, and these haveconsequences for interpreting the d15N values of consumers athigher trophic levels. It is important to emphasize that the d15Nvalues expressed in different species of wild or domestic herbi-vores depends on their dietary preferences and seasonality ofavailable food sources (Balasse et al., 2001; Darimont andReimchen, 2002; Makarewicz, 2014; Sponheimer et al., 2003a).For example, differences of ca. 3‰ have been reported betweenmean d15N values of different herbivore species feeding within thesame ecotone (Codron et al., 2005), and a similar degree of varia-tion has been observed along the tail hair of elephants in EastAfrica, reflecting seasonal differences in d15N of food consumed(Cerling et al., 2009). In contexts where humans intentionallymanipulate domesticated plant and animal resources, there ispotential for an even wider range of nitrogen isotopic variation,reflecting human decisions to amend agricultural fields withexogenous sources of nitrogen through manuring, foddering ani-mals with agricultural by-products, or grazing herds in differentpastures characterized by different d15N values (Bogaard et al.,2007; Fraser et al., 2011; Makarewicz, 2014; Makarewicz andTuross, 2012).

Animal physiology and metabolism are also key factors deter-mining the nitrogen isotopic composition of consumers (Reitsema,2013). Controlled feeding studies have revealed that different her-bivore species ingesting identical diets exhibit values that maydiffer by as much as 4.5‰ (in hair keratin), suggesting that inter-specific differences in digestive physiology, e.g., foregut v. hindgutfermenting, can heavily influence body tissue nitrogen isotopevalues (Sponheimer et al., 2003a). Diet-to-tissue fractionation and


the factors that control this remain poorly understood for nitrogenisotopes, and there appears to be considerable variation dependingon, among other factors, the protein content of the diet (Caut et al.,2009; Hedges and Reynard, 2007; O'Connell et al., 2012;Sponheimer et al., 2003a,b).

Environmental, anthropogenic and metabolic factors have beenincreasingly acknowledged as sources of nitrogen isotopic distinc-tion in herbivores, and there are clearly numerous factors thatcontribute to nitrogen isotopic shifts in herbivores that do notnecessarily indicate trophic level differences. However, the notionpersists that herbivore d15N values should be low, when in fact,herbivore collagens may be highly enriched in 15N. The best knownexamples are from arid regions, where bone collagen and toothdentin d15N values consistently reaching over 10‰ due to ingestionof 15N enriched plants which characterize arid environments(Codron et al., 2005; Hartman, 2011; Hartman and Danin, 2010;Heaton et al., 1986; Makarewicz and Tuross, 2006; Murphy andBowman, 2006; Pate, 1998). Herbivores inhabiting moretemperate environments generally exhibit lower d15N values,reflecting different environmental inputs, but some can still exhibitelevated nitrogen isotope values relative to the total local foodweb(e.g., Bocherens et al., 2005). More often than not, herbivoresexhibit d15N values higher than the 3‰e6‰ called for in the ca-nonical trophic level isotope pyramid still consulted, eitherimplicitly or explicitly, by archaeologists and some stable isotopeanalysts alike. This, combined with overall nitrogen isotopic vari-ation in plants and animals, has important implications for inter-preting the d15N values of consumer at higher trophic levels, whichintegrate dietary source 15N into their tissues.

Accordingly, particuarly in cases where herbviores are charac-terized by high d15N values and/or nitrogen isotopic variability, itbecomes increasingly difficult to precisely define what is drivingthe nitrogen isotopic composition of higher level consumers thateat these herbivores: ingestion of higher amounts of meat or con-sumption of some animal products characterized by high d15Nvalues. This outdated heuristic device should be discarded in favorof more flexible models that are keyed in to local environmentalconditions while accounting for use by humans of exploitationstrategies that involved modification of plant and animal resources.

Along these lines, it is essential that palaeodietary studies usingnitrogen isotopes adequately characterize the range of nitrogenisotope variation in foods consumed through robust sampling ofcontemporaneous plant and animal tissues. Attempts to use stableisotopes to reconstruct the diets of Neanderthal and early modernhumans illustrate these inherent challenges. In these studies, as inmany others, the trophic level of consumers and the relative con-tributions of plant and animal sources are estimated by comparingthe d15N values of the species of interest with the values of herbi-vores and carnivores occupying the same ecosystem. d15Ncoll valuesof Neanderthals in Western and Central Europe are high e highereven than contemporaneous carnivores. This has been interpretedas evidence of diets that centred on the consumption of meat,including large animals such as mammoth and woolly rhino(Bocherens, 2009; Drucker and Bocherens, 2004; Richards andTrinkaus, 2009; Salazar-García et al., 2013).

Prompted, in part, by these somewhat provocative suggestionsabout Neanderthal carnivory, researchers have set out to investi-gate dental microwear, plant microfossils trapped in dental calcu-lus, fecal biomarkers, and other innovative approaches topalaeodietary reconstruction (El-Zaatari et al., 2011; Fiorenza et al.,2015; Henry et al., 2011, 2014; Sistiaga et al., 2014). Plants tend tocontain less protein (i.e., less nitrogen) than meat, so plant foodswill be less visible in consumer d15N values. This work hasdemonstrated that Neanderthals did indeed consume plant foods,despite their apparent invisibility in collagen d15N values, and

highlights one of the difficulties in interpreting stable isotopevalues: to what extent can they provide quantitative informationabout diet? We return to this question below.

