Human dietary assessment in the Pre-colonial Lesser Antilles: New stable isotope evidence from Lavoutte, Saint Lucia

Journal of Archaeological Science: Reports 5 (2016) 168–180

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Human dietary assessment in the Pre-colonial Lesser Antilles: New stable isotope evidence from Lavoutte, Saint Lucia Jason E. Laffoon a,b,⁎, Menno L.P. Hoogland a, Gareth R. Davies b, Corinne L. Hofman a a b

Faculty of Archaeology, Leiden University, 2333 CC, Leiden, The Netherlands Faculty of Earth and Life Sciences, VU University Amsterdam, 1081 HV, Amsterdam, The Netherlands

a r t i c l e

i n f o

Article history: Received 18 August 2015 Received in revised form 28 October 2015 Accepted 14 November 2015 Available online 5 December 2015 Keywords: Diet Stable isotope analysis Carbon Nitrogen Caribbean archeology Ceramic age

a b s t r a c t Dietary assessment of Late Ceramic Age inhabitants (~AD 1200–1500) from Lavoutte, Saint Lucia, Lesser Antilles was undertaken on human skeletal remains using stable carbon and nitrogen isotope analysis of bone collagen (δ13Cco and δ15N), and bone and enamel bioapatite (δ13Cap and δ13Cen). The isotope data were interpreted in the context of regional food webs, intra-population dietary variation was assessed relative to individual demographic variables and the results were compared to contemporaneous populations at multiple scales. Moderately enriched δ13Cco and elevated δ15N point to substantial contributions from both non-marine and marine protein sources, including nitrogen-enriched pelagic resources. The lack of correlations between δ13Cco and δ13Cap (or δ13Cen) suggests distinct isotopic differences between protein and energy sources. The smaller range and variance of δ13Cco and δ15N values relative to δ13Cap and Δ13Cap-co indicate greater inter-individual heterogeneity in dietary energy sources relative to protein sources. Intra-population dietary variation was not, however, correlated with either age or sex, consistent with communal-based food consumption practices. From a broader perspective, the collagen isotope results are comparable with several islands in the Lesser Antilles but are distinct from others, indicating a large degree of regional variation in dietary protein sources, while the bone (and enamel) apatite results are more variable and overlap with many islands in both the Greater and Lesser Antilles indicating wider variation in average (whole) diets. The relative enrichment in δ13Cap and higher Δ13Cap-co values are strongly indicative that C4 (e.g., maize) or CAM plants were important dietary components. Overall, the isotopic evidence suggests that the Lavoutte population consumed mixed diets including substantial contributions of both C3 and C4/CAM plant resources, as well as terrestrial and marine protein sources. © 2015 Elsevier Ltd. All rights reserved.

1. Introduction Owing to the central role of food in human biology, culture, and society, dietary and subsistence reconstruction have long been and continue to be important foci of archeological investigations (Metheny and Beaudry, 2015; deFrance, 2009; Parker-Pearson, 2003; Roosevelt, 1987; Ross, 1987; Twiss, 2007; and references therein). Dietary patterns not only shed light on aspects of food procurement and consumption but can also provide indirect evidence concerning many other aspects of past human societies such as economic systems, ecology, adaptation, exchange, and health conditions. A wide range of evidence has been used in traditional archeological models of dietary reconstruction including ethnographic, documentary, architectural, iconographic, and material. In particular, analysis of botanical and faunal remains has contributed greatly to our current understandings of past human dietary practices. These data sources, however, possess inherent limitations in

⁎ Corresponding author at: Faculty of Archaeology, Leiden University, 2333 CC Leiden, The Netherlands. E-mail address: [email protected] (J.E. Laffoon).

http://dx.doi.org/10.1016/j.jasrep.2015.11.020 2352-409X/© 2015 Elsevier Ltd. All rights reserved.

that they generally reflect an accumulation of remains from diverse activities spanning long periods of time and are heavily biased by differential preservation and taphonomic factors. The development of stable carbon and nitrogen isotope analysis over the last few decades has revolutionized our understanding of human paleodiet by permitting investigations of previously elusive aspects of past dietary patterns such as relative contributions of different resource types to individual diets; intra-population variation related to sex, age, status, or chronology; intra-individual changes in diet over time; and inter-population differences in food-ways just to name a few (reviewed in Ambrose and Krigbaum, 2003; Katzenberg, 2008; Lee-Thorp, 2008). Stable isotope analysis of human skeletal remains is a wellestablished method for dietary reconstruction in the Caribbean region (see reviews in Pestle, 2013a; Stokes, 2005). In fact, early important contributions to the theoretical and methodological development of stable isotope dietary studies derive from studies using archeological skeletal materials from the circum-Caribbean region (Keegan, 1985; Keegan and DeNiro, 1988; Schoeninger et al., 1983; Van Klinken, 1991). Large-scale stable isotope studies of various insular Caribbean populations have revealed spatial patterns in dietary variation linked to geographic and ecological parameters (Stokes, 1998, 2005). In recent

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years, a large number of paleodiet isotope studies have been conducted in the region (Antón, 2008; Buhay et al., 2013; Healy et al., 2013; Krigbaum et al., 2013; Laffoon and de Vos, 2011; Laffoon et al., 2013; Norr, 2002; Pestle, 2010a, 2010b, 2013a; Pestle and Colvard, 2012) and these have expanded not only the size of the overall isotopic datasets but also have shed further light on the spatial, chronological, and social aspects of dietary variation in the pre-colonial Caribbean. Specifically, these and related studies have revealed a high degree of regional dietary diversity, particularly in the relative contributions of animal protein (terrestrial/aquatic) and to a lesser extent of plant energy (C3/C4) resources to indigenous diets. This study contributes to this line of research by focusing on an understudied area of the Caribbean, namely the island of St. Lucia and the Windward Islands archipelago. Results of previous research focusing on osteological, mortuary, ecological and material cultural analyses of the Lavoutte site, including strontium isotope analyses of human burials to investigate natal origins, have been published elsewhere (Hofman et al., 2012). In this study, we expand on those initial results through the application of multiple stable isotope analyses of human bones and teeth from the Lavoutte burial population. The aims of this study are to 1) shed light on the dietary patterns of the ancient inhabitants of Lavoutte, Saint Lucia; 2) assess the isotope results in reference to food web data; and 3) explore dietary patterning at multiple scales from intra-societal (relative to sex, age, origins), to archipelagic (Lesser Antilles), to macro-regional (broader Caribbean). 2. Archeological context The site of Anse Lavoutte (hereafter Lavoutte) is located on the Atlantic (eastern) coast near the northern tip of Saint Lucia (Fig. 1). The site was first discovered in the late 1950s and is well-known for the so-called ‘Lady of Lavoutte’, a large distinctive ceramic figurine of a seated female. Excavations carried out by the University of Florida in cooperation with the Saint Lucia Archaeological and Historical Society (SLAHS) in the 1960s uncovered substantial quantities of Suazan Troumassoid ceramics, including numerous fragments of ceramic figurines (Bullen and Bullen, 1970). Based on these remains and a single radiocarbon date the site was proposed to have been a large settlement site occupied during the Late Ceramic Age (~ AD 800–1500) and

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possibly even a ‘Carib ceremonial centre’ (Bullen and Bullen, 1970:75). The presence of artifacts which bear a striking resemblance to ‘Taíno’ material culture suggests interactions with contemporaneous populations from the Greater Antilles (Allaire, 1999). Subsequent small-scale excavations at the site in the 1980s were conducted by a team from the University of Vienna, which recovered the remains of three human burials and additional cultural remains (Fabrizii-Reuer and Reuer, 2005). In recent years, both natural (hurricanes, coastal erosion) and human (hotel construction, increased tourist activity) factors have contributed to large scale alteration of the local landscape (Hofman et al., 2012; Siegel et al., 2013). Routine monitoring of the site in 2009 revealed numerous exposed human burials and an initial impact assessment revealed that the site was under serious and immediate threat of destruction owing to its location, proximity to the coast, elevation, and the increased use of the beach by both locals and tourists as a thoroughfare for horse and vehicular traffic (Hofman and Branford, 2011). In response to the urgent need for archeological remediation, an international team of researchers comprised of the Caribbean Research Group from Leiden University, the Florida Museum of Natural History, the SLAHS, the Saint Lucia Government, and local volunteers carried out large-scale rescue excavations in 2009 and 2010 (Hofman et al., 2012). These excavations recovered significant quantities of material remains including ceramics, lithic raw materials and tools, shell, coral, and animal bone, and revealed the presence of numerous features including post-holes from houses and other built structures in addition to more than 48 burials. Preliminary paleo-environmental reconstruction of the site's surroundings (Hofman et al., 2012) indicated the presence of xerophytic scrub or low woodland on the surrounding slopes including, amongst other plants, agave and several varieties of cacti. Multiple small freshwater channels drain into the bay near the site and were the locations of small mangrove swamps, remnants of which persist today. The site sits on a promontory overlooking the partially sheltered bay of Casen-Bas and is surrounded on three sides by steeply sloping hills. Dispersed patches of coral reef can be found in the immediate surroundings, and the there is a sharp drop off in the sea floor just outside of the bay. These combinations of conditions would have provided ready access to a wide range of environments and food resources in the immediate vicinity of the settlement.

Fig. 1. Map of the Caribbean with inset of Saint Lucia and location of the Lavoutte site.

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A prominent focus of previous multi-disciplinary studies at Lavoutte is bioarcheological research of the burial population including mortuary, osteological, dental, radiocarbon, and biogeochemical analysis (Hofman et al., 2012; Laffoon, 2012; Mickleburgh, 2013; Weston, 2011). Carbon-14 dating of human bone material from twelve separate individuals from Lavoutte produced median calibrated dates ranging from circa AD 1250 to 1500 and confirms that the burial population dates to the terminal Late Ceramic Age (Hofman et al., 2012). Results of the osteological analysis have also revealed important aspects pertaining to demography, health and disease (Weston, 2011). Previous isotope research has also included strontium isotope analysis of human and faunal remains and these results indicate that most individuals have 87 Sr/86Sr signals consistent with a local origin but that two individuals possess nonlocal Sr isotope signals indicating nonlocal origins (Hofman et al., 2012; Laffoon, 2012). The Lavoutte burial population comprises the largest and most intensively researched pre-colonial skeletal assemblage ever recovered from the Windward Islands (the islands of the southern Lesser Antilles, defined here as including Dominica, Martinique, St. Lucia, St. Vincent and the Grenadines, and Grenada). Owing to a combination of poor preservation and the fact that rescue excavations were focused on recovery of imminently threatened burial features, no botanical remains and relatively few faunal remains have been recovered from the site, hampering efforts to assess many aspects of pre-colonial lifeways, such as dietary and subsistence practices. Recovered faunal materials include marine and terrestrial shell, and animal bone but these have yet to be systematically subjected to zooarcheological analysis. Stable isotope analyses of the Lavoutte skeletal remains, not only affords the opportunity to address these lacunae but also represents the first large-scale stable isotope study of St. Lucian archeological collections and provides important comparative data for the Windward Islands and Lesser Antilles, both understudied regions of the Caribbean. 3. Stable isotopes and dietary assessment Dietary assessment based on stable isotope analyses is based on the principle that the isotopic composition of dietary resources is reflected in the tissues of consumers. The stable isotope method was first applied in archeology to track the intensification of maize agriculture in the eastern woodlands of North America (van der Merwe and Vogel, 1978; Vogel and van de Merwe, 1977). Subsequent research focused on quantifying trophic level enrichment and dietary patterning in different ecosystems (DeNiro and Epstein, 1978, 1981) and provided for a more rigorous foundation for stable isotope dietary reconstruction (Schoeninger and DeNiro, 1984; Schoeninger et al., 1983). Methodical and theoretical developments in stable isotope studies have continued in subsequent decades to such an extent that they have become a routine and widely utilized tool in archeological research (Ambrose and Krigbaum, 2003; Katzenberg, 2008; Lee-Thorp, 2008). One of the most significant advances in this regard is a number of controlled feeding studies that have greatly enhanced our knowledge of the relationships between the isotopic values of dietary constituents and various consumer tissues and the preferential routing of certain dietary macronutrients to different tissues during biosynthesis (Ambrose and Norr, 1993; Hare et al., 1991; Howland et al., 2003; Jim et al., 2006; Tieszen and Fagre, 1993; Warinner and Tuross, 2009). These studies generally confirmed the routing model originally proposed by Krueger and Sullivan (1984) whereby, under conditions of sufficient protein consumption (Schwarcz, 2000), collagen carbon isotopes (δ13Cco) will primarily reflect dietary protein sources, while apatite carbon isotopes (δ13Cap) better reflect the average of the whole diet (carbohydrates, fats, and proteins). Recent studies have argued that ~ 75% of carbon atoms in bone collagen derive from dietary protein sources (Fernandes et al., 2012; Froehle et al., 2010), and owing to the fact that animal flesh contains much higher concentrations of protein than plants by mass, δ13Cco is a reliable indicator of consumed animal protein

