The Uncertain Origins of Mesoamerican Turkey Domestication

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1 J Archaeol Method Theory DOI /s The Uncertain Origins of Mesoamerican Turkey Domestication Erin Kennedy Thornton 1 & Kitty F. Emery 2 # Springer Science+Business Media New York (outside the USA) 2015 Abstract The turkey (Meleagris gallopavo) is the only domesticated vertebrate to originate from North America. Accurate reconstructions of the timing, location, and process of its domestication are thus critical for understanding the domestication process in the ancient Americas. A substantial amount of recent research has been devoted to understanding turkey domestication in the American Southwest, but comparatively little research has been conducted on the subject in Mesoamerica, despite the fact that all modern domestic turkeys descend from birds originally domesticated in Mexico during pre-colonial times. To address this disparity, we have conducted a review of the available literature on early turkeys in the archaeological record of Mesoamerica. We evaluate the evidence in terms of its accuracy and use this evaluation as a stepping off point for suggesting potential avenues of future research. Although the lack of available data from Mesoamerica currently precludes detailed cross-cultural comparisons, we briefly compare the origins and intensification of turkey rearing in Mesoamerica with the American Southwest to generate more dialogue among researchers independently studying the topic in these two distinct but interconnected cultural regions. Keywords Turkeys. Domestication. Mesoamerica. Maya. Preclassic Within Mesoamerican society, turkeys were both mundane food resources and symbolically charged ritual animals commonly used in ceremonial feasts and as sacrificial offerings (Pohl and Feldman 1982; Tozzer and Allen 1910). Turkey feathers were incorporated into headdresses, capes, and fans, and bones and other by-products were * Erin Kennedy Thornton erin.thornton@wsu.edu 1 2 Kitty F. Emery kemery@flmnh.ufl.edu Department of Anthropology, Washington State University, College Hall 150, PO Box , Pullman, WA , USA Florida Museum of Natural History, University of Florida, Gainesville, FL , USA

2 Thornton and Emery used to produce medicines, tools, and musical instruments (Corona Martínez 2005, 2008; Pohl 1983). The turkey(meleagris gallopavo) is also the only domestic vertebrate to originate from North America. Its history of use and domestication is thus of great importance to understanding the process and timing of New World animal domestication. Although recent research in the American Southwest has provided many details regarding the history and practice of turkey husbandry in this region (e.g., Badenhorst et al. 2012; McCafferyet al. 2014; McKusick2001; Munro 2006, 2011; Newbold et al. 2012; Rawlings and Driver 2010; Spelleret al. 2010), comparatively little research has been conducted on the subject in Mesoamerica despite the fact that all modern domestic turkeys descend from birds originally domesticated in Mexico during pre-colonial times (for exceptions, see Monteagudo et al. 2013; Speller et al. 2010; Thorntonet al. 2012). To address this disparity, we conducted a review of the published and available unpublished archaeological evidence for early turkey use in the region. We use this information to assess the state of our current understanding of Mesoamerican turkey domestication and to suggest potential avenues for future research to document the timing, location, and process of turkey domestication in Mesoamerica. Identifying when, where, and how turkeys were domesticated is critical to reconstructing pre-colonial Mesoamerican subsistence systems and why ancient populations decided to adopt animal management. Careful comparisons must also be drawn between the process, timing, and spread of turkey domestication in Mesoamerica and the American Southwest to fully understand animal domestication in ancient North America. Our work provides a first step toward this goal by evaluating our current state of knowledge and both the hurdles and opportunities that remain. Mesoamerican Turkey Species Taxonomy and Description Two species of wild turkeys were present in pre-hispanic Mesoamerica: the smallerbodied and more colorful ocellated turkey 1 (Meleagris ocellata) native to the northern half of the Maya cultural region, and the wild turkey (Meleagris gallopavo), originally found throughout much of central and northern Mexico and into the continental USA (Fig. 1). Of the six currently recognized subspecies of wild turkey, three range into Mesoamerica (M. g. mexicana, M. g. intermedia, and M. g. gallopavo). Genetic analysis confirms that the southern Mexican subspecies (M. g. gallopavo) gave rise to the domestic turkeys bred and reared throughout the world today (Monteagudo et al. 2013). Genetic evidence also supports the independent pre-columbian domestication of at least one other subspecies of wild turkey (M. g. intermedia and/or silvestris)in the American Southwest or an unidentified region to the east (Speller et al. 2010). Domestic turkeys from the Southwest do not contribute to the genetic stock of modern domestic turkeys (Speller et al. 2010). indicating that it is the timing of dissemination of the southern Mexican subspecies (M. g. gallopavo) that is of most importance to understanding the origins of modern domestic turkeys. 1 In Ornithology, standardized common names of species are capitalized.