How did the diets of early modern humans compare with thoseof Neanderthals? Were modern humans better than Neanderthalsat accessing food and other resources, contributing to their successas a species (Conard, 2006; d’Errico and S�anchez Go~ni, 2003)?d13Ccoll and d15Ncoll values appear more variable and more positiveamong early modern humans than among Neanderthals, suggest-ing that dietary breadth probably was greater, very likely includingmore fish and other aquatic resources (Fu et al., 2014; Richardset al., 2001; Richards and Trinkaus, 2009).

Comparisons such as these are valid only if the environmentremains stable over the period of interest. We have known for someyears that there was a Pleistocene/Holocene shift in d13C and ananomaly in d15N just after the Last Glacial Maximum (Hedges et al.,2004; Richards and Hedges, 2003; Stevens and Hedges, 2004), butthese do not bear directly on the Neanderthal/modern humantransition. Very recently, Bocherens et al. (2014) have reported that,at least in the Dordogne, terrestrial fauna from the early Aurigna-cian show elevated d15Ncoll compared with preceding and suc-ceeding periods. If this pattern is widespread, more positive d15Ncollin early modern humans may simply reflect environmental ratherthan behavioural (dietary) differences. The same study did not findchanges in carbon isotopes over time, although d13Ccoll values inearly modern humans are also more positive (Richards andTrinkaus, 2009; Fig. 1).

It is challenging to understand isotopic patterning in environ-ments of the distant past for which there may be no modern ana-logues. Even in a modern context, we need a better understandingof nitrogen cycling in vivo, and in animals consuming different di-ets. If we are to interpret isotope measurements of consumers inany ecosystem (ancient or modern) properly, we require detailed,geographically and temporally resolved isotopic surveys ofcontemporaneous environments. This will require heavy invest-ment in the analysis of large numbers of archaeological faunalspecimens, well beyond the token sampling of a few bones thatcharacterizes many studies, in order to establish the extent of ni-trogen isotope variation within and between species and provide abetter means to interpret d15N values in consumers.

4. Mobility, isoscapes, and serial sampling

The isotopic analysis of biogenic tissues for the purposes ofelucidating the movement of people, animals, goods, and foods hasbeen a major focus of recent archaeological research, (Balasse et al.,2002; Bentley, 2007; Britton et al., 2009; Dupras and Schwarcz,2001; Hedges et al., 2005; Meiggs, 2007; Price et al., 2006).Reconstructing the mobility patterns of both people and the ani-mals they exploited is key to understanding resource acquisitionstrategies, social relationships, and the permeability of politicalboundaries, while tracking the transport of goods and foodstuff canhave important implications for delineating the shape of distribu-tion systems and trade networks (Barrett et al., 2008; Guiry et al.,2012; Meiggs, 2007). Isotope-based mobility studies on ancienthumans and animals are increasingly abundant, but many areconstrained by coarse scales of analysis, limiting interpretations tostatements about the ‘local’ or ‘non-local’ origins of sampled in-dividuals. This simple framework does not reflect complexities orsubtleties of movement. Accurately defining modes of movementin the ancient past depends on the use of isotopic systems appro-priate for the environment in question, a biogenic substrate thatprovides the required scale of resolution, and a robust estimation ofthe spatially linked isotopic variation on the landscape.


Strontium has long been the isotope of choice for examiningancient human mobility because 87Sr/86Sr is determined by un-derlying bedrock and therefore strongly geographically defined.This value is directly assimilated into co-localized soils, plants, andanimals and is not fractionated in biological systems (Bentley,2006; Blum et al., 2000; Knudson and Price, 2007; Montgomeryet al., 2005). Local 87Sr/86Sr values cannot, however, be estimatedsolely from geological maps, because bioavailable strontium doesnot necessarily mirror bedrock strontium. Sources of variation inbioavailable 87Sr/86Sr include differential weathering of rocks withdifferent 87Sr/86Sr, variably mixed local soils, precipitation levels,and the relative importance of atmospheric Sr inputs, which maychange over time (Hartman and Richards, 2014; Price et al., 2012;Sillen et al., 1998). Recently, 87Sr/86Sr ratios have increasinglybeen paired with other isotopes in order to better constrain thegeographical origins of sampled individuals (Chase et al., 2014;Chenery et al., 2010; Lamb et al., 2014; Müldner et al., 2011; Sealyet al., 1995). This approach can reduce the problem of conver-gence that may occur when different localities in the area of studyexhibit similar strontium isotope ratios.