δ13C amongst omnivores, while δ13Cap is generally a more sensitive indicator of plant δ13C (Harrison and Katzenberg, 2003). Variation in 13C of plants is conditioned by a wide range of factors but primarily by different photosynthetic pathways (Bender, 1971; O'Leary, 1981; Smith and Epstein, 1971). The vast majority of plants, including most plants consumed by humans, are C3 plants with average δ13C of −26.5‰ but a small minority, mostly represented by sedges and tropical grasses, are C4 plants with average δ13C of −12.5‰ (Smith and Epstein, 1971). A third pathway, known as CAM, is used by succulents which possess moderately enriched δ13C values that can overlap with those of C4 plants (Bender et al., 1973). In New World contexts, maize was the most common and economically important C4 crop and likely the only C4 (or CAM) plant that would have been widely utilized as a staple crop. Carbon isotopes also vary between terrestrial and aquatic (marine and freshwater) ecosystems owing to differences in the primary source of carbon at the base of the food web (Chisholm et al., 1982; DeNiro and Epstein, 1978; Schoeninger and DeNiro, 1984; Schoeninger et al., 1983), and the greater number of trophic levels in the latter. In general, marine organisms possess higher (more enriched) δ13C than terrestrial organisms. Nitrogen isotopes (δ15N) in plants average ~ 3‰, although certain plants such as legumes which directly fix nitrogen tend to have lower values around 0‰, and in animals δ15N is enriched stepwise between each trophic level in a food chain (DeNiro and Epstein, 1978; Minigawa and Wada, 1984). The patterning of δ15N within a particular ecosystem is such that plants possess the lowest values, while herbivores, omnivores, and carnivores possess increasingly higher values. Marine ecosystems generally possess higher δ15N as marine plants have higher δ15N than terrestrial plants, averaging around 7‰, and there are often more trophic levels in marine ecosystems (DeNiro and Epstein, 1978; Schoeninger et al., 1983). The combination of both higher δ15N and less negative δ13C values in marine organisms relative to terrestrial ones are such that coupled isotope systematics can be used to infer the relative contributions of marine and terrestrial protein sources to human diet based on analyses of bone (or dentine) collagen (Hedges and Reynard, 2007; Richards et al., 2003). Some ecological contexts, however, are exceptions to these general patterns of isotopic variation. These include tropical reef ecosystems based on nitrogen-fixing bluegreen algae that have much lower basal δ15N isotope values (~ 0‰) than other marine ecosystems (Capone and Carpenter, 1982; Keegan and DeNiro, 1988; Schoeninger et al., 1983; Schoeninger and DeNiro, 1984). Sea grasses also display less negative δ13C values that overlap with terrestrial C4 plants potentially further complicating attempts to distinguish between marine and C4 consumption (Keegan and DeNiro, 1988; Schoeninger and DeNiro, 1984). In contexts where these combinations of different resources with overlapping isotope ranges exist, some researchers have utilized differences between collagen and apatite δ13C values (Δ13Cap-col) to attempt to disentangle these dietary complexities (e.g., Ambrose et al., 1997; Harrison and Katzenberg, 2003; Lee-Thorp et al., 1989; Norr, 2002; Stokes, 2005). Such approaches are based on observations that for monoisotopic diets, δ13Cco is enriched relative to δ13Cdiet by ~ 5‰ while δ13Cap is enriched relative to δ13Cdiet by ~9.5‰, leading to Δ13Cap-col of ~4.5‰ (Ambrose and Norr, 1993; Tieszen and Fagre, 1993). Individuals with Δ13Cap-col substantially deviating from this value indicate mixed diets with different δ13C for the protein and energy components. For example, Δ13Cap-col less than ~3‰ have been interpreted as indicative of diets with C3 energy and C4/marine protein (Ambrose et al., 1997; Harrison and Katzenberg, 2003; Iacumin et al., 1996; Krueger and Sullivan, 1984; Lee-Thorp et al., 1989). This approach has been criticized by Kellner and Schoeninger (2007) who have proposed that it is more informative to use bivariate plots of comparison of δ13Cco and δ13Cap relative to regression lines representing different dietary combinations. The model's lack of discrimination between C4 and marine diets (Froehle et al., 2010; Kellner and Schoeninger, 2007), however, led to the development of multivariate models which incorporate δ15N, in addition to δ13Cco and δ13Cap (Froehle et al., 2012).

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4. Pre-colonial Antillean diets The general consensus is that Late Ceramic Age Antilleans would have had horticultural based subsistence strategies involving some combination of slash and burn land clearance, house gardens containing domesticated and managed plants, gathering of wild plants, forest hunting, and frequent fishing and shellfish collecting (Newsom and Wing, 2004; Rouse, 1992; Wilson, 2007). Although the pre-colonial Lesser Antilles (and the Windward Islands particularly) were poor in terrestrial faunal diversity, they possess rich and varied ecosystems for the exploitation of forest, tidal, mangrove, reef, and pelagic resources (Newsom and Wing, 2004; Woods and Sergile, 2001). Rice rats (Oryzomyini) represent one of the few widespread native mammals that were hunted and possibly managed, while dogs were the only common domesticated animal. The only other clearly domesticated animal species was the guinea pig (Cavia porcellus) but their skeletal remains do not occur in abundance, display a very spotty distribution, and are fairly uncommon in the Lesser Antilles overall (LeFebvre and deFrance, 2014). Other mainland taxa that were clearly introduced or translocated to the islands from the mainland by Amerindians in pre-colonial times include agouti (Dasyprocta sp.), armadillo (Dasypus sp.), and opossum (Didelphis sp.), and the geographic distribution of these three translocated species was restricted with most documented cases deriving from the Windward Islands (Giovas et al., 2012; Newsom and Wing, 2004; Wing, 2012). Owing to issues of preservation, which are generally poor in the tropics, macro-botanical remains are very sparse within the Caribbean archeological record (Newsom and Pearsall, 2003; Newsom and Wing, 2004). Therefore, zooarcheological analysis has been one of the primary sources of information concerning human paleodiet (Carlson, 1999; Giovas, 2013; Grouard, 2001; Newsom and Wing, 2004; Steadman and Jones, 2006; Steadman and Stokes, 2002; Wing, 2001; reviewed in deFrance, 2013). Although highly valuable, particularly for providing essential data concerning the range and types of faunal resources exploited by ancient peoples, the zooarcheological record is biased in several important regards that limit certain important aspects of paleodietary reconstruction. For example, there is often a strong preservation bias for shell remains such that they are generally by far the largest recovered components of faunal assemblages. Additionally, quantitative and comparative assessments of faunal assemblages are further hampered by variable research designs, collection strategies, and sieving techniques. Most importantly, refuse or midden deposits, from which most macroscopic evidence derives, represent cumulative records of various activities relating to many decades or even generations of discard, and therefore are too course-grained to permit analysis of dietary patterns amongst or between individuals or social groups. Early stable isotope studies of Caribbean paleodiet revealed that Bahamian individuals had much lower δ15N values than would be expected for a population that was heavily reliant on marine resources (Keegan and DeNiro, 1988; Schoeninger et al., 1983). This pattern has been attributed in part to nitrogen fixation at the base of tropical reef ecosystems that characterize much of the Caribbean, such that many shellfish and reef fish species have lower than expected δ15N values that overlap substantially with terrestrial animals (Keegan and DeNiro, 1988; Schoeninger et al., 1983). This lack of discrimination in δ15N values between terrestrial and reef resources complicates the interpretation of stable isotope data for paleodietary reconstruction in the Caribbean. Additionally, owing to the presence and likely consumption of both marine resources and C4 plants (both enriched in 13C) in various cultural contexts, including the pre-colonial Caribbean, and the aforementioned complexities of 15N variation, most stable isotope studies in the region no longer rely solely on collagen as a sampling material and instead also incorporate carbon isotope analysis of bone apatite. Combined human skeletal stable isotope results are most appropriately interpreted relative to regionally specific food web data (Fig. 2).

Fig. 2. Coupled carbon–nitrogen isotope ecology of the circum-Caribbean region (adapted from data available inKeegan and DeNiro, 1988; Norr, 2002; Pestle, 2010a; Schoeninger and DeNiro, 1984; Stokes, 1998; Van Klinken, 1991).

Bivariate plots of Antillean collagen isotope results cluster geographically with relatively little overlap in the ranges of isotope values at the scale of archipelagoes (Greater vs. Lesser Antilles) and in many cases even at smaller scales (individual islands or sites). Furthermore, with the possible exception of Punta Candelero, Puerto Rico, where a temporal shift in diet has been reported (Pestle, 2010a; Pestle, 2013b), these spatial dietary patterns seem to persist over long periods of time. Whether this chronological consistency in stable isotope results, and presumably dietary practices, extends into earlier time periods is difficult to assess owing to a general lack of data from Pre-Ceramic (Archaic Age) contexts in the circum-Caribbean, although recent studies have begun to address this deficiency (Buhay et al., 2013; Chinique de Armas et al., 2015; Mickleburgh and Laffoon, in press). Overall, the available stable isotope evidence indicates a high degree of temporal conservatism in dietary behavior (Stokes, 1998) that contrasts with zooarcheological (Carlson and Keegan, 2004; Fitzpatrick et al., 2008) and dental anthropological (Mickleburgh, 2013, 2014) evidence for changes to human food-ways during this period, and the rather largescale cultural, social, environmental, and climatic changes that occurred throughout the Ceramic Age (~2500–500 BP) (Burney et al., 1994; Curet and Oliver, 1998; Curtis et al., 2001; Hodell et al., 1991; Hofman, 2013; Keegan, 2000; Lane et al., 2011; Rouse, 1992; Siegel, 2010). One possible explanation for this consistency is that there were strong ecological or biogeographical factors conditioning pre-colonial dietary practices such that contemporaneous, culturally similar populations from different islands had distinctive diets and that these distinctions persisted for several centuries (Newsom and Wing, 2004; Stokes, 1998). Alternatively, the lack of stable isotope evidence (from human skeletal remains) for chronological change may reflect an inherent limitation of stable isotope data which primarily reflects large-scale changes in the relative proportions of specific food groups, and thus is unlikely to detect changes in dietary components between resources with similar isotope values or variations in food preparation, processing, and consumption. In terms of dietary assessment, results of stable isotope analyses in addition to starch grain analyses of residues on ceramics, lithics, and in human dental calculus have called into question several aspects of the traditional model of pre-colonial subsistence in the Caribbean (Berman and Pearsall, 2000, 2008; Chinique de Armas et al., 2015; Mickleburgh and Pagán Jiménez, 2012; Pagán-Jiménez, 2011; Pagán Jiménez, 2013; Pagán-Jiménez and Oliver, 2008; Pagán-Jiménez and Rodríguez Ramos, 2007; Pearsall, 2002). First, overall convergence of multiple lines of evidence indicates that most, if not all, Ceramic Age communities relied on a broad spectrum of plant and animal resources. In contrast to previous models, however, which emphasized the