3 The Uncertain Origins of Mesoamerican Turkey Domestication Fig. 1 Modern geographic ranges of Mesoamerican turkey species (The IUCN Red List of Threatened Species. Version ). The southeastern corner of the M. gallopavo range was expanded slightly to conform to other modern range descriptions (Howell and Webb 1995; Navarro and Peterson 2007) Unlike the wild turkey, the ocellated turkey was never domesticated (Schorger 1966:6). Instead, it served as an important wild game species for both ancient and modern hunters throughout Mexico s Yucatan Peninsula and northern Belize and Guatemala. Although this species is not classified as domesticated, it may have been reared in captivity at certain sites by ancient Maya populations (Hamblin 1984; Masson and Peraza Lope 2008; PohlandFeldman1982; Pollock and Ray 1957). Living wild and ocellated turkeys are distinguished by their body size, and plumage. Ocellated turkeys are more colorful with iridescent bronze, green, and blue body feathers, deep red legs, and bright blue heads covered with large orange and red nodules or carnucles. Ocellated turkeys also lack the long clusters of hair-like feathers known as beards, which characterize adult male wild turkeys. Plumage differences include the ocellated turkeys broader white bands on the outer edges of the secondary flight feathers, and the presence of eye-shaped spots on the tail feathers. Ocellated turkeys are also generally smaller than wild turkeys, with average adult hens weighing kg and adult males between 4.3 and 5 kg. In contrast, female and male wild turkeys weigh kg and kg, respectively (Leopold 1959). Although the natural geographic ranges of wild M. g. gallopavo and M. ocellata do not overlap, the diffusion of domesticated wild turkeys to the Maya cultural region brought the two turkey species into coexistence. Determining when and where humans caused the ranges of M. g. gallopavo and M. ocellata to overlap in the Maya region is significant because the two species of turkey may be able to hybridize (Schorger 1966:67; Williams et al. 2010:11). However, many reports of hybridization are anecdotal and hybridization appears uncommon (Leopold 1948:399). It is currently unclear whether hybridization occurred in the past, but the topic warrants further investigation, especially in the Maya region where both species may have been raised together in captivity.

4 Thornton and Emery Challenges to Identifying Turkey Domestication in the Archaeological Record One challenge to reconstructing the process of Mesoamerica turkey domestication is the dispersed nature of much of the current zooarchaeological literature relevant to the subject. From the current literature, it is also difficult to generate accurate temporal and spatial comparisons. We discuss additional methodological and conceptual challenges and considerations relevant to turkey domestication in the following sections as background for our assessment of the current state of knowledge, and our suggestions for future research. Morphometric and Genetic Markers of Domestication Recent reviews emphasize that animal domestication is a prolonged process of humananimal mutualism that results in a continuum of states from wild to fully domestic (Zeder 2006). Along this continuum are various levels of human control over a species movement, diet, and reproduction ranging from taming and confinement to directed breeding. Over time, this relationship often results in morphological and genetic changes within a species, but these changes typically do not all occur at once and instead appear at different points throughout the domestication process (Zeder 2006). In some cases, genetic and morphological changes do not occur at all. This is especially true early in the domestication process or when there is ongoing breeding between wild and captive populations of a species (Munro 2011). To date, morphological studies have been unable to distinguish between wild and pre-colonial domestic forms of M. gallopavo (Breitburg 1988; Munro 2006). Genetic markers for Mesoamerican domestic turkeys are also unknown, although some distinctions are now possible among American Southwest turkeys (Speller et al. 2010). In the absence of direct morphological and genetic markers for Mesoamerican turkey domestication, it is difficult to identify early examples of tamed or captive reared turkeys that appear within the natural range of the species wild progenitor. Documenting turkey domestication throughout the Americas is further complicated by the fact that wild turkey hunting likely occurred alongside domestic rearing in both the American Southwest and Mesoamerica (Götz 2008; Hamblin 1984; McCaffery et al. 2014; Speller et al. 2010). Indirect Evidence of Husbandry: Abundance, Demographics, Pathology, Diet, and Penning The current lack of identified morphometric and genetic markers for domestication requires archaeologists to rely on indirect evidence of turkey husbandry such as species abundance, demographic profiles, paleopathology, dietary shifts, and the presence of pen structures. For example, overall turkey abundance at a site in comparison to wild taxa may be used to infer husbandry although such increases may not occur until late in the domestication process (Zeder 2006). Instead of being immediately used as staple resources, domesticates may serve as occasional insurance against scarcity, or as feasting or ceremonial items to be harvested when needed. This pattern appears to be typical of the North American transition to horticulture, which is characterized by a long period of low dependence on domesticated

5 The Uncertain Origins of Mesoamerican Turkey Domestication species alongside continued use of wild resources (Piperno 2011). Consequently, turkey husbandry may have been in practice long before we observe increasing abundances. Demographic (age and sex) profiles have also been cited as evidence for turkey husbandry (Badenhorst et al. 2012; Fothergill 2012). Very young individuals (e.g., poults) are not expected to appear in the population of turkeys obtained through hunting, which typically targets large adult individuals. Similarly, the recovery of eggshells may indicate that turkeys were born on-site. To differentiate between onsite laying and wild egg collection, researchers can now identify eggshells that hatched versus those that broke early in development (Beacham and Durand 2007; Lapham et al. 2013b). Other demographic features of domestic flocks include the culling of young or subadult males, but age and sex profiles have not been widely used due to the difficulty of accurately assigning age and sex to fragmentary and subadult bird remains. The use of domestic turkeys for ceremonial purposes or feather production (Fothergill 2012; McKusick 1974) could also result in different culling or demographic profiles. This suggests that researchers also need to focus on reconstructing the role turkeys played in Mesoamerican society through time to better understand expected patterns of flock management and size. Healed fractures and other pathologies are an additional line of evidence that may be used to infer captive turkey rearing. Besides indicating human care or protection of injured birds, repeated pathologies on specific elements, such as the ulna, may indicate repeated feather harvesting (Fothergill 2012). To date, paleopathologies have only been reported for pre-colonial turkeys from the American Southwest (Akins 1985; Fothergill 2012; McKusick 1974; Steen 1966) and Post- Medieval Britain (Fothergill 2014). In comparison to the American Southwest, the indirect evidence for turkey domestication in Mesoamerica is relatively meager, due in part to taphonomy and archaeological preservation. In the Southwest, the record of turkey domestication includes finds such as whole mummified turkeys, turkey coprolites, feathers, eggshells, and fragile juvenile skeletal remains (e.g., Arakawa et al. 2001; Breitburg 1988; Lang and Harris 1984; Lipe 1979; Olsen 1990; Reed 2006). Equivalent finds are rare to nonexistent in Mesoamerica although eggshell has been recovered from a few sites (Lapham et al. 2013b; Storey 1992:64; Widmer 1987:346). Turkey pens, such as those found at sites including Mesa Verde and Paquimé (Casas Grandes) (Breitburg 1993) are also more common in the Southwest than farther south in Mesoamerica. The presence or absence of turkey pens in both regions warrants further investigation as it may be indicative of rearing practices, and whether turkeys were confined to pens or raised using free-range techniques as in many rural Central American villages today (Mallia 1999). Regardless of whether turkeys were free-range or penned, their residence within human habitation areas likely resulted in a dietary shift for the birds that may be documented through stable carbon isotope (δ 13 C) analysis of their remains. In both Mesoamerica and the American Southwest, much of the animal fodder and human food waste would be maize (Zea mays), the most important staple crop cultivated in the region. Maize utilizes a C4 photosynthetic pathway which makes it isotopically distinct from most other foods consumed by turkeys (Leopold 1959:273; Rawlings and Driver 2010:2435). Tame or captive reared turkeys should therefore have higher carbon isotope ratios (δ 13 C) than wild foraging turkeys. Significantly, this method has good potential to identify the earliest stages of turkey domestication