Characterizing the degree to which different isotopic systemsare sensitive to spatially defined environmental inputs lies at theheart of studies attempting to track human and animal movement.In cases where human mobility is of interest, such an evaluationusually entails analysis of contemporaneous faunal remains, usu-ally the bones and teeth of herbivores. Typically, the isotopic valuesexhibited by these animals are interpreted as reflective of the localisotopic structuring in natural vegetation communities and hy-drological systems, which then provides a point of comparisonagainst which potentially mobile humans can be compared.Increasingly, these values (largely derived from herbivores, butoccassionaly also omnivores such as pigs) are referred to as anisotopic ‘baseline’, a term that has cropped up in the archaeologicalstable isotopic literature with increasing frequency. Unfortunately,many ‘baselines’ violate basic sampling requirements, relying onbut a few isotope values to describe the isotopic composition oflocal food webs. This practice in no way characterizes the potentialrange of isotopic diversity and structuring in local biomes, which inmany environments can be surprisingly large.

Faunal and botanical remains recovered from a particulararchaeological context (e.g., a settlement) potentially provide somemeasure of isotopic variation expressed in the local or mid-rangeenvironment in question, provided that sample sizes areadequate. However, if we wish to investigate mobility in a sophis-ticated way, as a series of controlled moves that can be linked tohuman decisions relating to subsistence or society, we need toconsider isotopic variation on the landscape in a broader contextand control more carefully for spatially defined inputs such aslatitudinal positioning, altitude, precipitation levels, and tempera-ture in research designs. Use of faunal remains requires carefulconsideration of the degree to which the isotopic composition oflocal biomes might shift over the short- and long-term, and thepotential range of movement of sampled animals. While it may bepossible to estimate a priori the type and scale of movement forwild taxa depending on the behavior of modern analogues (e.g.long-distance migration or limited movement due to territorialbehavior), there is a lurking assumption that domesticated animalswill faithfully produce a ‘local’ signal unadulterated by human in-fluences (see, for example, Murphy et al., 2013). However, the iso-topic values of husbanded animals frequently depend quite heavilyupon the type of husbandry strategy enacted on livestock (Balasseand Tresset, 2002; Balasse et al., 2002; Makarewicz, 2014;Makarewicz and Tuross, 2012; Tornero et al., 2013). For example,domesticates herded to distant pastures or foddered with wildplants or agricultural by-products that are isotopically distict from

the seasonally available vegetation or sourced from distant pas-tures are unlikely to provide an entirely ‘local’ signal.

In light of the dynamic intersection between regional climate,local environment, and culturally modified landscapes and re-sources in defining the isotopic composition of natural andanthropogenic biomes, it seems sensible to abandon the term‘isotopic baseline’ in favor of ‘isoscape’. Isoscapes are spatially andtemporally sensitive maps of the distribution of isotopes on thelandscape that are generated through predictive modelling ofisotope-fractionating processes and the use of environmental datasets (Bowen, 2010; West et al., 2010). The strength in isoscapemodelling lies in its capacity to estimate isotopic values in placeswhere observed data are not available, although it is important tokeep in mind that the accuracy and precision of the isoscape indata-poor regions will be less reliable. The approach likely worksbetter for isotopes that are highly sensitive to spatially definedenvironmental inputs. For example, modern isoscapes for deute-rium and d18O in precipitation are well developed, while those forother isotopes are still preliminary (Bowen, 2010;West et al., 2010).

The potential for higher spatial and temporal resolution offeredby isoscapes is a promising analytical direction, but its usefulness isdependent on the resolution of both the isoscape itself and thedistribution of isotopes in bones and teeth. One of the challenges inusing isotope signals recovered from human or animal skeletalremains to reconstruct mobility has to do with the period capturedin the tissue in question, compared with the longevity of the in-dividual. It is possible to trace mobility in early life stages throughisotopic analysis of teeth with different formation times, and also todetect movements that occur later in life by analysing differentskeletal elements with different bone turnover rates (Ericson,1985;Montgomery, 2010; Schroeder et al., 2009; Schweissing and Grupe,2003; Sealy et al., 1995).

The discovery that teeth preserve an isotopic time-seriesderived from enamel and dentin has supported burgeoningresearch investigating dietary change and mobility in wild un-gulates and domesticated livestock exploited by ancient commu-nities (Balasse et al., 2001, 2002; Bendrey et al., 2014; Britton et al.,2009; Makarewicz, 2014; Passey and Cerling, 2002). Hypsodontungulate tooth crowns, in particular, provide an excellent record ofisotopic shifts associated with seasonal changes in environment,diet, and movement over short time spans, from less than a year upto a few years depending on the taxon sampled (Balasse et al., 2002;Passey and Cerling, 2002). Recent work examining isotopic se-quences derived from pig incisors suggests that change in birthseason, diet, and mobility patterns may be detectable in omnivores(Fr�emondeau et al., 2012).