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ubiquitous reliance on marine protein, closer examination of regional patterning of isotope data indicates a substantial degree of variation in marine orientation and that at some sites in the Greater Antilles, particularly on Puerto Rico, freshwater and/or terrestrial protein sources may have been as, if not more, important (Norr, 2002; Pestle, 2010a; Stokes, 1998; Stokes, 2005). Secondly, root crops in general (and manioc in particular) have long been considered the primary staple of Amerindian plant economies in the insular Caribbean, while maize was generally considered to have been a minor or supplemental food source (Newsom and Pearsall, 2003; Newsom and Wing, 2004; Rouse, 1992). Recent starch grain studies have revealed the presence of maize (Zea mays) starch grains in a wide range of substrates (ceramics; lithic, coral and shell tools; and human dental calculus) at a large number of pre-colonial sites with a broad geographic distribution in the circumCaribbean (summarized in, Pagán Jiménez, 2013). Evidence of maize consumption has even been recovered from Archaic Age individuals that were previously thought to predate the arrival of maize to the insular Caribbean (Chinique de Armas et al., 2015; Mickleburgh and Pagán Jiménez, 2012; Pagán-Jiménez et al., 2015). Importantly, in terms of overall abundance and distribution, maize starch grains are just as, if not more, widespread and ubiquitous as those of roots crops (Pagán Jiménez, 2013). Stable isotope evidence has shed further light on questions concerning maize consumption in the pre-colonial Antilles. More specifically, elevated bone apatite δ13Cap and enamel δ13Cen results and high Δ13C ap-co values (Healy et al., 2013; Laffoon et al., 2013; Pestle, 2010a; Stokes, 1998) provide evidence that average whole diets (including carbohydrates, lipids, and proteins) amongst some Caribbean populations were enriched in 13C. Although other C4 and CAM plants, such as amaranths and pineapple, were available in the pre-colonial Caribbean (Newsom and Wing, 2004), maize would likely have been the only plant to have been consumed in sufficient quantities to elevate δ13C of skeletal tissues representing several years or decades of consumption patterns depending on the tissue type. Interestingly, compared to the relatively pronounced spatial clustering of collagen δ 13C and δ 15N values, δ13 Cap values tend to display larger ranges and more overlap between populations, likely indicating that the relative contributions of different plant resources (C 3/CAM/C4 ) were less variable between populations and more variable within them, compared to dietary protein. Additionally, Δ13Cap-co values also display pronounced spatial patterning and are generally higher (N 4‰) in the Greater Antilles compared to the Lesser Antilles (b 4‰), which is consistent with greater reliance on marine resources amongst the latter (Stokes, 1998). In summary, although considerable progress has been made in terms of our understandings of pre-colonial Caribbean diet and subsistence strategies, several important gaps in our knowledge persist. Traditionally, reconstructions of indigenous Caribbean food-ways have been overly reliant on zooarcheological and ethnohistorical evidence, both of which have multiple inherent biases that limit research into social aspects of human diet such as individual variation and are not readily amenable to detailed comparative analyses. Furthermore, region-wide generalizations are often made on the basis of relatively few case studies or samples. In terms of stable isotope evidence in particular, to our knowledge there is no available data from St. Lucia (except a single individual reported in Stokes, 1998), and only one previous study based on the analyses of 14 individuals from Grand Bay, Carriacou has been conducted in the entire Windward Islands archipelago (Krigbaum et al., 2013). This lack of data obviously hampers efforts to better assess smaller scale (inter-island or intra-island) spatial variation or chronological changes in paleodiet. Within the Lesser Antilles, the largest published stable isotope data set (n = 42) derives from the site of Anse a la Gourde, Guadeloupe (Laffoon and de Vos, 2011; Stokes, 1998) and as its burial population is broadly contemporaneous with that of Lavoutte, it provides a basis for comparative analysis and explorations of Late Ceramic age dietary diversity in this region.

5. Materials and methods Sampling of bone material from Lavoutte focused on ribs or long bone fragments. Pieces of cortical bone were mechanically cleaned using a diamond-tipped rotary blade attached to a hand-held drill on low speed to remove residual soil, encrustations, and cancellous bone. Bone fragments weighing ca. 0.5 g were broken into a coarse powder using a manual crushing press. For collagen isotope analysis, sample processing followed a modified version (Ambrose, 1990; Brown et al., 1988) of the Longin (1971) method. Bone particles were placed in pre-cleaned vials and demineralized with 0.6 M hydrochloric acid (HCl) at 4 °C for 5–10 days with the acid being refreshed every 48 h. Afterwards, the solutions were centrifuged, the acid was decanted, and the samples were rinsed to neutral pH in DDI-H2O (distilled, deionized water). Subsequently, 0.125 M sodium hydroxide (NaOH) was added to remove humic acids and other possible organic contaminants, and after 20 h the samples were centrifuged and rinsed with DDI-H2O. Collagen samples were then gelatinized in 0.001 M HCl at 80 °C for 24 h, purified with Ezee filters (Elkay©), condensed, and lyophilized (freeze-dried). For apatite isotope analysis, sample processing followed a modified version of the protocol reported by Bocherens et al. (2011). Fragments of bone or enamel were mechanically cleaned and rinsed with DDI-H2O, placed into pre-cleaned vials and chemically oxidized for 4 h in 2.5% sodium hypochlorite (NaOCl). Samples were then vortexed, centrifuged, decanted, and rinsed to neutral with DDI-H2O. Then 0.1 M acetic acid (CH3COOH) was added to remove secondary carbonates and after 4 h these were vortexed, centrifuged, decanted, rinsed, air-dried, condensed and freeze dried. Measurements of isotopic compositions of bone collagen, and bone and enamel apatite samples were conducted in the Stable Isotope Lab, Faculty of Earth and Life Sciences, VU University, Amsterdam. Collagen samples were analyzed on a ThermoQuest IRMS Delta XP plus interfaced with a Flash elemental analyzer after combustion to produce CO2 and N2 gases that were purified via GC column using He as the carrier gas. International standards USGS40 and USGS 41, and IAEA310(A) and IAEA-NO3 were used for sample calibration for δ13C and δ15N isotope analyses respectively. Bone and enamel apatite samples were analyzed on a Finnigan DeltaPlus IRMS, following reaction of the sample with orthophosphoric acid (H3PO4) [100%] for 24 h at 45 °C and isolation of the produced carbon dioxide (CO2), with a Gasbench II universal automated interface. Long term reproducibility of the international reference material (NBS19) for δ13C is b 0.1‰. Carbon and nitrogen isotope results are reported in the δ notation, in parts per thousand (‰) relative to the international PDB and AIR standards respectively. Typical analytical uncertainty for both collagen δ13C and δ15N averages b0.2‰, and for apatite δ13C b0.15‰. 6. Results and discussion The relevant sampling information and isotope results can be found in Table 1. It should be noted that comprehensive stable isotope analyses could not be conducted on every skeleton as some individuals did not possess intact teeth, while others had very poor bone preservation. For collagen isotope analyses 32 samples were originally processed, three yielded insufficient collagen for isotope analyses and a fourth had a very low yield and excessively high weight % C and weight % N (wt.% C and wt.% N). These four samples are excluded from further consideration. Overall, collagen content was somewhat variable ranging from 0.8% to 10.5% (mean = 4.3%), consistent with reported variations in archeological bone collagen preservation from tropical contexts (Pestle and Colvard, 2012). Despite variation in collagen content, the remaining 29 samples reported herein had good quality control indicators with C:N ratios falling within the accepted range of 2.9–3.6 (Ambrose, 1990; DeNiro, 1985), wt.% C and wt.% N above the preferred ranges of 13.0% and 4.8%, respectively (Ambrose, 1990), and a strong positive linear correlation between wt.% C and wt.% N (R2 = 0.9551) as would be

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Table 1 Demographic information and stable isotope results from Lavoutte, St. Lucia. Sample ID

Age cat.

Age

Sex

δ13Cco

δ15N

‰ VPDB

‰ AIR

‰ VPDB

‰ VPDB

‰ VPDB

‰ VPDB

F57-02 F57-03 F57-04 F57-08 F57-10 F57-11 F57-17 F57-23 F58-22 F58-23A F58-23B F67-05 F67-11 F67-12 F67-14 F67-18 F67-19 F67-25 F67-27 F67-30 F67-31 F67-33 F68-01 F68-04 F68-05 F68-06 F68-07 F68-08 F68-11 F68-20 F68-21 F68-29 F69-01 F69-02 F69-05 A1 A2 A3 A4

Adult Juvenile Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Juvenile Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Adult Juvenile Adult Adult Adult Adult Adult Juvenile Adult Adult Adult Adult Juvenile

26–35 10–12 18+ 18+ 18+ 18+ 36–45 26–35 36–45 18–25 26–35 26–35 18+ 18+ 4–5 18+ 18–25 18+ 18+ 18+ 36–45 46+ 26–35 26–35 36–45 18+ 26–35 4–5 46+ 36–45 26–35 18+ 36–45 14–16 36–45 36–45 46+ 18+ 5–6

Male Indeter. Male Indeter. Indeter. Female Female Female Male Female Male Female? Indeter. Female? Indeter. Indeter. Indeter. Indeter. Male Indeter. Male Male Male Male Female Indeter. Male Indeter. Female Female Female? Female? Male Female? Indeter. Male Male Female Indeter.