6 Thornton and Emery since a change from a wild C3 diet to human-provided fodder high in maize likely occurred very early in the domestication process before genetic or morphological features would appear. This method has successfully distinguished between wild and domestic turkeys in the American Southwest (McCaffery et al. 2014; Rawlings and Driver 2010). but its potential has not yet been realized in Mesoamerica (Thornton and Emery 2014). Geographic Range Reconstructions The appearance of turkeys outside their natural geographic ranges has also been used to infer that they were under some level of human control at the time these range shifts or introductions occurred. The pre-colonial ranges of wild turkeys throughout the Americas, however, are not always clear since modern ranges have been shaped and modified by several centuries of human hunting and landscape alteration (Mock et al. 2002:653; Newbold et al. 2012). Within Mesoamerica, it is especially difficult to reconstruct the past range of wild turkeys since wild populations of M. g. gallopavo have been largely extirpated over the last several centuries (Schorger 1966). Within two classic natural history publications there also has been some disagreement over the past range of wild turkeys. The map generated by Leopold (1959:269) only extends the historic (post-conquest) range of M. gallopavo as far south as the Río Balsas Valley near the modern border between the Mexican states of Guerrero and Michoacán, and excludes areas around the Valley of Mexico (Fig. 2). Leopold s range map is primarily based on modern and historic (post-conquest) evidence, and he acknowledges that wild turkeys may have been extirpated from certain areas, such as the Valley of Mexico, prior to colonial times (Leopold 1959:270). A later publication by Schorger (1966:49) disagrees with Leopold s original range map and extends the historic and pre-columbian range of M. gallopavo throughout central Mexico as far south as southern Guerrero, central Veracruz, and the Puebla-Oaxaca border (Fig. 2). In his revised map, Schorger plots known locations from both published literature and collected biological specimens to support this farther southern range expansion. Significantly, Schorger s map conforms to both previous and more recent reports of the natural range of M. gallopavo, which includes the pine-oak upland forests extending from Jalisco and northern Veracruz through Guerrero and into northern Oaxaca (Camacho-Escobar et al. 2011; Clementset al. 2014; Friedmannet al. 1950; Navarro and Peterson 2007). In the archaeological literature, this debate has led to some confusion regarding the interpretation of the region s zooarchaeological record. Some researchers argue for the wild turkey s natural absence from areas such as the Valley of Mexico, based on Leopold s (1959) more restricted range map and a lack of Pleistocene and Archaic period turkey specimens from the region (Valadez Azúa and Arrellín Rosas 2000:314). However, the Pleistocene record of Mexican bird species is far from complete and nearly all of the bird taxa found in paleontological assemblages from the Valley of Mexico are aquatic species due to the fossil sites location on the shores of an ancient lake system (Corona Martínez 2002b). Support for Schorger s suggested nonanthropogenic presence of turkeys in and around the Valley of Mexico comes from the mention of both wild and domestic turkeys in the

7 The Uncertain Origins of Mesoamerican Turkey Domestication Fig. 2 Comparison of Meleagris gallopavo range reconstructions (shaded) in Mexico: a Leopold (1959:269, Fig. 102); b Schorger (1966:49, Fig. 8) ethnohistoric accounts of Sahagún (1963:29, 53), and the overlooked identification of turkey remains in late Pleistocene/early Holocene pre-occupation levels at the site of Cuanalan (Manzanilla 1985:135) and Archaic period ( B.C.) deposits at Texcal Cave near modern-day Puebla (Alvarez 1975). Until further evidence emerges, it may be safest to assume that the wild progenitor of the domestic turkey conforms to modern descriptions and ranged from northern Mexico through southern Guerrero and eastward to include Puebla and northern Veracruz (Clements et al. 2014). Determining whether turkeys identified at central Mexican archaeological sites were domestic versus wild will therefore require additional lines of evidence beyond the mere presence of the species in human settlements.