In order to establish if different patterns of dietary change arevisible in an assemblage of archaeological herd animals, thesestudies require direct comparison of isotopic time-series obtainedfrom multiple individuals. This is typically achieved by aligningmultiple isotopic curves at the crown-tooth junction. It is now wellknown, however, that the temporal resolution of intra-tooth iso-topic variation obtained from sequentially sampled teeth is atten-uated by several factors including duration and geometry of tissueformation, and this complicates inter-individual comparisons.Enamel mineralization, for example, is a prolonged process thatinvolves initial enamel formation from the tooth crown towards thecervix, followed by an extended maturation stage in which sec-ondary mineralization occurs at various appositional fronts(Balasse, 2002; Passey and Cerling, 2002; Suga, 1982). This hasimportant implications for interpreting intra-tooth isotopic changeassociated with mobility, or dietary change more broadly, as longermineralization processes will result in under estimations of theamplitude of isotopic variation (Balasse, 2003). In addition, inter-individual comparison of isotopic time-series assumes constant


rates of tooth growth rates and similar tooth geometry within asingle taxon. Recent work has demonstrated, however, that inter-individual isotopic variability expressed in a seasonal cycle isinfluenced by tooth type, size, and geometry, and differences in thetimescales over which tooth growth occurs. Divergence in isotopiccurves may therefore be ontogenic in origin, rather changes in dietor mobility (Balasse et al., 2012; Bendrey et al., 2014; Blaise andBalasse, 2011; Tornero et al., 2013; Towers et al., 2014; Zazzoet al., 2012).

These issues are likely compounded by current samplingmethods, which cross-cut growth fronts by sampling a series ofhorizontal bands that are roughly perpendicular to the primary axisof tooth growth (Balasse, 2003). Differences in the width and depthof sampled bands result in samples containing different amounts ofmaterial from different moments of growth which adds anotherdimension of isotopic complexity. Sampling discrete incrementsthat better reflect biological growth structures remains a primarygoal of studies that use tooth tissues to detect isotopic changesassociated with diet and mobility. The continued development ofthis field is critical to archaeological questions that explore dietarychange and mobility in wild animals, livestock, and humans, as itwill provide a more accurate representation of change over smalltimescales meaningful to human activity. Mathematical trans-formation models developed to better fit isotope data to toothgrowth rate and geometry may produce what seem to be onlymoderate changes on the periodicity of isotopic change visible inthe time series (cf. Balasse et al., 2012. Bendrey et al., 2014; Zazzoet al., 2012), but the modelled information provides informationat a scale that is critical for detecting human decision-making andtherefore has real consequences for better understanding man-agement and mobility decisions.

Bone offers much lower temporal resolution than tooth enameland dentine, as the isotopic consequences of dietary change andspecific movements are ‘averaged out’ through bone remodelling(Hedges et al., 2007). Consequently, detecting distinct patterns ofmovement in human and animal skeletal tissues once tooth for-mation is completed is difficult. However, recent developments inmicro-sampling of bone osteons suggest that capturing smallertimescales within the later portion of the lifetime of an individual ispossible (Scharlotta et al., 2013). Achieving this finer temporalresolution is necessary and critical to answering more refinedquestions relating to the timing and spatial outcome of moves.Indeed, continued development of analytical techniques thatimprove the temporal resolution of isotopic data, down to theimportant scale of seasonal and sub-seasonal variation, enables usto move further away from the concept of an isotopic baseline andexamine human and animal behaviour within a framework of dy-namic seasonal isoscapes. Being able to examine summer andwinter isoscapes, for example, is a significant step forward in thearchaeological study of mobility.

5. Dietary quantification and mixing models

The archaeological record is rich in the discarded remains ofhuman subsistence activities and food waste. Carbonized seeds,animal bones, starch, phytoliths, and molecular-level food residues(e.g., lipids) are recovered from archaeological sites around theworld. However, establishing the precise contribution of thesevarious foodstuffs to past human diets is a difficult, if not impos-sible, task. Various taphonomic and site formation processes,including density-mediated attrition, food processing, and discardpractices heavily modify the remains of food items, leading to dif-ferential representation in the archaeological assemblage. Stableisotope analysis, as a direct measure of diet, may offer a way todetect the consumption of dietary constituents that are otherwise

difficult to discern in the archaeological record. However, stableisotope values currently offer only a semi-quantitative measure ofdietary intake, and detection of the precise contribution of dietarysources to the total mixed diet remains a challenge.

This issue was central to the earliest application of isotope-based dietary tracing to an archaeological question e the spreadof maize agriculture from Mesoamerica into the woodland areas ofeastern North America (Van der Merwe and Vogel, 1978; Vogel andvan der Merwe, 1977). Carbonized plant remains provided directevidence for the use of maize at many settlements, but especially inearlier sites, it was unclear whether maize had already become adietary staple or whether it was consumed only occasionally(Staller et al., 2006). Woodland North America was a C3 environ-ment, and the d13C values of bone collagen from skeletons pre-dating the arrival of maize agriculture measured about �21‰.Maize is a C4 plant, so the skeletons of maize farmers showed moreenriched d13C values, providing direct evidence of maize con-sumption (Van der Merwe and Vogel, 1978; Vogel and van derMerwe, 1977). The most positive values (up to �11‰) clearlyreflect a substantial reliance on maize e but how substantial? Canwe quantify an individual's maize intake based on collagen d13C?Does collagen that exhibits enrichment of 10‰ in d 13C signal twiceas much maize in the diet as collagen enriched by only 5‰?