−15.85 −15.01 −17.04 −15.15 −15.37 −16.26 −15.66 −16.10 −14.89 – – – – −15.49 −14.96 −17.18 −15.71 −15.88 −17.11 −15.81 – −16.62 −15.46 −16.17 −15.87 −15.50 −16.04 −15.47 −17.36 −15.75 −16.34 – −16.84 – −16.37 – – – –

10.86 10.96 10.07 11.93 11.80 11.93 11.38 11.37 11.60 – – – – 12.26 11.70 11.63 11.76 12.02 11.63 11.32 – 11.29 11.78 11.22 11.84 12.54 11.60 11.45 11.00 11.49 12.05 – 11.86 – 12.22 – – – –

−6.72 −8.07 −10.32 −9.87 −5.52 −6.99 −9.72 −10.43 −10.03 – – – – −9.69 −10.77 −10.48 −7.92 −10.80 −8.30 −9.99 – −8.42 −10.69 −9.20 −8.93 −8.44 −10.41 −10.67 −10.81 −7.40 −7.72 – −8.05 – −9.44 – – – –

9.13 6.95 6.72 5.28 9.85 9.27 5.93 5.68 4.87 – – – – 5.79 4.19 6.70 7.78 5.08 8.82 5.82 – 8.20 4.77 6.97 6.94 7.06 5.63 4.80 6.56 8.35 8.62 – 8.79 – 6.94 – – – –

– – – −11.28 – −9.82 −11.51 −9.71 −9.38 −9.07 −12.05 −12.49 −9.50 −9.98 −9.09 −12.09 −9.55 – – – −10.03 −9.56 −10.39 −9.60 −9.98 – −9.46 −11.26 −9.57 −9.38 – −9.59 −10.95 −9.25 −10.94 −9.66 −9.67 −9.58 −9.32

– – – −1.41 – −2.83 −1.79 0.71 0.65 – – – – −0.29 1.68 −1.61 −1.63 – – – – −1.14 0.30 −0.40 −1.05 – 0.95 −0.59 1.24 −1.98 – – −2.90 – −1.50 – – – –

excepted from biogenic collagen (Pestle and Colvard, 2012). Based on the assumption that the preservation of bone apatite and collagen are linked (e.g., Nelson et al., 1986; Tütken et al., 2008), δ13Cap from individuals with poor collagen content are considered suspect and have been excluded from the dataset. Summary statistics of the isotope results are presented in Table 2, and individual isotope values are plotted in Fig. 3. Carbon and nitrogen isotope analysis of Lavoutte bone collagen produced δ13Cco ranging from − 17.4 to − 14.9‰ (mean − 16.0‰) and δ15N ranging from 10.1 to 12.5‰ (mean 11.6‰). Carbon isotope analysis of bone and enamel apatite yielded δ13Cap values ranging from − 10.8 to − 5.5‰ (mean −9.1‰), and δ13Cen ranging from −12.5 to − 9.1‰ (mean −10.1‰). Spacing between bone collagen and apatite values was variable, with Δ13Cap-co ranging from 4.2 to 9.8‰ (mean 6.9‰). Clear differences were also observed between some bone and enamel apatite carbon isotope values, with Δ13Cen-ap ranging from −2.9 to +1.7‰ (mean 0.8‰). The collagen isotope results display considerably less variation than bone or enamel apatite, which may indicate greater variability in whole diet relative to dietary protein. The range of Lavoutte δ13Cco values are intermediate relative to the overall possible range of δ13Cco recorded in humans and are broadly consistent with mixed diets containing both terrestrial and marine protein sources. In contrast, the δ15N values are higher than average values for terrestrial diets and are more comparable with populations consuming higher trophic level protein. The δ13Cap and δ13Cen values display notable enrichment in 13C, are somewhat intermediate within the overall spectrum of human variation, and indicate that dietary energy components was likely derived from

δ13Cap

Δ13Cap-co

δ13Cen

Δ13Cen-ap

Uncalib. 14C years B.P.

– – – – – – – 740 ± 30 750 ± 30 720 ± 35 – – 770 ± 35 – – – – – – – – – 920 ± 25 790 ± 35 – 865 ± 35 – – 745 ± 30 820 ± 35 – – − 620 ± 40 960 ± 35 – – – –

Table 2 Summary statistics of multiple stable isotope results from Lavoutte, St. Lucia. Statistic

δ13Cco

‰ VPDB ‰ AIR ‰ VPDB ‰ VPDB

‰ VPDB ‰ VPDB

Mean Std. dev. Min. Max. n Female Mean Std. dev. Min. Max. n Male Mean Std. dev. Min. Max. n Indeterm. Mean Std. dev. Min. Max. n Juveniles Mean Std. dev. Min. Max. n

−16.0 0.7 −17.4 −14.9 28 −16.1 0.6 −17.4 −15.5 8 −16.2 0.8 −17.1 −14.9 9 −15.9 0.6 −17.2 −15.1 8 −15.1 0.3 −15.5 −15.0 3

−10.1 1.0 −12.5 −9.1 29 −10.1 1.0 −12.5 −9.1 11 −10.1 0.8 −12.0 −9.4 10 −10.7 1.1 −12.1 −9.5 5 −9.7 1.5 −11.3 −9.1 2

Category

All

δ15N

11.6 0.5 10.1 12.5 28 11.7 0.4 11.0 12.3 8 11.3 0.6 10.1 11.9 9 11.9 0.4 11.3 12.5 8 11.4 0.4 11.0 11.7 3

δ13Cap

−9.1 1.4 −10.8 −5.5 27 −9.3 1.5 −11.5 −7.0 10 −9.2 1.3 −10.7 −6.7 10 −9.3 1.6 −10.9 −5.5 10 −9.8 1.5 −10.8 −8.1 3

Δ13Cap-co δ13Cen

6.9 1.6 4.2 9.8 27 7.1 1.4 5.7 9.3 8 7.1 1.7 4.8 9.1 9 4.9 5.8 −10.0 9.8 9 5.3 1.4 4.2 6.9 3

Δ13Cen-ap

−0.8 1.4 −2.9 1.7 18 −0.8 1.3 −2.8 1.2 9 −0.4 1.3 −2.9 1.0 7 −1.1 0.9 −1.6 0.5 5 0.5 1.6 −0.6 1.7 2

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mixed diets dominated by C3 plants but with substantial proportions of C4/CAM plants. The Δ13Cap-co results are consistently greater than 4‰ indicating that the protein component of the diet was depleted relative to the whole diet. Considering that the δ13Cco and particularly the δ15N results indicate substantial contributions of marine protein, this result is somewhat surprising and suggests that the C4/CAM contributions to whole diets must have been considerable. The isotope data were assessed for linear correlations using Pearson's r correlation coefficients. No statistically significant correlation was found between collagen and apatite δ13C, which is a good indication that these each reflect different sources of dietary carbon, namely protein and energy (dominated by carbohydrates) respectively. Of the four bone isotope indicators, the only statistically significant correlations exist between δ13Cco and Δ13Cap-co (R2 = 0.16, n = 28, p = 0.017), and between δ13Cap and Δ13Cap-co (R2 = 0.81, n = 28, p b 0.001). These strong correlations are not surprising given that δ13Cco and δ13Cap are used to calculate the Δ13Cap-co spacing. The lack of correlation between δ15N and Δ13Cap-co, however, is perhaps surprising since they might be excepted to be inversely correlated with each other and to both reflect marine resource consumption. A likely explanation for this pattern is that the variation in Δ13Cap-co is primarily resulting from differences in the δ13C of dietary energy. It should be noted, that the same pattern and lack of correlation exists between δ15N and the enamel to collagen δ13C spacing (Δ13Cen-co). Additionally, neither δ13Cen and δ13Cap (R2 = 0.035) nor δ13Cen and 13 δ Cco (R2 = 0.039) were strongly correlated. The lack of correlation between δ13Cen and δ13Cap contrasts with the expectation that both form from blood (bi)-carbonate. The lack of correlation between these sample types, however, has been noted by other researchers (e.g., Loftus and Sealy, 2012) and may result from variation in how the materials respond to sample treatment, age-related differences between childhood and adult diets, diagenic alteration of bone apatite, or a combination of these factors (e.g., Garvie-Lok et al., 2004; King et al., 2011; Koch et al., 1997; Lee-Thorp and Sponheimer, 2003; Metcalfe et al., 2009; Nielsen-Marsh and Hedges, 2000a, 2000b; Pestle et al., 2014; Wright and Schwarcz, 1996). Furthermore, no systematic offset in δ 13C was found as some individuals possessed positive Δ13Cen-ap offsets while others had negative values (Fig. 4), and the mean offset (0.8‰) was not substantial. Various Δ13Cen-ap spacings have been reported for humans and animals (Clementz et al., 2007; Loftus and Sealy, 2012; Warinner and Tuross, 2009) and clearly the exact nature of the relationship between δ13Cen and δ13Cap remains poorly understood. Despite the presence of substantial absolute offsets in paired δ13Cen and δ13Cap for some individuals, these two data sets possess comparable mean values, ranges, and deviations. Part of the reasoning behind analyzing δ13Cen is that because enamel is more resistant to diagenesis, it should provide more reliable data than bone apatite which is generally less resistant. These results indicate that although the two tissues do not provide equivalent absolute isotopic signals, at least in this case they lead to similar assessments of dietary patterns and that δ13Cen measurements provide useful additional information. Lastly, the lack of correlation between δ13Cco and δ13Cen, is perhaps more unexpected as moderate to strong correlations have been reported for populations both with (Loftus and Sealy, 2012) and without (France and Owsley, 2013) access to marine resources. However, assessment of similar data sets from archeological populations in Mesoamerica (e.g., Rand et al., 2013; Somerville et al., 2013) and in the Caribbean (Laffoon et al., 2013; Laffoon and de Vos, 2011; Norr, 2002; Stokes, 1998) reveals that δ13Cco and δ13Cen are often not well correlated, consistent with the notion that these two isotope proxies reflect different aspects of diet in biocultural contexts where dietary energy and protein sources vary independently in δ13C.

Fig. 3. Human bone stable isotope values from Lavoutte, St. Lucia: A) δ13Cco and δ15N; B) δ13Cap and δ13Cco; C) Δ13Cap-co and δ15N.

J.E. Laffoon et al. / Journal of Archaeological Science: Reports 5 (2016) 168–180

More direct assessments of diet can be made by applying offsets or discrimination factors to isotope values obtained from biological tissues to estimate the isotopic composition of dietary sources. Here we combine the formula for estimating δ13C of dietary protein from δ13Cco and Δ13Cap-co [δ13Cprotein (‰) = (0.78× δ13Cco) − (0.58 × Δ13Cap-co) − 4.7 (r2 = 0.86)] from Pestle et al. (2015) with the reported mean offset in δ15N between diet and bone collagen of 3.6 ± 1.2‰ (Ambrose, 2000; DeNiro and Epstein, 1981; Hare et al., 1991; Howland et al., 2003; Sponheimer et al., 2003; Warinner and Tuross, 2010), and the formula for estimating the δ13C of (whole) diet from δ13Cap [δ13Cdiet = (1.04 × δ13Cap) − 9.2 (r2 = 0.97)] from Ambrose and Norr (1993). Applying these formulas to the isotope results from Lavoutte generates the following estimates for mean protein and whole diet: δ13 Cprotein = − 21.1 ± 2.5 (2σ); δ15Nprotein = 8.0 ± 1.0 (2σ); δ13Cdiet = −18.7 ± 3.0 (2σ). Based on comparative assessment with regional isotopic food web (e.g., Krigbaum et al., 2013; Norr, 2002; Pestle, 2010a) these dietary isotope estimates from Lavoutte are intermediate in terms of the overall range of variation for both δ13C and δ15N. This patterning in itself is revealing in that it rules out dietary practices falling at the extreme ends of the isotopic spectrum (e.g. pure C3 or C4, or pure terrestrial or marine). Intermediate isotope values, however, may indicate either the consumption of food items possessing intermediate isotopic values or mixed diets comprised of comparable quantities of foods with high and low isotopic compositions. Nonetheless, some combinations are more likely than others based on general expectations of human feeding behavior. For example, the mean δ15N value of the Lavoutte population (11.6‰) is highly enriched relative to the mean δ15N values for either C3 (3.3‰) or C4 (4.4‰) plants (Pestle, 2010a) and the enrichment greatly exceeds the estimated trophic level effect of (~3.6‰). As such, the possibility of purely herbivorous (vegetarian) diets can be ruled out. Furthermore, the Lavoutte mean δ15N value is also higher than almost all mean δ15N values of available animal protein sources (e.g., birds, freshwater fish, marine fish, bivalves, gastropods, terrestrial animals). Thus, assuming that the Lavoutte population consumed both plants and some (unknown) amounts of freshwater and terrestrial animal protein, then it is highly likely that only the regular consumption of higher trophic level marine resources (e.g., pelagic fish) could counter-balance the 15N depleted components of their diets and account for their moderately high bone δ15N values. In support of this conclusion, faunal evidence for the intensive exploitation of tuna has been recovered from other locations in the Windward Islands such as Carriacou in the Grenadines (Giovas, 2013) and on Tobago (Steadman and Jones, 2006) but has not been widely reported for other areas of the Antilles, which may indicate a regional marine-oriented strategy focused on offshore fishing of pelagic species. Lastly, the fact that the Lavoutte δ15N are moderately high and their δ13C are only mildly to moderately elevated, while not ruling out the consumption of shellfish (bivalves and gastropods), which are generally nitrogen depleted and carbon enriched,

Fig. 4. Chart of intra-individual variation in δ13Cap and δ13Cen values from Lavoutte, Saint Lucia.