8 Thornton and Emery Distinguishing Between Wild and Ocellated Turkeys Although living wild and ocellated turkeys are not easily confused, archaeologists must differentiate the two species based solely on their very similar skeletal remains. Distinguishing between the two species in archaeological deposits is a particular challenge in the Maya cultural region where ocellated turkeys occur naturally and domesticated turkeys (M. g. gallopavo) were introduced during pre-colonial times. Taphonomic factors, a lack of appropriate comparative skeletal specimens, and confusion over what skeletal features characterize each species often precludes identification of turkey remains to the species level (Bochenski and Campbell 2006;Steadman 1980). Morphological identifications are further complicated by the fact that hunted game also included other species of large galliforms within the family Cracidae (e.g., great curassow Crax rubra), which are similar in size and gross morphology to turkeys. Consequently, all large galliform bird remains at Mesoamerican archaeological sites should be identified with care. Accurately distinguishing between the two species is important for determining when domesticated turkeys diffused to the Maya area, how wild and domestic turkeys may have been used differently, and whether the ancient Maya maintained captive, or tame populations of the locally available ocellated turkey. Ocellated turkey husbandry has been suggested for several Postclassic Maya sites including Mayapan and Cozumel in Mexico s northern Yucatan Peninsula. On the offshore island site of Cozumel, rearing is inferred based on the presence of pen structures and immature individuals, as well as the fact that turkeys do not naturally occur on the island (Hamblin 1984:93). Animal pens are also present at the site of Mayapan (Masson and Peraza Lope 2008). but evidence of ocellated turkey rearing derives primarily from the large quantity of turkey remains recovered at the site, as well as the fact that many of the individuals are larger and stouter than typical ocellated turkeys, which may imply captive rearing and intentional feeding (Breitburg 1988; Pollock and Ray 1957; Steadman 1980:150). Determining whether the Maya maintained captive populations of ocellated turkeys may reveal much about the process and diffusion of turkey domestication. If the ancient Maya husbanded both ocellated and introduced domesticated wild turkeys, it indicates that the Maya independently experimented with turkey management. In contrast, if the Maya only reared the introduced domestic turkeys, it suggests that turkey husbandry was integrated into the economy primarily through the adoption of a non-local domesticated species. Ocellated turkeys may have been reared alongside or instead of introduced domestic turkeys due to some cultural preference related to plumage color, size, symbolism, or some other attribute. Both zooarchaeological data and ethnohistoric accounts confirm that Mesoamerican cultures experimented with the captive rearing of wild species such as macaws (Ara sp.), parrots (Psittacidae), rabbits (Sylvilagus sp.), deer (Odocoileus virginianus), and peccaries (Tayassuidae) (Corona Martínez 2002a, 2013; Hamblin 1984; Valadez Azúa 1993, 2003; White et al. 2004). The extent of these practices is unclear, as is whether the captive animals were maintained primarily for subsistence or elite display and ceremonial purposes. Regardless, ocellated turkey husbandry would not be out of place within this cultural framework. The question of ocellated turkey husbandry thus

9 The Uncertain Origins of Mesoamerican Turkey Domestication warrants further investigation as it relates to larger questions regarding the function, extent, and meaning of managed animal resources in the Mesoamerican economy. The Origins of Mesoamerican Turkey Domestication: an Assessment of Current Evidence Central Mexican Highlands A Preclassic period turkey (M. g. gallopavo) found at the Tehuacan Valley site of Coxcatlan Cave (Flannery 1967)(Fig.3) is typically, and often uncritically, cited as the earliest example of a domestic turkey in Mesoamerica. At this site, a single bone of M. gallopavo was dated to the earlier part of the Palo Blanco phase ( A.D. 180) (Flannery 1967:155, 163) (Table 1). The bird was interpreted as a domestic turkey based on Leopold s (1959) range map, which does not extend the historic range of the wild turkey as far south as the Tehuacan Valley. No other evidence for turkey domestication was observed at the site, although the absence of turkeys in earlier site deposits was cited in support of the turkey s nonlocal origin (Flannery 1967:155). Turkeys are not the only species with a somewhat patchy temporal distribution in the Tehuacan Valley deposits so their absence in earlier archaeological deposits is suggestive, but not conclusive evidence that turkeys are nonlocal to the region. Although the Coxcatlan Cave turkey may very well represent an early example of introduced or diffusing domestic turkeys, we cannot completely rule out the presence of wild turkey populations within or near the Tehucan Valley during ancient times since the valley lies at the southern edge of the wild turkey s currently accepted geographic range. Although not commonly cited, earlier examples of Preclassic turkeys have been reported from archaeological sites within and around the Valley of Mexico and nearby Morelos (Alvarez 1975, 1976; Corona Martínez 2006; Grove 1974; Manzanilla 1985; Fig. 3 Map of Mesoamerica showing major cultural areas and sites mentioned in text