The question remains unresolved to this day. Mixing modelsmay provide a way of tackling this and similarly important ques-tions, and the use of suchmodels has become increasingly commonover the past decade (e.g., Bocherens et al., 2005, 2006; Coltrain,2009; Coltrain and Janetski, 2013; Coltrain et al., 2004; Newsomeet al., 2004). In their simplest form, mixing models consider twofood sources with clearly separated isotope values. If the isotopevalues of the consumer, the foods ingested, and the extent offractionation in the formation of consumer tissue are known, it ispossible to calculate the proportion of each food in the total diet.Early work employed simple linear mixing models to estimate thecontribution of different sources to consumer diets, but these couldnot estimate variability in sources or accommodate the high di-versity of most human diets.

More recent mixing models, surveyed in Phillips et al. (2014),may better estimate food source contributions in the diets ofancient consumers. These models allow for a greater number ofpotential sources, accommodate varying concentrations of the el-ements of interest (e.g., different percentages of nitrogen in animaland plant food), and consider uncertainties in diet-to-tissue frac-tionation (Erhardt and Bedrick, 2013; Hopkins and Ferguson, 2012;Ogle et al., 2014; Parnell et al., 2010; Ward et al., 2011). Bayesianstatistical frameworks, a standard feature of current mixingmodels, describe the uncertainties in food source and consumerisotope values as well as estimated source contributions, and alsoincorporate prior distributions to futher constrain dietary sources(Phillips et al., 2014). However, a fundamental problemwithmixingmodels remains in that they cannot deal with the complexities ofmetabolic routing, discussed below. Since these are currentlypoorly understood, they cannot at present be incorporated intomixing models. There is therefore a risk that mixing models maygive overly simplistic answers to complex questions.

Mixing models are a powerful exploratory tool that can helpcharacterize dietary structure, but they are by no means a panaceafor dietary source determination and trophic level positioning,particularly for dietary generalists such as humans. It is importantto keep inmind thatmixingmodels are only as good as the estimateof the isotopic composition of foods ingested by the consumer.Models that incorporatemeans and variance of sources, rather thansingle data points, will better describe isotopic structuring withinsource populations and the diet (Semmens et al., 2009). The pre-cision of model-based parameter estimates is also influenced by the


number of samples contributed to each structural level (i.e., con-sumers and sources) (Phillips et al., 2014). In cases where eachsource is represented by only a small number of isotopic values, asis often the case for archaeological data sets, the degree of uncer-tainty in source mean and variance values can be high (Phillipset al., 2014). This presents a challenge for modeling ancient hu-man diets, as the number of plant and animal samples derived fromappropriate archaeological contexts are often too small toadequately characterize the extent of isotopic variation in foodsources. Conversely, modeling source contributions in the humandiet that exhibit similar isotope value and variance, but representvastly different resource pools, present a different type of chal-lenge. For example, cereal cultivars may be isotopically similar tolocally available and exploitable wild grass resources. Dependingon local environmental strategies and degree of human interven-tion with cultivar growth, the isotopic distinction between culti-vated and wild seeds may be minimal, but the contrast betweencultivation and gathering as a subsistence strategy is enormous.

If mixing models are to be used to explore human dietarysources, more detailed attention to the isotopic complexity ofancient food webs is required, and more traditional types of evi-dence for subsistence considered. The integration of data fromzooarchaeological and paleobotanical assemblages, which providea partial, albeit flawed (for reasons discussed above) measure of therelative importance of plant and animal resources exploited byancient human groups, with stable isotope data obtained fromthose assemblages, would help to further constrain dietary sourceparameters. Keeping in mind that we are interested not only in theconstituents that make up the human diet but also strategies usedto procure diets, it may be that the real value in stable isotopicmixing models is not that they provide an exact calculation of di-etary resource contributions to consumers but that they are anexploratory tool with the potential to generate new lines ofarchaeological inquiry and questions.

6. Compound-specific analysis and dietary routing

Compound-specific analysis of body tissues is one of the currentfrontiers of isotopic dietary studies. This approach offers a way ofcircumventing some of the problems of mixing models in the questto estimate the relative contributions of different dietary sources inconsumer diets. Once taken into the body, the major macronutri-ents in the diet (proteins, fats, and carbohydrates) are metabolisedvery differently, entering into different metabolic pathwaysresponsible for, among others, glycolysis, gluconeogenesis, aminoacid synthesis, and fatty acid b eoxidation. All of these biochemicalreactions involve isotopic fractionation. Consequently, the isotopiccompositions of consumer tissues depend on both the isotope ra-tios of nutrients consumed and on metabolic fractionation duringtissue synthesis.

d13C and d15N values obtained from bulk collagen represent anaverage of distinct values derived from single amino acids, thevalues of which reflect the isotopic composition of their respectivesource dietary pools and the fractionation factors involved in theirmetabolism (Hare et al., 1991; Macko et al., 1987). In the case ofcarbon, the d13C values of essential amino acids e supplied by thediet - should directly reflect the values of dietary protein sourcesand exhibit very similar isotope ratios in food and consumer (Corret al., 2005). Non-essential amino acids may also be incorporateddirectly from the diet, especially if this is high in protein, or theircarbon skeletons may be synthesized from a range of macronutri-ents (carbohydrates and lipids as well as proteins). Carbon derivedfrom maize in the diet therefore shows up more strongly in non-essential, compared with essential, amino acids (Fogel andTuross, 2003).