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certainly indicates that they must have formed only a minor component of dietary protein. On the assumption that δ13Cap correlates well with whole diet and is likely dominated by carbohydrates, the Lavoutte δ13Cdiet estimate (−18.7‰) should provide some indication of the relative contributions of different plant foods. Pestle (2010a) has compiled a large database of food isotope values for the Caribbean, including δ13C of C3 (mean − 25.5‰; range − 30.2 to − 21.1‰); C4 (mean − 9.6‰; range − 12.5 to −8.6‰); and CAM (mean −11.4‰; range −13.2 to −10.2‰) plants. The Lavoutte whole diet δ13C estimate falls between these ranges and considering the mild to moderately enriched δ13Cco values and that the Δ13Cap-co spacings are fairly high, this indicates that whole diet is enriched in 13C relative to protein and cannot result simply from marine resource consumption. We therefore interpret the isotopic data as indicating substantial contributions of C4 and/or CAM plants to Lavoutte diet(s). This conclusion is further illustrated by plotting δ13Cap and δ13Cco (Fig. 5) against modeled dietary regression lines (Froehle et al., 2010; Kellner and Schoeninger, 2007). The two regression lines represent C3 and C4/marine dietary protein, while the endpoints of each line represent 100% C3 (left) and 100% C4 (right) whole diets, respectively. Perhaps unexpectedly, the Lavoutte data plot in the middle of the chart with most samples falling outside of the 95% confidence intervals of both regression lines or within the intervals for the C3-protein linear regression, with only two samples plotting within the confidence intervals for the C4/marine protein linear regression. The possibility that there are unresolved limitations with these published regression lines has to be considered (e.g., Rand et al., 2013). However, if accurate, these patterns suggest a large non-marine protein component to the diet at Lavoutte. The component remains unidentified but most likely represents a combination of terrestrial and freshwater animal, and C3 plant protein. Also notable is the position of most data points approximately mid-way between the ‘pure diet’ end-members, which again are consistent with mixed whole diets consisting of significant contributions various plant food groups. This conclusion is further corroborated by the recent recovery and identification of maize starch grains in the dental calculus of multiple individuals from Lavoutte (Pagán-Jiménez and Mickleburgh, 2015). It is also worth noting that various edible CAM plants including century plants (Agave sp.) and prickly pear cactus (Opuntia sp.) can be found growing in the vicinity of Lavoutte and would also likely have been exploited by the local population. The intra-population dietary variation is presented in Table 2. The stable isotope results were assessed relative to age (at death) and sex estimates derived from osteological analyses of the skeletal assemblage conducted by Darlene Weston (Hofman et al., 2012; Weston, 2011) using standard methods for the estimation of sex and age from skeletal remains. Student t-tests revealed no statistically significant differences in any of the stable isotope results between females and males. There were also no significant differences based on age, except for collagen carbon, with juveniles possessing significantly (Student's t-test, p = 0.025) more enriched δ13Cco signatures (− 15.1 ± 0.3‰) than adults (−16.1 ± 0.7‰). This observed difference, however, is based on only three juvenile samples and likely reflects age-related dietary differences (e.g., Richards et al., 2002). This general lack of clear differences in isotope values between different demographic groups is also observed in other pre-colonial populations from the Caribbean (Laffoon and de Vos, 2011; Laffoon and Hoogland, 2012; Norr, 2002; Pestle, 2010a; Stokes, 1998) and may indicate either communal food consumption practices or that dietary variation was structured by factors other than sex or age (e.g., Laffoon, in press; Pestle, 2013c). Lastly, no correlations were observed between the radiocarbon dates and any of the stable isotope results. This may indicate a lack of temporal diet change but may also result from insufficient data as only twelve of the individuals have been radiocarbon dated thus far. The isotopic evidence from Lavoutte can be placed into a broader regional perspective through comparison with published isotope datasets of pre-colonial populations from the Caribbean at various scales

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(Table 3; Fig. 6). Stokes (1998) reports stable isotope results from one individual from St. Lucia (Grand Anse site) but does not discuss these results. This individual's stable isotope values are very similar to those reported here for Lavoutte. Stokes (1998) also reports stable isotope results from two individuals from the Pearls site, Grenada, which possess comparable collagen δ13C and δ15N values to Lavoutte but much lower δ13Cap values. Although both sites are situated in broadly similar ecological settings, the Pearls skeletal materials likely date to the Early Ceramic Age (Saladoid/Huecoid) and the observed differences in δ13Cap signals may relate to chronological changes in the relative importance of C4 crops. If so, this would provide tentative evidence for increased reliance over time on C4 crops, such as maize, or CAM plants in this region. The nearest island with comparable pre-colonial isotope data is Carriacou in the Grenadines, where Krigbaum et al. (2013) reported bone collagen and apatite results from the site of Grand Bay (n = 14). Interestingly, there are some pronounced differences in isotope results between these sites with much higher δ13Cco, lower δ15N, similar δ13Cap, and much lower Δ13Cap-co at Grand Bay than at Lavoutte. These overall differences in stable isotope data indicate distinct differences in diet and subsistence strategies between these two (burial) populations. More specifically, lower trophic level (nitrogen depleted) reef and shellfish seemed to have been more important sources of dietary protein at Grand Bay then at Lavoutte, while C4/CAM plants were larger contributors to dietary energy at Lavoutte then Grand Bay. The Lavoutte collagen isotope data (Fig. 6A) are very comparable to Anse à la Gourde, Guadeloupe and are in fact more similar to populations from the northern Lesser Antilles (Leeward Islands) than they are to the geographically closer population from Grand, Bay, Carriacou. Conversely, in terms of the apatite δ13C data (Fig. 6B), the Lavoutte population are most similar with both Grand Bay, Carriacou and Anse à la Gourde, Guadeloupe (as well as various populations from Puerto Rico, and Haiti) but differ the most from other populations in the Leeward Islands and the Bahamas, although there is considerable overlap between most sites and regions. From a broader regional perspective, the collagen isotope data from Lavoutte clusters most closely with geographically dispersed locations in the Greater Antilles (Dominican Republic), and northern Lesser Antilles (e.g. Guadeloupe, Saba, St. Thomas). In fact, the Lavoutte collagen isotope data occupy an intermediate position in isotope space relative to Greater and Lesser Antillean populations. This pattern suggests that while there is some degree of spatial patterning to pre-colonial protein

Fig. 5. Mean ± 1σ and individual δ13Cco values plotted against δ13Cap (×) and δ13Cen (+) values from Lavoutte, St. Lucia plotted against the Froehle et al. (2010) model regression lines and their 95% prediction intervals.

sources in the Caribbean (e.g., Stokes, 1998) it is not structured solely by geographic parameters per se, but is more likely influenced by a combination of ecological and cultural factors. Whereas Antillean collagen isotope values overall display a weak positive correlation, those from the Lesser Antilles are slightly negatively correlated. This negative correlation could result from differences in the relative importance of lower trophic level reef and sea grass species, which tend to possess both enriched carbon and depleted nitrogen isotope values. In terms of apatite isotope data and collagen-apatite differences, there is overall less geographic clustering in isotope space and more overlap between different populations, indicating that biogeographic factors may have had a reduced influence on whole diet patterns, relative to protein. Here, the Lavoutte data once again cluster most closely with certain sites in both the Greater (Haiti, Puerto Rico) and Lesser Antilles (Guadeloupe), yet display more dispersion consistent with higher inter-individual variation in whole diet δ13C. In other words, within the circum-Caribbean, the overall patterning of collagen and apatite isotope data strongly suggest that local environmental structuring placed greater constraints on animal protein availability and consumption, while the affordances of terrestrial habitats placed less constraints on plant production and consumption strategies which were more flexible and overlapping. 7. Conclusions Stable isotope data from the burial population of Lavoutte, St. Lucia permit assessment of pre-colonial dietary patterning. Comparative analyses with isotopic food web data using conversion formulae to estimate tissue-diet fractionation reveal that dietary protein comprised both terrestrial and marine components. The substantial terrestrial protein component indicates that the Lavoutte population, although well adapted to marine resource exploitation, were not entirely dependent upon it. The relatively elevated nitrogen values indicate frequent consumption of animal foods and that marine contributions to dietary protein may have been dominated by higher trophic level (nitrogen-15 enriched) taxa, such as larger pelagic fish. Within a regional context, the high degree of inter-island variation in collagen stable isotope ratios is inconsistent with the notion that all pre-colonial Antillean populations were equally reliant on the sea for protein but rather that the importance of marine resources was highly variable between different islands and sites. In contrast, the relatively low degree of intrapopulation variation in collagen stable isotope data at Lavoutte is notable especially considering that the sample population includes both adults and sub-adults, in addition to non-contemporaneous individuals. This likely attests to the consistency of protein consumption within the population and a fairly conservative subsistence strategy whereby the relative contributions of fishing and hunting to dietary protein did not vary greatly over time. The lack of correlation between δ13Cco and δ13Cap, or δ13Cco and 13 δ Cen provides evidence that dietary protein and energy varied independently. The relatively elevated δ13Cap-co values indicate that dietary energy was enriched in 13C relative to protein and demonstrates C4/ CAM plant contributions to diet at Lavoutte. In contrast to the collagen results, the larger variance and range of δ13Cap and δ13Cap-co indicates a greater degree of intra-population heterogeneity in dietary energy and variable importance of different plant resources to individual dietary regimes. The source of this variation is not immediately apparent, as no statistically significant differences were observed between adults or sub-adults, or between males and females, for δ13Cap and δ13Cap-co. Whatever the source of this variation it does not appear to be clearly linked to these demographic variables and may instead be associated with chronology, status, natal origins, ethnic identity, or some other unidentified factor(s). If these patterns result from variations in C4 plant consumption, related directly to differential consumption of maize, this may partly explain observed differences in the presence/absence of maize starch grains between contemporaneous individuals at some pre-colonial Caribbean populations. In order to disentangle

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Table 3 Average (±1σ) stable isotope values of human bone from various pre-colonial populations in the Caribbean. Island

Site

Bahamas1 La Tortue (Haiti)1 Dominican Rep.1 Dominican Rep.1 Puerto Rico1 Puerto Rico2 Puerto Rico2 Puerto Rico2 Puerto Rico3 St. Thomas4 Anguilla1 St. Martin1 Saba1 St. Kitts5 Grande-Terre1,6 La Desirade1 St. Lucia1 Carriacou7 Grenada1 St. Lucia*

Multiple Manigat Cave Juan Dolio Boca del Soco Maisabel Paso del Indio Punta Candelero Tibes Rio Tanamá Tutu Multiple Hope Estate Multiple Bloody Point Anse à la Gourde Petite Riviere Grande Anse Grand Bay Pearls Lavoutte

δ13Cco

δ15Nco

δ13Cap

Δ13Cap-co

Mean

S.D.

Mean

S.D.

Mean

S.D.

Mean

S.D.