10 Thornton and Emery Table 1 Summary of earliest turkeys (Meleagris gallopavo) identified in Mesoamerican faunal assemblages organized chronologically by region Site (location) Time period (dates) Quantity: NISP (%*)/MNI (%*) Context Wild vs. domestic classification and evidence for domestication noted in publications Citation Central Mexican Highlands: Cuanalan (Estado de Mexico) Pleistocence (upper parts of precultural sequence) Texcal Cave (Puebla) Archaic ( B.C.) Tlatilco (Estado de Mexico) Middle Preclassic ( B.C.) Nexpa (Morelos) Middle Preclassic ( B.C.) Oaxtepec Km 27.5 (Morelos) Middle Preclassic ( B.C.) Cuicuilco (Distrito Federal) Middle/Late Preclassic (700 B.C. A.D. 150) Terremote-Tlaltenco (Distrito Federal) Temamatla (Estado de Mexico) Late Preclassic ( B.C.) Late Preclassic ( 400 B.C.) Present (quantity unknown) Pre-cultural No comments presumed wild Manzanilla (1985:135) 3 NISP (<1 %) Stated as unknown wild vs. domestic Alvarez (1975) 4NISP(<1%) 1inahumanburial; 3 in truncated conical pits Present (quantity unknown) 46 NISP /1 MNI (all from single partial turkey skeleton; burned) At least 1 MNI (from partial skeleton) No comments regarding wild vs. domestic status (Alvarez 1976). Classified as domestic by Valadez and Arrellín (2000: 313) Domestic/living areas No comments regarding wild vs. domestic status Alvarez (1976:7 8); Alvarez and Ocaña (1999). Valadez and Arrellín (2000) Grove (1974:42) Human burial Stated as unknown wild vs. domestic Corona Martínez (2006) Human burial No comments regarding wild vs. domestic status. Classified as domestic by Valadez and Arrellín (2000: 313) 43 NISP (14 %) Some individuals interred as complete or semicomplete burials/offerings Suggested to be domestic/captivereared based on abundance, and presence of males, females, and juveniles (Serra Puche and Valadez Azúa 1985:195). Classified as domestic by Valadez and Arrellín (2000: 313) Alvarez and Ocaña (1999). Valadez and Arrellín (2000) Serra Puche and Valadez Azúa (1985). Valadez and Arrellín (2000). Alvarez and Ocaña (1999) 6MNI(2%) ValadezAzúaandRamirez (1991). Valadez and Arrellín (2000)

11 The Uncertain Origins of Mesoamerican Turkey Domestication Table 1 (continued) Site (location) Time period (dates) Quantity: NISP (%*)/MNI (%*) Context Wild vs. domestic classification and evidence for domestication noted in publications Citation Cuanalan (Estado de Mexico) Late Preclassic ( B.C.) Texcal Cave (Puebla) Late Preclassic ( 200 B.C.) Coxcatlan Cave (Tehuacan Valley, Puebla) Oaxaca: Late Preclassic ( A.D. 180) San Jose Mogote Early/Middle Preclassic ( B.C.) Tayata Early Formative ( 900 B.C.) Nochixtlan Valley Middle/Late Preclassic ( B.C.) Gulf Coast: La Joya (Veracruz, MX) Early Preclassic ( B.C.) Present (quantity unknown) 1 individual from human burial; others from unspecified contexts No comments regarding wild vs. domestic status (Valadez Azúa and Ramirez 1991). Classified as domestic by Valadez and Arrellín (2000: 313) No comments regarding wild vs. domestic status (Manzanilla 1985: ). Classified as domestic by Valadez and Arrellín (2000: 313) 7 NISP (1 %) Stated as unknown wild vs. domestic Alvarez (1975) 1NISP(<1%)/1 MNI (3 %) 4 NISP (<1 %) 2 from a high status residence (living area); 2 from a neighborhood midden Interpreted as an imported domestic turkey Interpreted as wild turkeys. Suggestedtorepresentdried specimens imported for feathers from central Mexico 1NISP(<1%) Specimennotidentifiedto species (could be either M. gallopavo or M. ocellata). No comments regarding wild vs. domestic status 1 % NISP No comments regarding wild vs. domestic status 2NISP(<1%)/1 MNI (4 %) No comments regarding wild vs. domestic status Manzanilla (1985: ); Valadez and Arrellín (2000) Flannery (1967:155,163,175) Flannery and Marcus (2005: 188, 245,251): Lapham et al. (2013b) Lapham et al. (2013a) Breitburg (1988:33, 73) citing unpublished data by Spores. Vanderwarker (2006). Peres et al. (2013)

12 Thornton and Emery Table 1 (continued) Site (location) Time period (dates) Quantity: NISP (%*)/MNI (%*) Context Wild vs. domestic classification and evidence for domestication noted in publications Citation La Joya (Veracruz, MX) Late Preclassic (400 B.C. A.D. 100) Bezuapan (Veracruz, MX) Late Preclassic (400 B.C. A.D. 300) Santa Luisa (Veracruz, MX) Late Preclassic Early Classic (300 B.C. A.D. 300) Patarata 52 (Veracruz, MX) Late Preclassic Early Classic (300 B.C. A.D. 300) Maya Region: El Mirador (Petén, Guatemala) Late Preclassic (327 B.C. A.D. 54) Zaculeu Early/Late Classic (A.D ) 1NISP(<1%)/1 MNI (2 %) 11 NISP (1 %)/3 MNI (6 %) No comments regarding wild vs. domestic status No comments regarding wild vs. domestic status 3 MNI (4.4 %) No comments regarding wild vs. domestic status Vanderwarker (2003; 2006). Peres et al. (2013) Vanderwarker (2003; 2006). Peres et al. (2013) Wing (1978: 35) 3 MNI (6.3 %) Stated as unknown wild vs. domestic Wing (1977: 210),(1978: 34) 5NISP(<1%)/3 MNI (4 %) Construction fill within site ceremonial core 2 NISP Tomb bones found inside bowls Identified as domestic/captive-reared based on carbon isotope values indicative of high maize consumption Bones were identified by Dr. Alexander Wetmore (Smithsonian Institution) Thornton et al. (2012). Thornton and Emery (2014) (Woodbury and Trik 1953: 87; 278) *Percent (%) of entire faunal assemblage