Much more research focused on defining the relationship be-tween metabolic pathways and the isotope values of consumertissues is needed. It remains unclear the extent to which differentpathways operate under different conditions. For example, whatproportion of the carbon skeletons of non-essential amino acids inbone collagen are derived directly from dietary amino acids? Whatproportion are synthesised in the consumer's body, thus probablyincorporating carbon from dietary carbohydrates and fats? Howdoes this change according to the nature of the diet, the balancebetween energy intake and expenditure, whether the individual isgrowing, pregnant, or lactating? The lack of answers to thesequestions imposes major limitations on any attempts to achieveprecise quantitative reconstructions of diet, particularly if onewishes to distinguish energy and protein foods with different car-bon isotopic compositions. This problem is, of course, most acutefor carbon isotopes, given that nitrogen (and sulphur) derives onlyfrom dietary protein.

One way to gain a better understanding of these metabolicpathways and their isotopic consequences is through the analysis oftissues from animals raised on controlled diets. Animals fedspecially manufactured diets inwhich the nutrients of interest havedifferent isotope ratios (e.g., C3 protein/C4 carbohydrate) offer awayto trace the origins of dietary macronutrients in consumer tissues.Such experiments afford insights into factors that might, at firstglance, be thought unlikely to affect isotope patterning but haveturned out to play a significant role, e.g., quantity of fibre in the diet(Howland et al., 2003). Difficult to design and manage, andextremely expensive, such studies have nonetheless yielded veryvaluable results for both bulk tissues (Ambrose and Norr, 1993;Ayliffe et al., 2004; Passey et al., 2005; Sponheimer et al.,2003a,b; Tieszen and Fagre, 1993) and compound-specific work(Howland et al., 2003; Jim et al., 2004, 2006).

Another approach to understanding isotopic information fromindividual amino acids involves mapping d13C and d15N of aminoacids in free-ranging organisms with different diets and/or origi-nating from different environments. Useful patterns have beendocumented: for example, the difference between the d15N valuesof the non-essential amino acid glutamate and the essentialphenylalanine (D15Nglu-phe) is larger in marine compared withterrestrial organisms, and this spacing is transmitted to the tissuesof consumers. This provides a way of estimating marine proteinintake independent of factors that can confound interpretation ofbulk collagen measurements, such as elevated terrestrial d15N incoastal deserts (Naito et al., 2010, 2013a; Styring et al., 2010).Recently, it has been shown to be useful in assessing trophic level(Naito et al., 2013b; Styring et al., 2015), although more work isneeded to assess D15Nglu-phe across different environments. Givenour difficulties in detecting plant foods using isotope methods, thisis a promising development.

Compound-specific analysis is applicable to a wide range ofmolecules, including waxes, lipids, and similar molecules which arestable over long periods of time. Compound-specific stable isotopicanalysis of lipids is most commonly associated with residuesrecovered from unglazed ceramics (Evershed, 2008; Evershed et al.,1999) and has providedmajor insights into the antiquity of dairying(Evershed et al., 2008). Lipids have also been recovered fromarchaeological bones and teeth, especially cholesterol (Stott et al.,1999). Cholesterol has much potential as a palaeodietary proxy, asit is synthesized primarily from dietary fat and carbohydrates.Moreover, it reflects dietary inputs during approximately the lastyear of life, which may be useful for comparison with the longer-term signal obtained from bone (Hedges et al., 2007; see Richardset al., 2007; Corr et al., 2008 for applications of this approach).

The goal of compound-specific work on the body tissues ofconsumers is essentially to obtain greater resolution in dietary


reconstruction. There has already been considerable progress inresolving dietary mixes such as marine/C4 terrestrial, for whichanalysis of bulk tissues yields ambiguous results (Corr et al., 2005)and identifying the consumption of otherwise difficult-to-detectfreshwater foods (Naito et al., 2013a). Ultimately, it may bepossible to track particular foods, e.g., key cultigens.

7. Oxygen and hydrogen isotopes in bone collagen and toothdentin

Oxygen isotopes derived from carbonates and phosphates inbone and tooth apatite are now routinely used to track mobility inhumans and animals, sometimes in conjunction with an indepen-dent, geospatially reactive isotopic system such as strontium (Chaseet al., 2014; Chenery et al., 2010; Lamb et al., 2014; Müldner et al.,2011). The d18O values of animal tissues reflect the isotopiccomposition of oxygen in body water, which is sourced from at-mospheric water, food, and perhaps most crucially, ingested waterderived largely from meteoric waters (Ayliffe and Chivas, 1990;Bryant and Froelich, 1995; Longinelli, 1984; Luz et al., 1984). Asmeteoric waters are influenced by a combination of environmentalinputs including source water, temperature, humidity, altitude, andcontinental positioning (Clark and Fritz, 1997), oxygen isotopes inanimal tissues can potentially delineate long- and short-rangepatterns of movement in environments characterized by high ox-ygen isotopic variability.