−13.4 −16.5 −17.1 −18.0 −18.1 −19.1 −17.5 −17.6 −19.6 −15.5 −14.4 −15.7 −15.7 −15.3 −14.8 −14.1 −16.3 −12.8 −17.0 −16.0

1.4 1.2 0.7 0.5 1.0 0.5 1.0 0.6 0.3 0.9 1.1 0.1 0.6 0.3 0.8 0.4 na 0.9 0.2 0.7

9.8 8.7 11.9 11.9 9.7 9.8 9.9 9.5 9.1 12.2 10.1 10.4 10.8 11.0 10.9 10.3 12.4 11.1 12.6 11.6

1.0 0.8 0.5 0.4 0.7 0.9 0.9 0.7 0.7 0.9 0.7 0.1 0.5 0.3 0.7 0.6 na 0.5 0.5 0.5

−10.8 −9.9 −12.5 −11.9 −9.9 −9.4 −8.3 −8.6 −10.5 −10.5 −10.7 −10.8 −11.0 −9.9 −8.4 −8.2 −8.1 −8.6 −13.0 −9.1

2.1 1.4 2.9 1.8 0.9 1.1 1.2 1.0 0.1 0.9 1.7 0.1 1.3 1.2 1.3 0.6 na 0.6 3.6 1.4

2.6 6.6 4.6 6.2 8.2 9.7 9.2 9.0 9.1 5.0 3.7 4.9 4.7 5.4 6.4 5.9 8.3 4.1 4.0 6.9

1.6 1.5 3.2 2.0 1.0 1.3 1.6 1.2 0.5 1.3 2.0 0.1 1.6 1.3 1.5 0.9 na 1.1 3.4 1.6

Data sources: 1Stokes (1998); 2Pestle (2010a); 3Antón (2008); 4Norr (2002); 5Farr, 1996; 6Laffoon and de Vos (2011); 7Krigbaum et al. (2013); *this study.

chronological and spatial variation in dietary practices and to investigate various hypotheses concerning consistency and change in human foodways in the circum-Caribbean, a larger scale dietary study

incorporating paleobotanical (macro- and micro-), zooarcheological, dental anthropological (wear and pathology), and isotopic (radiocarbon and stable) analyses has been initiated. To make a more quantitative assessment of the data, future research will also focus on: the application of dietary mixing models; further testing of different possible sources of inter-individual variation in dietary practices; and continued spatial and temporal expansion of human and foodweb stable isotope data sets in the circum-Caribbean. Acknowledgments The research leading to these results has received funding from the European Research Council under the European Union's Seventh Framework Programme (FP7/2007-2013)/ERC grant agreement no. 319209. At the VU University Amsterdam, we wish to acknowledge the contributions of Suzan Verdegaal-Warmerdam and Remy van Baal, who conducted the stable isotope measurements, and Robin van der Velde for her assistance with the preparation and processing of the skeletal samples. We are very grateful to Eric Milton Branford and the Saint Lucia Archaeological and Historical Society for granting access to study the Lavoutte archeological collections. This paper was improved by the valuable feedback of two anonymous reviewers. References

Fig. 6. A) Collagen δ13Cco and δ15N; and B) δ13Cap and Δ13Cap-co isotope values from Lavoutte, St. Lucia compared to other pre-colonial populations in the circum-Caribbean. Symbols with error bars represent population means ±1σ; symbols without error bars represent individual data points. Data from Krigbaum et al. (2013), Laffoon and de Vos (2011), Norr (2002), Pestle (2010a) and Stokes (1998).

Allaire, L., 1999. Archaeology of the Caribbean Region. In: Solomon, F., Schwartz, S.B. (Eds.), The Cambridge History of the Native Peoples of the Americas. Vol. III: South America. Cambridge University Press, Cambridge, pp. 668–733. Ambrose, S.H., 1990. Preparation and characterization of bone and tooth collagen for isotopic analysis. J. Archaeol. Sci. 17 (4), 431–451. Ambrose, S.H., 2000. Controlled diet and climate experiments on nitrogen isotope ratios of rats. In: Ambrose, S.H., Katzenberg, M.A. (Eds.), Biogeochemical Approaches to Paleodietary Analysis. Kluwer Academic, New York, pp. 243–259. Ambrose, S.H., Butler, B.M., Hanson, D.B., Hunter-Anderson, R.L., Krueger, H.W., 1997. Stable isotopic analysis of human diet in the Marianas Archipelago, Western Pacific. Am. J. Phys. Anthropol. 104 (3), 343–361. Ambrose, S.H., Krigbaum, J., 2003. Bone chemistry and bioarchaeology. J. Anthropol. Archaeol. 22 (3), 193–199. Ambrose, S.H., Norr, L., 1993. Experimental evidence for the relationship of the carbon isotope ratios of whole diet and dietary protein to those of bone collagen and carbonate. In: Lambert, J.B., Grupe, G. (Eds.), Prehistoric Human Bone: Archaeology at the Molecular Level. Springer, Berlin, pp. 1–37. Antón, S.C., 2008. Human remains from the Río Tanamá Sites (AR-38 and AR-39). In: Carlson, L.A. (Ed.), A Multidisciplinary Approach to Site Testing and Data Recovery at Two Village Sites (AR-38 and AR-39) on the Lower Río Tanamá, Municipality of Arecibo, Puerto Rico. Southeastern Archaeological Research, Inc., Jacksonville, pp. 149–190.

178

J.E. Laffoon et al. / Journal of Archaeological Science: Reports 5 (2016) 168–180

Bender, M.M., 1971. Variations in the 13C/12C ratios of plants in relation to the pathway of photosynthetic carbon dioxide. Phytochemistry 10, 1239–1244. Bender, M.M., Rouhani, I., Vines, H.M., Black, C., 1973. 13C/12C ratio changes in Crassulacean acid metabolism plants. Plant Physiol. 52 (5), 427–430. Berman, M.J., Pearsall, D.M., 2000. Plants, people, and culture in the prehistoric Central Bahamas: a view from the three Dog site, an Early Lucayan settlement on San Salvador Island, Bahamas. Lat. Am. Antiq. 11 (3), 219–239. Berman, M.J., Pearsall, D.M., 2008. At the crossroads: starch grain and phytolith analyses in Lucayan prehistory. Lat. Am. Antiq. 19 (2), 181–203. Bocherens, H., Sandrock, O., Kullmer, O., Schrenk, F., 2011. Hominin palaeoecology in Late Pliocene Malawi: first insights from isotopes (13C, 18O) in mammal teeth. S. Afr. J. Sci. 107 (3–4), 1–6. Brown, T.A., Earle Nelson, D., Vogel, J.S., Southon, J.R., 1988. Improved collagen extraction by modified Longin method. Radiocarbon 30 (2), 171–177. Buhay, W.M., Chinique de Armas, Y., Rodríguez Suárez, R., Arredondo, C., Smith, D.G., Armstrong, S.D., Roksandic, M., 2013. A preliminary carbon and nitrogen isotopic investigation of bone collagen from skeletal remains recovered from a Pre-Columbian burial site, Matanzas Province, Cuba. Appl. Geochem. 32, 76–84. Bullen, A.K., Bullen, R.P., 1970. The Lavoutte Site, St. Lucia: a Carib Ceremonial Centre. Proceedings of the 3rd International Congress for the Study of Pre-Columbian Cultures of the Lesser Antilles. Grenada National Museum, Grenada, pp. 61–86. Burney, D.A., Burney, L.P., DE MacPhee, R., 1994. Holocene charcoal stratigraphy from Laguna Tortuguero, Puerto Rico, and the timing of human arrival on the island. J. Archaeol. Sci. 21 (2), 273–281. Capone, D.G., Carpenter, E.J., 1982. Nitrogen fixation in the marine environment. Science 217, 1140–1142. Carlson, Lisabeth Anne. 1999. Aftermath of a Feast: Human Colonization of the Southern Bahamian Archipelago and Its Effects on the Indigenous Fauna. PhD diss., Department of Anthropology, University of Florida, Gainesville. Carlson, L.A., Keegan, W.F., 2004. Resource depletion in the prehistoric northern West Indies. In: Fitzpatrick, S.M. (Ed.), Voyages of Discovery: The Archaeology of Islands. Greenwood Publishing Group, Westport, pp. 85–107. Chinique de Armas, Y., Buhay, W.M., Rodríguez Suárez, R., Bestel, S., Smith, D.G., Mowat, S.D., Roksandic, M., 2015. Starch analysis and isotopic evidence of consumption of cultigens among fisher–gatherers in Cuba: the archaeological site of Canímar Abajo, Matanzas. J. Archaeol. Sci. 58, 121–132. Chisholm, B.S., Earle Nelson, D., Schwarcz, H.P., 1982. Stable-carbon isotope ratios as a measure of marine versus terrestrial protein in ancient diets. Science 216, 1131–1132. Clementz, M.T., Koch, P.L., Beck, C.A., 2007. Diet induced differences in carbon isotope fractionation between sirenians and terrestrial ungulates. Mar. Biol. 151 (5), 1773–1784. Curet, L.A., Oliver, J.R., 1998. Mortuary practices, social development, and ideology in precolumbian Puerto Rico. Lat. Am. Antiq. 9 (3), 217–239. Curtis, J.H., Brenner, M., Hodell, D.A., 2001. Climate change in the circum-Caribbean (Late Pleistocene to Present) and implications for regional biogeography. In: Woods, C.A., Sergile, F.E. (Eds.), Biogeography of the West Indies: Patterns and Perspectives. CRC Press, Boca Raton, pp. 35–54. deFrance, S.D., 2009. Zooarchaeology in complex societies: political economy, status, and ideology. J. Archaeol. Res. 17, 105–168. deFrance, S.D., 2013. Zooarchaeology in the Caribbean: current research and future prospects. In: Keegan, W.F., Hofman, C.L., Rodríguez Ramos, R. (Eds.), The Oxford Handbook of Caribbean Archaeology. Oxford University Press, Oxford, pp. 378–390. DeNiro, M.J., 1985. Postmortem preservation and alteration of in vivo bone collagen isotope ratios in relation to palaeodietary reconstruction. Nature 317, 806–809. DeNiro, M.J., Epstein, S., 1978. Influence of diet on the distribution of carbon isotopes in animals. Geochim. Cosmochim. Acta 42 (5), 495–506. DeNiro, M.J., Epstein, S., 1981. Influence of diet on the distribution of nitrogen isotopes in animals. Geochim. Cosmochim. Acta 45 (3), 341–351. Fabrizii-Reuer, S., Reuer, E., 2005. Die Gräber aus den “shellmiddens” der präkolumbianischen Siedlung von Pointe de Caille, St. Lucia, West Indies. Verlag der Österreichischen Akademie der Wissenschaften, Vienna. Farr, Starr, 1996. Unpublished Report on the Results of Stable Isotope Analysis of Human Remains from Bloody Point, St. Kitts. Report on file. St. Christopher National Trust, St. Kitts. Fernandes, R., Nadeau, M.-J., Grootes, P.M., 2012. Macronutrient based model for dietary carbon routing in bone collagen and bioapatite. Archaeol. Anthropol. Sci. 4, 291–301. Fitzpatrick, S.M., Keegan, W.’a.F., Sullivan Sealey, K., 2008. Human impacts on marine environments in the West Indies during the Middle to Late Holocene. In: Rick, T.C., Erlandson, J.M. (Eds.), Human Impacts on Ancient Marine Ecosystems: A Global Perspective. University of California Press, Berkeley, pp. 147–164. France, C.A.M., Owsley, D.W., 2013. Stable carbon and oxygen isotope spacing between bone and tooth collagen and hydroxyapatite in human archaeological remains. Int. J. Osteoarchaeol. 25 (3), 299–312. Froehle, A.W., Kellner, C.M., Schoeninger, M.J., 2010. Focus: effect of diet and protein source on carbon stable isotope ratios in collagen: follow up to Warinner and Tuross (2009). J. Archaeol. Sci. 37, 2662–2670. Froehle, A.W., Kellner, C.M., Schoeninger, M.J., 2012. Multivariate carbon and nitrogen stable isotope model for the reconstruction of prehistoric human diet. Am. J. Phys. Anthropol. 147, 352–369. Garvie-Lok, S.J., Varney, T.L., Anne Katzenberg, M., 2004. Preparation of bone carbonate for stable isotope analysis: the effects of treatment time and acid concentration. J. Archaeol. Sci. 31 (6), 763–776.