13 The Uncertain Origins of Mesoamerican Turkey Domestication Serra Puche and Valadez Azúa 1985; Valadez Azúa and Arrellín Rosas 2000; Valadez Azúa and Ramirez 1991)(Table1,Fig.3). Many of these sites (Caunalan, Cuicuilco, Tlatilco, Temamatla, and Terremote-Tlatenco) are listed among those with evidence for turkey domestication in a recent synthesis of the region s zooarchaeological data (Valadez Azúa and Arrellín Rosas 2000:313). In this assessment, evidence for domestication at these sites comes from high turkey abundance (at Temamatla and Terremote-Tlatenco, Valadez Azúa and Arrellín Rosas 2000:314), the presence of subadults, and a lack of observed morphological differences between the archaeological specimens and modern domestic turkeys (Valadez Azúa and Arrellín Rosas 2000: ). The original reports from each site only allow us to confirm relatively high turkey abundance at the site of the Terremote-Tlaltenco where turkeys composed over 13 % of the site s number of identified specimens (NISP) (Serra Puche and Valadez Azúa 1985). At all other sites, turkeys were either equal to or less abundant than other wild bird species such as ducks (Anatidae) (Alvarez 1976; Manzanilla1985; Valadez Azúa and Ramirez 1991; Valadez Azúa and Arrellín Rosas 2000). Subadult turkeys and eggshell are also reported from Terremote-Tlaltenco, but the published images in the original report included no clear examples of subadult remains (Serra Puche and Valadez Azúa 1985:196, photo 6). The eggshell, although presented in the site report s figures (Serra Puche and Valadez Azúa 1985:206, 210, 211) is not discussed in the accompanying text, and it is unclear what species the eggshell belongs to since multiple bird species were found at the site. Valadez Azúa and Arrellín Rosas (2000) do not appear to have assessed the morphological details of the skeletal specimens identified by the original authors as morphologically similar to modern domestic turkeys, and no images were available in the site reports to allow us to do so. However, since previous studies of turkey morphology (Breitburg 1988;Munro2011)havefailedto identify clear morphological distinctions between wild and domestic turkeys, the lack of distinguishing characters for domestication at Terremote-Tlaltenco and other sites is inconclusive. Although it is possible that the Middle to Late Preclassic sites from the Valley of Mexico and Morelos contain domesticated turkeys, the current evidence requires further investigation of the original materials. In the absence of clear evidence to support geographic displacement, or morphometric changes, additional evidence related to penning, dietary shifts, or the presence of juvenile turkeys will be needed to confirm that the presence of tame or domesticated turkeys. Oaxaca, Southern Gulf Coast and Maya Region Small numbers of Preclassic turkey remains have also been reported from sites falling well outside the widely accepted natural range of wild M. gallopavo (Clements et al. 2014). The significance of these finds to interpreting the timing, location, and diffusion of Mesoamerican turkey domestication has not yet been adequately considered. Based on their geographic displacement, these specimens lend support to turkeys being under some degree of human control during the Preclassic. This line of evidence, however, is not without its faults as wild animals were also traded, either dead or alive, throughout Mesoamerica during pre-hispanic times (e.g., Lopez Luján 2005; Thornton 2011). We evaluate the Preclassic evidence for turkey rearing or trade from southern Mesoamerica in the following paragraphs.

14 Thornton and Emery In central Oaxaca, just south of the species presumed southern range limit, turkeys have been identified in Preclassic deposits at San Jose Mogote, Tayata and the Nochixtlan Valley (Breitburg 1988; Flannery and Marcus 2005; Laphamet al. 2013a) (Table1). At each site, turkeys are very rare (NISP <5). Based on such low abundance and in the absence of other evidence for on-site rearing such as pens, it is possible that these turkeys represent trade goods rather than established breeding populations. Flannery and Marcus (2005:96, 188) specifically suggest that the turkeys at San Jose Mogote represent dried birds imported for their feathers from central Mexico. Alternate explanations include the onsite rearing of captive/domestic turkeys, the import of live turkeys from Central Mexico, or the acquisition of wild or domestic turkeys from closer source populations. Turkey bones were also present at two nonelite residential Gulf Coast Olmec sites (La Joya and Bezuapan) dating to the Early and Late Preclassic, but their status as wild versus domestic, or their potential nonlocal origin is not discussed by the analysts (Peres et al. 2013; Vanderwarker 2003, 2006) (Table1). The Olmec area is a wet subtropical broadleaf forest ecoregion very different from the upland pine-oak forests known to provide suitable habitat for wild M. g. gallopavo. The turkey remains from these sites thus represent probable examples of imported or introduced turkeys, with the presence of wing, sternum, and distal leg elements suggesting on-site rearing or transport of whole animals. Since turkey remains are absent at the much larger and higher status lowland Olmec sites of Tres Zapotes and San Lorenzo (Peres et al. 2013). M. gallopavo appears to have been present, but not widespread in the southern Gulf Coast region during the Preclassic. Another clear example of Preclassic turkey diffusion, transport, or trade comes from the site of El Mirador located in the central Maya Lowlands of Petén, Guatemala (Table 1). This site lies far outside the natural range and habitat requirements of wild M. gallopavo. At El Mirador, the incomplete remains of at least three M. gallopavo individuals were found in deeply buried, sealed contexts within structures surrounding the site s second largest temple pyramid. The M. gallopavo specimens represent at least one female and two males, which were distinguished from local ocellated turkeys based on combined morphological, osteometric, and ancient DNA evidence (Thornton et al. 2012). The El Mirador turkeys are currently the earliest examples of M. gallopavo identified in the Maya region as all other confirmed examples date to the much later Postclassic period (A.D ). To date, the only other suggestion of domestic turkeys in the Maya area prior to the Postclassic come from unconfirmed M. gallopavo specimens identified in a Classic period (A.D ) elite tomb at the highland Guatemala site of Zaculeu (Table 1) (Woodbury and Trik 1953:278). Comparing Mesoamerican and Southwest Origins of Turkey Domestication As reviewed above, the earliest evidence for Mesoamerican turkey domestication currently relies primarily on the identification of anthropogenic range expansions and nonlocal introductions. This is due to the absence of recognized morphological and genetic features that distinguish wild from domestic Mesoamerican turkeys, and a lack of Preclassic sites with clear evidence of on-site rearing such as the presence of pen