There are, however, analytical challenges associated with themeasurement of oxygen isotopes in bone. Structural carbonate isnotoriously susceptible to diagenetic alteration due to the smallcrystal structure of bone hydroxyapatite, which creates a largesurface area for bicarbonate adsorption (Koch et al., 1997). Conse-quently, carbonate d18O values derived frombonemay be unreliable(Koch et al., 1997; Lee-Thorp and van der Merwe,1987, 1991; Quadeet al., 1992; Wang and Cerling, 1994). FTIR spectra of bones can helpestablish whether or not apatite recrystallation in bone hasoccurred, but measurements of crystallinity do not reliably predictthe degree of diagentic alteration of isotopes, particularly oxygen, inbone (Trueman et al., 2008). Different pretreatment protocols usedto remove organic matter and exogenous secondary carbonatesimpose additional isotopic offsets that further remove bone car-bonate d18O values from their original composition (Balasse et al.,2002; Koch et al., 1997). Oxygen isotopes in tooth enamel and inbone phosphate are less likely to suffer diagenetic alteration butmay be affected in some circumstances (Zazzo et al., 2004a,b).

One arena of stable isotope analysis in archaeology that hasgreat potential for expanding our understanding of seasonality,mobility, and the geographic origins of food in ancient societies liesin organic hydrogen and oxygen isotope analyses of proteins.Hydrogen and, to a lesser extent, oxygen isotope analyses of hairand feather keratin are relatively common in ecological studiestracking animal mobility and human travel on continental scales(Hobson et al., 1999; Kelly et al., 2002; O'Brien and Wooller, 2007;Sharp et al., 2003), but remain infrequently used in archaeologicalcontexts, and then primarily as a climate proxy and way to detectthe trade of textiles (Leyden et al., 2006; von Holstein, 2013;Wilsonet al., 2007). Studies using the dD and d18O values of bone collagenand dentine are limited to a relatively small number of modern andarchaeological applications (Crowley, 2014; Cui et al., 2015;Kirsanow et al., 2008; Koon and Tuross, 2013; Leyden et al., 2006;Makarewicz and Tuross, 2012; Topalov et al., 2013; Warinneret al., 2012). Hydrogen and oxygen isotopes are well-suited toarchaeological questions relating to mobility, as the dD and d18Ovalues of collagen are correlated with the isotopic composition ofingested waters (Cormie et al., 1994; Leyden et al., 2006), andorganic oxygen and hydrogen bound to carbon appear to record

principally ingested waters with some contribution from dietaryhydrogen and oxygen (Kirsanow et al., 2008; Reynard and Hedges,2008; Tuross et al., 2008). However, these isotopes remainunderutilized and underexplored, in part because ca. 20% of totalorganic hydrogen in collagen is exchangeable with ambient atmo-spheric moisture (Cormie et al., 1994), and the question of how todeal with hydrogen exchange remains unresolved. Several pre-treatment protocols attempt to control the isotopic compositionof exchangeable hydrogen by isotopically equilibrating proteinswith water vapors of known isotopic composition (Chesson et al.,2009; Kelly et al., 2009; Sauer et al., 2009; Wassenaar andHobson, 2000), although the differences between keratin dDvalues measured with steam equilibriation and those determinedby equilibriation in dessicators at ambient temperature suggeststhe structure of keratinmight be alteredwith steaming (Coplen andQi, 2013). The application of temperature steam-equilibrationtechniques to separate non-exchangeable hydrogen fromexchangeable hydrogen may further reduce the hydrogen isotopicintegrity of already fragile ancient proteins.

The measurement of hydrogen isotopes faces some veryfundamental analytical challenges that must be overcome if dDvalues are to gain currency as a method appropriate for exploringchange in dietary and drinking water sources in animals. First andforemost, the issue of exchangeable hydrogen is a significantchallenge that needs to be solved. In cases where samples areequilibrated with ambient moisture in the laboratory, dD valuesmeasured at different geographic locations or points in time are notreliably comparable because humidity levels, and therefore thehydrogen isotopic of ambient moisture, change according to localenvironmental inputs throughout the year (Meier-Augenstein et al.,2013). Systematic application of corrections that account for frac-tionation during sample preparation and hydrogen exchange withatmospheric moisture would help facilitate intra-laboratory com-parison of dD values (see Meier-Augenstein et al., 2013 for a reviewof the analytical challenges associated with measuring hydrogenisotopes and recommendations for best practices).