Giovas, Christina Marguerite. 2013. Foraging Variability in the Prehistoric Caribbean: Multiple Foraging Optima, Resource Use, and Anthropogenic Impacts on Carriacou, Grenada. PhD diss., Department of Anthropology, University of Washington. Giovas, C.M., LeFebvre, M.J., Fitzpatrick, S.M., 2012. New records for prehistoric introduction of Neotropical mammals to the West Indies: evidence from Carriacou, Lesser Antilles. J. Biogeogr. 39 (3), 476–487. Grouard, Sandrine. 2001. Subsistance, Systèmes Techniques et Gestion Territoriale en Milieu Insulaire Antillais Précolombien. Exploitation des Vertébrés et des Crustacés aux époques Saladoïdes et Troumassoïdes de Guadeloupe (400 av. J.-C. à 1 500 ap. J.-C.). PhD diss., U.F.R. Sciences Sociales et Administration, University of Paris, Paris. Hare, P.E., Fogel, M.L., Stafford, T.W., Mitchell, A.D., Hoering, T.C., 1991. The isotopic composition of carbon and nitrogen in individual amino acids isolated from modern and fossil proteins. J. Archeol. Sci. 18, 277–292. Harrison, R.G., Katzenberg, M.A., 2003. Paleodiet studies using stable carbon isotopes from bone apatite and collagen: examples from Southern Ontario and San Nicolas Island, California. J. Anthropol. Archaeol. 22 (3), 227–244. Healy, P.F., Keenleyside, A., Dorst, M.C., 2013. Isotope analysis and radiocarbon dating of Prehistoric human bone from the Manzanilla (SAN 1) Site, Trinidad. Caribbean Connections 3 (1), 30–45. Hedges, R.E.M., Reynard, L.M., 2007. Nitrogen isotopes and the trophic level of humans in archaeology. J. Archaeol. Sci. 34 (8), 1240–1251. Hodell, D.A., Curtis, J.H., Jones, G.A., Higuera-Gundy, A., Brenner, M., Binford, M.W., Dorsey, K.T., 1991. Reconstruction of Caribbean climate change over the past 10, 500 years. Nature 352 (6338), 790–793. Hofman, C.L., 2013. The Post-Saladoid in the Lesser Antilles (A.D. 600/800–1492). In: Keegan, W.F., Hofman, C.L., Rodríguez Ramos, R. (Eds.), The Oxford Handbook of Caribbean Archaeology. Oxford University Press, Oxford, pp. 205–220. Hofman, C.L., Branford, E.M., 2011. Lavoutte revisited, preliminary results of the 2009 rescue excavations at Cas-en- Bas, St. Lucia. In: Rebovich, S.A. (Ed.), Proceedings of the 23rd International Congress for Caribbean Archaeology. Dockyard Museum, English Harbour, Antigua, pp. 690–700. Hofman, C.L., Hoogland, M.L.P., Mickleburgh, H.L., Laffoon, J.E., Weston, D.A., Field, M.H., 2012. Life and death at precolumbian Lavoutte, Saint Lucia, Lesser Antilles. J. Field Archaeol. 37 (3), 209–225. Howland, M.R., Corr, L.T., Young, S.M.M., Jones, V., Jim, S., Van der Merwe, N.J., Mitchell, A.D., Evershed, R.P., 2003. Expression of the dietary isotope signal in the compound-specific δ13C values of pig bone lipids and amino acids. Int. J. Osteoarchaeol. 13, 54–65. Iacumin, P., Bocherens, H., Mariotti, A., Longinelli, A., 1996. An isotopic palaeoenvironmental study of human skeletal remains from the Nile Valley. Palaeogeogr. Palaeoclimatol. Palaeoecol. 126, 15–30. Jim, S., Jones, V., Ambrose, S.H., Evershed, R.P., 2006. Quantifying dietary macronutrient sources of carbon for bone collagen biosynthesis using natural abundance stable carbon isotope analysis. Br. J. Nutr. 95 (6), 1055–1062. Katzenberg, M.A., 2008. Stable isotope analysis: a tool for studying past diet, demography, and life history. In: Katzenberg, M.A., Saunders, S.R. (Eds.), Biological Anthropology of the Human Skeleton. John Wiley & Sons, Inc., New York, pp. 411–441. Keegan, William F. 1985. Dynamic Horticulturalists: Population Expansion in the Prehistoric Bahamas. PhD diss., Department of Anthropology, University of California, Los Angeles. Keegan, W.F., 2000. West Indian archaeology. 3. Ceramic age. J. Archaeol. Res. 8 (2), 135–167. Keegan, W.F., DeNiro, M.J., 1988. Stable carbon- and nitrogen- isotope ratios of bone collagen used to study coral-reef and terrestrial components of prehistoric Bahamian diet. Am. Antiq. 53 (2), 320–336. King, C.L., Tayles, N., Gordon, K.C., 2011. Re-examining the chemical evaluation of diagenesis in human bone apatite. J. Archaeol. Sci. 38, 2222–2230. Koch, P.L., Tuross, N., Fogel, M.L., 1997. The effects of sample treatment and diagenesis on the isotopic integrity of carbonate in biogenic hydroxylapatite. J. Archaeol. Sci. 24, 417–429. Kellner, C.M., Schoeninger, M.J., 2007. A Simple Carbon Isotope Model for Reconstructing Prehistoric Human Diet. Am. J. Phys. Anth. 133, 1112–1127. Krigbaum, J., Fitzpatrick, S.M., Jamie Bankaitis, 2013. Human paleodiet at Grand Bay, Carriacou, Lesser Antilles. J. Island Coastal Archaeol. 8 (2), 210–227. Krueger, H.W., Sullivan, C.H., 1984. Models for carbon isotope fractionation between diet and bone. In: Turnlund, J., Johnson, P.E. (Eds.), Stable Isotopes in Nutrition. American Chemical Society, Washington, DC, pp. 205–222. Laffoon, Jason E. 2012. Patterns of Paleomobility in the Ancient Antilles: An Isotopic Approach. PhD diss., Faculty of Archaeology, Leiden University, Leiden. Laffoon, Jason E. (in press). Human mobility and dietary patterns in precolonial Puerto Rico: integrating multiple isotope data. In Cuban Archaeology in the CircumCaribbean Context, edited by I. Roksandic. Gainesville, University Press of Florida. Laffoon, J., de Vos, B., 2011. Diverse origins, similar diets: an integrated isotopic perspective from Anse à la Gourde, Guadeloupe. In: Hofman, C.L., van Duijvenbode, A. (Eds.), Communities in Contact. Essays in Archaeology, Ethnohistory & Ethnography of the Amerindian Circum-Caribbean. Sidestone Press, Leiden, pp. 187–204. Laffoon, Jason E., Menno L.P. Hoogland. 2012. Migration and mobility in the CircumCaribbean: integrating archaeology and isotopic analysis. In Population Dynamics in Prehistory and Early History. New Approaches Using Stable Isotopes and Genetics, edited by E. Kaiser, J. Burger, W. Schier, 337–353. Topoi: Berlin Studies of the Ancient World 5. Berlin: de Gruyter. Laffoon, J.E., Rojas, R.V., Hofman, C.L., 2013. Oxygen and carbon isotope analysis of human dental enamel from the Caribbean: implications for investigating individual origins. Archaeometry 55 (4), 742–765. Lane, C.S., Horn, S.P., Orvis, K.H., Thomason, J.M., 2011. Oxygen isotope evidence of Little Ice Age aridity on the Caribbean slope of the Cordillera Central, Dominican Republic. Quat. Res. 75 (3), 461–470. Lee-Thorp, J.A., 2008. On isotopes and old bones*. Archaeometry 50 (6), 925–950.

J.E. Laffoon et al. / Journal of Archaeological Science: Reports 5 (2016) 168–180 Lee-Thorp, J.A., Sealy, J.C., Van der Merwe, N.J., 1989. Stable carbon isotope ratio differences between bone collagen and bone apatite, and their relationship to diet. J. Archaeol. Sci. 16 (6), 585–599. Lee-Thorp, J., Sponheimer, M., 2003. Three case studies used to reassess the reliability of fossil bone and enamel isotope signals for paleodietary studies. J. Anthropol. Archaeol. 22 (3), 208–216. LeFebvre, M.J., deFrance, S.D., 2014. Guinea pigs in the Pre-Columbian West Indies. J. Island Coastal Archaeol. 9 (1), 16–44. Loftus, E., Sealy, J., 2012. Technical note: interpreting stable carbon isotopes in human tooth enamel: an examination of tissue spacings from South Africa. Am. J. Phys. Anthropol. 147 (3), 499–507. Longin, R., 1971. New method of collagen extraction for radiocarbon dating. Nature 230, 241–242. Metcalfe, J.Z., Longstaffe, F.J., White, C.D., 2009. Method-dependent variations in stable isotope results for structural carbonate in bone bioapatite. J. Archaeol. Sci. 36 (1), 110–121. Metheny, K.B., Beaudry, M.C. (Eds.), 2015. Archaeology of Food: An Encyclopedia. Rowman and Littlefield, London. Mickleburgh, Hayley L. 2013. Reading the Dental Record. A Dental Anthropological Approach to Foodways, Health and Disease, and Crafting in the pre-Columbian Caribbean. PhD diss., Faculty of Archaeology, Leiden University, Leiden. Mickleburgh, H.L., 2014. Dental wear and pathology in the precolonial Caribbean: evidence for dietary change in the ceramic age. Int. J. Osteoarchaeol. http://dx.doi.org/ 10.1002/oa.2421. Mickleburgh, H.L., Pagán-Jiménez, J.R., 2012. New insights into the consumption of maize and other food plants in the pre-Columbian Caribbean from starch grains trapped in human dental calculus. Journal of Archaeological Science 39 (7), 2468–2478. Mickleburgh, Hayley L., Jason E. Laffoon. (in press). Assessing dietary and subsistence transitions on prehistoric Aruba: preliminary bioarchaeological evidence. In The Archaeology of Caribbean and Circum-Caribbean Farmers (5000 BC–AD 1500), edited by B.A. Reid. Gainesville: University Press of Florida. Minigawa, M., Wada, E., 1984. Stepwise enrichment of 15N along food chains: further evidence and the relation between δ15N and animal age. Geochim. Cosmochim. Acta 48 (5), 1135–1140. Nelson, B.K., DeNiro, M.J., Schoeninger, M.J., De Paolo, D.J., Edgar Hare, P., 1986. Effects of diagenesis on strontium, carbon, nitrogen and oxygen concentration and isotopic composition of bone. Geochimica et Cosmochimica Acta 50 (9), 1941–1949. Newsom, L.A., Pearsall, D.M., 2003. Trends in Caribbean Island Archaeobotany. In: Minnis, P.E. (Ed.), Plants and People in Ancient Eastern North America. Smithsonian Books, Washington DC, pp. 347–412. Newsom, L.A., Wing, E.S., 2004. On Land and Sea: Native American Uses of Biological Resources in the West Indies. University of Alabama Press, Tuscaloosa. Nielsen-Marsh, C.M., Hedges, R.E.M., 2000a. Patterns of diagenesis in bone I: the effects of site environments. J. Archaeol. Sci. 27, 1139–1150. Nielsen-Marsh, C.M., Hedges, R.E.M., 2000b. Patterns of diagenesis in bone II: effects of acetic acid treatment and the removal of diagenetic CO−2 3 . J. Archaeol. Sci. 27, 1151–1159. Norr, L., 2002. Bone isotopic analysis and prehistoric diet at the Tutu Site. In: Righter, E. (Ed.), The Tutu Archeological Village Site: A Multidisciplinary Case Study in Human Adaptation. Routledge, London, pp. 263–273. O'Leary, M.H., 1981. Carbon isotope fractionation in plants. Phytochemistry 20 (4), 553–567. Pagán-Jiménez, J.R., 2011. Early phytocultural processes in the Pre-Colonial Antilles. A Pan-Caribbean survey for an ongoing starch grain research. In: Hofman, C.L., van Duijvenbode, A. (Eds.), Communities in Contact. Essays in Archaeology, Ethnohistory, and Ethnography of the Amerindian circum-Caribbean. Sidestone Press, Leiden, pp. 87–116. Pagán Jiménez, J.R., 2013. Human–plant dynamics in the precolonial antilles: a synthetic update. In: Keegan, W.F., Hofman, C.L., Rodríguez Ramos, R. (Eds.), The Oxford Handbook of Caribbean Archaeology. Oxford University Press, Oxford, pp. 391–406. Pagán-Jiménez, Jaime R., Hayley L. Mickleburgh. 2015. Starchy plant food consumption in the precolonial Caribbean: new evidence from ancient human dental calculus. Paper presented at the 26th Congress of the International Association of Caribbean Archaeology, 19–25, July, 2015. St. Maarten. Pagán-Jiménez, J.R., Rodríguez-Ramos, R., 2007. Sobre el Origen de la Agricultura en las Antillas. In: Reid, B. (Ed.), Proc. Int. Congr. Caribb. Archaeol. 1 (21). University of the West Indies, Trinidad, pp. 252–259 Pagán-Jiménez, J.R., Oliver, J.R., 2008. Starch residues on lithic artifacts from two contrasting contexts in North Central Puerto Rico: Los Muertos Cave and Vega Nelo Vargas Farmstead. In: Hofman, C.L., Hoogland, M.L.P., van Gijn, A.L. (Eds.), Crossing the Borders: New Methods and Techniques in the Study of Archaeological Materials from the Caribbean. The University of Alabama Press, Tuscaloosa, pp. 137–158. Pagán-Jiménez, J.R., Rodríguez-Ramos, R., Reid, B.A., van den Bel, M., Hofman, C.L., 2015. Early dispersals of maize and other food plants into the Southern Caribbean and Northeastern South America. Quat. Sci. Rev. 123, 231–246. Parker-Pearson, M., 2003. Food, Culture and Identity in the Neolithic and Early Bronze Age. Archaeopress, Oxford. Pearsall, D.M., 2002. Analysis of charred botanical remains from the Tutu site. In: Righter, E. (Ed.), The Tutu Archaeological Village Site: A Multidisciplinary Case Study in Human Adaptation. Routledge, London, pp. 109–134. Pestle, William J. 2010a. Diet and Society in Prehistoric Puerto Rico. PhD diss., Department of Anthropology, University of Illinois at Chicago, Chicago. Pestle, W.J., 2010b. Bone chemistry and paleodiet at the ceremonial center of Tibes. In: Curet, L.A., Stringer, L.M. (Eds.), Tibes: People, Power, and Ritual at the Center of the Cosmos. University of Alabama Press, Tuscaloosa, pp. 209–230.