15 The Uncertain Origins of Mesoamerican Turkey Domestication structures, juveniles, and eggshells. An exception to this may be the site of Terremote- Tlaltenco in the Valley of Mexico ( B.C.) where the authors report that eggshells and young turkeys may be present (Serra Puche and Valadez Azúa 1985). The species presence farther south in Mesoamerica more clearly indicates that the process of turkey domestication was underway by at least the Late Preclassic. Even earlier beginnings (in the Early or Middle Preclassic) are suggested by the species presence at sites along the southern Gulf Coast and in central Oaxaca, which are contemporary with the earliest turkeys in and around the Valley of Mexico. More research is needed, but the practice of turkey husbandry appears to have extended outside of central/northern Mexico fairly early in time, suggesting either a rapid cultural diffusion, or an early date of domestication. Our review of current evidence reveals that the earliest evidence for turkey domestication likely predates the Tehuacan Valley specimens (Flannery 1967)mostcommonly cited. It further reveals that the timing and location of Mesoamerican turkey domestication is still largely unknown, and that many of our present interpretations are based on incomplete datasets. Nevertheless, it appears that turkey husbandry was practiced at a fairly small scale throughout the Preclassic in many different parts of Mesoamerica, but it may not have formed part of the ritual or subsistence economy at each and every site. The timing and pattern is similar to the American Southwest where sparse evidence for turkey rearing appears across a fairly broad geographic area around 300 B.C. (Munro 2011: ). In the Southwest, some researchers argue that this prolonged period of early low-level husbandry reflects the initial use of domestic turkeys primarily as sources of feathers or as ritual offerings rather than as a meat resource (Breitburg 1988; McKusick 1986). The evidence for this comes from ethnohistoric descriptions, the recovery of feathers and feathered artifacts (e.g., blankets and feather wrapped cordage), and the absence of turkeys from general midden deposits dating to before the Pueblo II III periods (A.D ) (Muir and Driver 2002; Badenhorst and Driver 2009). In Mesoamerica, evidence of turkey use is primarily limited to more durable skeletal remains, but feathers appear frequently in iconography as part of headdresses and other regalia. The stylized depictions of the feathers, however, preclude the identification of the plumes as specifically belonging to turkeys, and other species such as quetzals (Pharomachrus mocinno) and macaws (Ara sp.) are clearly represented. Many site reports do not provide detailed contextual data, but the available information notes that Preclassic Mesoamerican turkeys were found in ritual contexts (i.e.,offerings and burials) as well as in middens and construction fill with some bones exhibiting evidence of burning or butchery (Alvarez and Ocaña 1999; Breitburg1988:73; Corona Martínez 2006; Flannery and Marcus 2005; Thornton et al. 2012; Valadez Azúa and Arrellín Rosas 2000). This suggests that turkeys were likely used as ritual offerings, as well as for food, similar to many other animals in Mesoamerica where ritual and dietary uses of species are rarely mutually exclusive. This contrasts with the archaeological record from the Southwest, which more strongly supports nondietary use in earlier time periods (Munro 2006). In Mesoamerica, it is possible that domestic turkeys fulfilled a role similar to that of domestic dogs (Canis lupus familiaris), which were used extensively during the Preclassic for both dietary and ritual purposes (Clutton-Brock and Hammond 1994; Wing 1978), and perhaps primarily for the ceremonies and feasts related to elite display and power negotiations (Emery et al. 2013; Rosenswig 2007).

16 Thornton and Emery Therefore, although turkeys were consumed during this early period of population growth, the scarcity of their remains suggests that they were not yet a dietary staple. Instead, they may have been kept on-hand to serve as occasional insurance against scarcity, or as components of elite ceremonial, feasting and status display activities. Classic to Postclassic Expansion of Turkey Husbandry Substantial increases in Mesoamerican turkey use, as evidenced by their proportional abundance in faunal assemblages, did not occur until the Classic and Postclassic periods (A.D ). A long delay between initial domestication and later intensification also occurred in the American Southwest where turkey rearing increased in intensity after A.D. 900 (Pueblo II III periods). In the Southwest, this increase may be attributed to resource depression associated with human population increases and aggregation (Badenhorst and Driver 2009; Kohler et al. 2012; Muir and Driver 2002; Munro 2011). Although large game (e.g., deer) depletion could have motivated greater dependence on domesticated turkeys in Mesoamerica (Lapham et al. 2013b). turkey use does not expand in all cases of major population increases. For example, in central Mexico, population size and aggregation increased substantially during the Early Classic (A.D ) with the rise of Teotihuacan (Cowgill 1997). Although turkeys were used for both dietary and ritual purposes at Teotihuacan, they were less abundant than other taxa such as deer, dogs, and rabbits (Starbuck 1975, 1987; Valadez Azúa 1993). Domestic turkey abundance may have been relatively low because they formed part of a suite of managed species used in northern Mesoamerica for dietary and/or ritual purposes. Domestic dogs are very common in Classic and Postclassic faunal assemblages from central Mexico, and rabbits may have been occasionally reared in captivity (Lapham et al. 2013b; Manzanilla 1996; Valadez Azúa 1993). Conversely, wildlife resources may not have been strained to the point of widespread local scarcity around Teotihuacan, or wild game could have been imported into the site from outlying areas. Regardless, the Teotihuacan residents did not invest in intensive turkey rearing as a food supplement for their growing populations. Similarly, we do not see a sudden adoption of turkey husbandry in the Maya region during human population peaks in the Late to Terminal Classic, even though largebodied prey populations may have been locally depleted around some sites (Emery 2007). Instead, turkey use does not increase in the Maya area until the Postclassic ( A.D ), when populations were substantially reduced in most areas (Emery 2004a:47). The species early presence at El Mirador (Thornton et al. 2012) suggests that the Maya were aware of turkey domestication prior to the Postclassic even if they chose not to adopt it. In the Maya area, the availability of wild fauna, including the local Ocellated Turkey, could have delayed the widespread adoption of nonlocal domestic turkeys (Valadez Azúa and Arrellín Rosas 2000:315). The species later, more widespread adoption in the Postclassic may relate to the nature of the Postclassic Maya economy which is characterized by increased long-distance exchange between central Mexico and the Maya area, and expansion of maritime trade routes around the Yucatan Peninsula between the Gulf of Mexico and Central America s Caribbean coast (Guderjan and Garber 1995; McKillop and Healy 1989). Increased Postclassic exchange throughout Mesoamerica could have facilitated the dispersal of domesticated turkeys to the Maya area through repeated introductions of breeding pairs and