Oxygen isotopes in collagen may be less labile, but additionalwork investigating oxygen isotopic exchange with ambient atmo-sphere is required. Measurement of organic oxygen would providea major analytical advantage over d18Ocarbonate in that it wouldallow analysts to circumvent the challenges relating to diagenesisin using bone carbonate d18O and permit direct comparison of d18Ovalues from tooth dentine andmetabolically active bone collagen toprovide insights into age-related shifts in human and animal oxy-gen isotopic composition. Previous research indicates some, but notcomplete, covariance between dD and d18O in various tissues fromthe same individual, which suggests that drinking water and foodcontribute differentially to oxygen and hydrogen in proteinaceoustissues (Ehleringer et al., 2008; Kirsanow et al., 2008; Podlesaket al., 2008; Tuross et al., 2008). Trophic level and mass-specificmetabolic rates related to body size appear to influence thehydrogen isotopic composition of animal collagens, although thecontribution of drinking water dD to the isotopic composition ofcollagen is still preserved (Birchall et al., 2005; Kirsanow andTuross, 2011; Pietsch et al., 2011; Topalov et al., 2013). Simulta-neous measurement of dD and d18O may therefore help disentanglethe isotopic composition of diet and drinking water in consumerdiets (Kirsanow et al., 2008).

At present, this potentially rewarding line of research remainsexperimental and much additional work is needed to further testthe utility of hydrogen and organic oxygen in ancient proteins. Inparticular, the role of diagenesis in influencing dD and d18O values,accounting for exchangeable hydrogen in measurements, themetabolic fractionation factors involved with the incorporation ofoxygen and hydrogen into organic matrices, and the precise


contribution of dietary hydrogen and oxygen relative to ingestedwater all have yet to be established.

8. Conclusion

Although the halcyon days when stable isotope analyses werehailed by archaeologists as a straightforward solution for recon-structing ancient diets may long be over (see Sillen et al., 1989),these approaches have made and continue to make importantnovel contributions to our understanding of ancient societies. Thispaper emphasizes aspects of the field that we believe warrantcloser attention, including dietary quantification, molecular-levelwork, and exploration of less well-understood isotopes. These is-sues notwithstanding, existing stable isotope approaches still offera powerful, at least semi-quantitative, method of investigating dietsand mobility. At a minimum, it is possible to use stable isotopeswithin a comparative framework in order to assess broad similar-ities and differences in diets through time and space, and inferwhether isotopically distinct categories of foods played a substan-tial role in the diet, or not. However, this is achievable only whenenvironmental factors that also influence the stable isotopiccomposition of dietary constituents are controlled for.

One of the core tenets of hypothesis-driven anthropologicalarchaeology is that it draws from several lines of evidence toaddress particular problems and test models. However, stable iso-topic analyses are too frequently conducted and data interpreted inisolation from other archaeological data sets. This partitioning isnot unique to stable isotope analysis, and the diversification andproliferation of sub-disciplines within archaeological science overthe past decademeans that it is increasingly difficult to ‘know it all’.Stable isotopic approaches would, however, benefit greatly from acloser working relationship with those approaches that examinedietary change through analyses of subsistence remains commonlyfound at archaeological sites, including fauna, carbonized seeds,phytoliths, starch, and other food residues. By more explicitlyconsidering the subsistence strategies employed by people in thepast and the potential dietary outcomes of those strategies, ascharacterized by zooarchaeological and paleoethnobotanical ana-lyses, the isotopic values expressed in human calcified tissues arebetter contexualized and a more holistic understanding of dietachieved. Other novel techniques that directly document dietaryintake, such as characterization of proteins and plant microfossilspreserved in dental calculus (Henry et al., 2011, 2014; Warinneret al., 2014), will also complement and strengthen the perspec-tives afforded by stable isotopic data sets.

The astronomical growth and routinization of stable isotopeanalysis in archaeology points to a healthy development of thefield, but it may be worth cautioning against over-exuberance. Therelatively low cost of analysis, particularly for d13C and d15N, ismaking these approaches much more accessible. Ironically, at thesame time, the increasing size and complexity of the stable isotopeliterature makes it more difficult for archaeologists to assess whatstable isotopes are and are not likely to be able to contribute in agiven project. The most successful stable isotope studies aredesigned in such a way as to achieve a good fit between thearchaeological question and the capabilities of the stable isotopetracers. This requires an intimate understanding of both archae-ology and isotopes, or very close collaboration between researcherswith complementary expertise. Although a collaborative path isprobably the most frequently chosen route, often to good effect,stable isotopes and archaeology are too frequently viewed andpracticed as domains of specialist knowledge. We need to aim forbetter integration and ensure that both archaeologists and stableisotope analysts have at least a basic familiarity with the questions,techniques, and challenges inherent to each discipline at the outset

of a research project. Like lithic, faunal, or paleobotanical analyses,the results from archaeological stable isotope research are only asgood as the research design and understanding of the processesinvolved. There is much to be done. We look forward to the ad-vances in stable isotope approaches that will certainly be made inthe years to come.

Acknowledgements

Makarewicz would like to acknowledge the Jordanian Depart-ment of Antiquities, whose local sponsorship of the 2013 WorldArchaeology Congress in Jordan led to fruitful discussions withJudith Sealy and Robin Torrence. CM would also like to thankRichard Klein for his support during her time at Stanford University.During the writing of this paper, Sealy was supported by the SouthAfrican Research Chairs Initiative of the Department of Science andTechnology and the National Research Foundation of South Africa.Opinions, findings, conclusions, and recommendations expressedin this paper are those of the authors, and the NRF does not acceptany liability in this regard.

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