179

Pestle, W.J., 2013a. Stable isotope analysis of paleodiet in the Caribbean. In: Keegan, W.F., Hofman, C.L., Rodríguez Ramos, R. (Eds.), The Oxford Handbook of Caribbean Archeology. Oxford University Press, Oxford, pp. 407–417. Pestle, W.J., 2013b. Fishing down a prehistoric Caribbean marine food web: Isotopic evidence from Punta Candelero, Puerto Rico. J. Island Coastal Archaeol. 8 (2), 228–254. Pestle, W.J., 2013c. Equals in death as in life? Mortuary and isotopic variation in Late Ceramic Age Puerto Rico. J. Caribb. Archaeol. 13, 27–42. Pestle, W.J., Colvard, M., 2012. Bone collagen preservation in the tropics: a case study from ancient Puerto Rico. J. Archaeol. Sci. 39 (7), 2079–2090. Pestle, W.J., Crowley, B.E., Weirauch, M.T., 2014. Quantifying inter-laboratory variability in stable isotope analysis of ancient skeletal remains. PLoS One 9 (7), e102844. Pestle, W.J., Hubbe, M., Smith, E.K., Stevenson, J.M., 2015. A linear model for predicting δ13 Cprotein. Am. J. Phys. Anthropol. 157 (4), 694–703. Rand, A.J., Healy, P.F., Awe, J.J., 2013. Stable isotopic evidence of Ancient Maya diet at Caledonia, Cayo District, Belize. Int. J. Osteoarchaeol. 25 (4), 401–413. Richards, M.P., Mays, S., Fuller, B.T., 2002. Stable carbon and nitrogen isotope values of bone and teeth reflect weaning age at the Medieval Wharram Percy site, Yorkshire, UK. Am. J. Phys. Anthropol. 119 (3), 205–210. Richards, M.P., Schulting, R.J., Hedges, R.E.M., 2003. Archaeology: sharp shift in diet at onset of Neolithic. Nature 425 (6956), 366. Roosevelt, A.C., 1987. The evolution of human subsistence. In: Harris, M., Ross, E.B. (Eds.), Food and Evolution: Toward a Theory of Human Food Habits. Temple University Press, Philadelphia, pp. 565–575. Ross, E.B., 1987. An overview of trends in dietary variation from hunter–gatherer to modern capitalist societies. In: Harris, M., Ross, E.B. (Eds.), Food and Evolution: Toward a Theory of Human Food Habits. Temple University Press, Philadelphia, pp. 7–55. Rouse, I., 1992. The Taínos: The Rise and Fall of the People Who Greeted Columbus. Yale University Press, New Haven. Schoeninger, M.J., DeNiro, M.J., 1984. Nitrogen and carbon isotopic composition of bone collagen from marine and terrestrial animals. Geochimica et Cosmochimica Acta 48 (4), 625–639. Schoeninger, M.J., DeNiro, M.J., Tauber, H., 1983. Stable nitrogen isotope ratios of bone collagen reflect marine and terrestrial components of prehistoric human diet. Science 220, 1381–1383. Schwarcz, H.P., 2000. Some biochemical aspects of carbon isotopic paleodiet studies. In: Ambrose, S.H., Katzenberg, M.A. (Eds.), Biogeochemical Approaches to Paleodietary Analysis. Kluwer Academic, New York, pp. 189–209. Siegel, P.E., 2010. Continuity and change in the evolution of religion and political organization on pre-Columbian Puerto Rico. J. Anthropol. Archaeol. 29 (3), 302–326. Siegel, P.E., Hofman, C.L., Bérard, B., Murphy, R., Hung, J.U., Rojas, R.V., White, C., 2013. Confronting Caribbean heritage in an archipelago of diversity: politics, stakeholders, climate change, natural disasters, tourism, and development. J. Field Archaeol. 38 (4), 376–390. Smith, B.N., Epstein, S., 1971. Two categories of 13C/12C ratios for higher plants. Plant Physiol. 47 (3), 380–384. Somerville, A.D., Fauvelle, M., Froehle, A.W., 2013. Applying new approaches to modeling diet and status: isotopic evidence for commoner resiliency and elite variability in the Classic Maya lowlands. J. Archaeol. Sci. 40 (3), 1539–1553. Sponheimer, M., Robinson, T., Ayliffe, L., Roeder, B., Hammer, J., Passey, B., West, A., Cerling, T., Dearing, D., Ehleringer, J., 2003. Nitrogen isotopes in mammalian herbivores: hair δ15N values from a controlled feeding study. Int. J. Osteoarchaeol. 13, 80–87. Steadman, D.W., Jones, S., 2006. Long-term trends in prehistoric fishing and hunting on Tobago, West Indies. Lat. Am. Antiq. 17 (3), 316–334. Steadman, D.W., Stokes, A.V., 2002. Changing exploitation of terrestrial vertebrates during the past 3000 years on Tobago, West Indies. Hum. Ecol. 30 (3), 339–367. Stokes, Anne V. 1998. A Biogeographic Survey of Prehistoric Human Diet in the West Indies Using Stable Isotopes. PhD diss., Department of Anthropology, University of Florida, Gainesville. Stokes, A.V., 2005. Ceramic-age dietary patterns in Puerto Rico: stable isotopes and island biogeography. In: Siegel, P.E. (Ed.), Ancient Borinquen: Archaeology and Ethnohistory of Native Puerto Rico. University of Alabama Press, Tuscaloosa, pp. 185–201. Tieszen, L.L., Fagre, T., 1993. Effect of diet quality and composition on the isotopic composition of respiratory CO2, bone collagen, bioapatite, and soft tissues. In: Lambert, J.P., Grupe, G. (Eds.), Prehistoric Human Bone: Archaeology at the Molecular Level. Springer, Berlin, pp. 121–155. Tütken, T., Vennemann, T.W., Pfretzschner, H.-U., 2008. Early diagenesis of bone and tooth apatite in fluvial and marine settings: constraints from combined oxygen isotope, nitrogen and REE analysis. Palaeogeogr. Palaeoclimatol. Palaeoecol. 266 (3), 254–268. Twiss, K.C., 2007. The Archeology of Food and Identity. Center for Archaeological Investigations, Southern Illinois University Carbondale, Carbondale. Van der Merwe, N.J., Vogel, J.C., 1978. 13C content of human collagen as a measure of prehistoric diet in woodland North America. Nature 276, 815–816. Van, Klinken, Dating, G.J., 1991. Dating and Dietary Reconstruction by Isotopic Analysis of Amino Acids of Fossil Bone Collagen—With Special Reference to the Caribbean. Publications Foundation for Scientific Research in the Caribbean Region, Amsterdam. Vogel, J.C., van de Merwe, N.J., 1977. Isotopic evidence for early maize cultivation in New York state. Am. Antiq. 42 (2), 238–242. Warinner, C., Tuross, N., 2009. Alkaline cooking and stable isotope tissue-diet spacing in swine: archaeological implications. J. Archeol. Sci. 36, 1690–1697. Warinner, C., Tuross, N., 2010. Brief communication: tissue isotopic enrichment associated with growth depression in a pig: implications for archaeology and ecology. Am. J. Phys. Anthropol. 141, 486–493. Weston, D.A., 2011. Human Skeletal Report: Anse Lavoutte, St. Leiden University, Lucia. Leiden. Wilson, S.M., 2007. The Archaeology of the Caribbean. Cambridge University Press, Cambridge.

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J.E. Laffoon et al. / Journal of Archaeological Science: Reports 5 (2016) 168–180

Wing, E.S., 2001. Native American use of animals in the Caribbean. In: Woods, C.A., Sergile, F.E. (Eds.), Biogeography of the West Indies: patterns and perspectives. CRC Press, Boca Raton, pp. 481–518. Wing, E.S., 2012. Zooarchaeology of West Indian land mammals. In: Borroto-Páez, R., Woods, C.A., Sergile, F.E. (Eds.), Terrestrial Mammals of the West Indies. Florida Museum of Natural History, Gainesville, pp. 341–356.

Woods, C.A., Sergile, F.E. (Eds.), 2001. Biogeography of the West Indies: patterns and perspectives, 2nd ed. CRC Press, Boca Raton. Wright, L.E., Schwarcz, H.P., 1996. Infrared and isotopic evidence for diagenesis of bone apatite at Dos Pilas, Guatemala: palaeodietary implications. J. Archaeol. Sci. 23 (6), 933–944.