17 The Uncertain Origins of Mesoamerican Turkey Domestication transmission of rearing information. In contrast, the rare earlier introduction of the bird might not have been sufficient to fully incorporate the species into the Maya economy. The adoption of the domestic turkey by the ancient Maya thus could have resulted largely from cultural interaction and exchange, rather than as a reaction to potential protein stress due to population expansion. Conclusions and Directions for Future Research This review clearly demonstrates the need for expanded research focusing on the timing, location, and process of turkey domestication and management in Mesoamerica. Greater attention to Mesoamerica turkey husbandry in will likely overturn some of our current interpretations, and yield new insights into the history of North America s only indigenous domesticated vertebrate animal. At the most basic level, there is a critical need for more thorough and systematic analysis and reporting of faunal remains from Mesoamerican archaeological sites. Our review reveals that many relevant faunal assemblages are underanalyzed and that the reported data are seldom presented in a way that allows for temporal and spatial comparison across sites or regions. More thorough analysis and reporting of zooarchaeological remains (e.g., see Emery 2004a,b; Grigson 1978), including sample quantification according to both NISP and minimum number of individuals (MNI), division of samples by chronology, and reporting of age, sex, skeletal element, and contextual data would greatly improve our ability to reconstruct the history of turkey domestication in Mesoamerica. With the increasing availability of 3-D scanning technology, it may also be possible to scan and share digital or printed replicas of turkey bones among researchers to confirm identifications, and conduct inter-site morphological comparisons (Emery et al. 2014). There is also a need to conduct combined morphometric and genetic analyses of a temporal sequence of archaeological and historic wild and domestic turkeys from Mesoamerica. Such combined analyses would allow us to evaluate potential changes in genetic and morphological traits (e.g., changes in body proportion or overall size) associated with the domestication process. Utilizing archaeological samples avoids the effects of recent artificial selection that may make modern turkeys poor morphological and genetic proxies for indigenous, pre-colonial domestic turkeys. It would then be productive to compare the genetic and morphological traits of archaeological turkeys to modern turkey populations from rural Mexico to trace more recent changes in the species. As with many domestic species, identifying more specifically when and where the domestication process began in Mesoamerican is likely to be problematic. In Mesoamerica, the transition from foraging to farming was a gradual one characterized by a very long period of low level domesticated plant use alongside wild resources (Piperno 2011). Based on the paucity of turkey remains in early archaeological deposits, it seems likely that turkey domestication also started very gradually, and was not intensified until much later in time. This pattern makes it difficult to determine when and where turkey rearing first began, since the earliest captive turkeys will appear within the geographic range of their wild counterparts, and because such low levels of husbandry or captive rearing are unlikely to leave any genetic or morphological markers until much later in the domestication process. Moreover, wild turkey hunting

18 Thornton and Emery may have continued alongside the earliest examples of turkey rearing so sites could be expected to contain the remains of both wild, and tame or captive-reared turkeys. In the absence of other lines of evidence such as pen structures, increased species abundance, morphological changes, and the presence of juveniles, one of the most promising methods for determining the spatial and temporal origins of turkey domestication is stable isotope analysis, which can be used to reconstruct past diets and determine geographic origins. Previous isotopic research on turkeys in the American SW (Rawlings and Driver 2010; McCaffery et al. 2014) and other domesticated taxa including pigs, sheep, goats, cattle, llamas, and alpacas (e.g., Makarewicz and Tuross 2012; Minagawaet al. 2005; Noe-Nygaard et al. 2005; Thorntonet al. 2010; Huet al. 2009) confirm the utility of such methods to document the timing of animal domestication and various types of husbandry practices. Among Mesoamerican turkeys, dietary shifts associated with confinement or habitation within human settlements may be one of the earliest indicators of domestication widely detectable in the archaeological record. As such, stable carbon (δ 13 C) and nitrogen (δ 15 N) isotopes may be critical to distinguishing between wild and domestic turkeys at various stages within the domestication process. Stable carbon isotope analysis will also be useful for reconstructing husbandry practices. For example, turkeys primarily kept in pens or cages would need to be fed on a regular basis, and would thus likely show very high δ 13 C values indicative of a diet consisting primarily of maize. In contrast, free range turkeys could be expected to show more diverse diets slightly lower in human provided fodder. Regardless of the husbandry pattern, tame or captive turkeys should be isotopically distinct from wild turkeys due to their much greater access to maize (Rawlings and Driver 2010; McCaffery et al. 2014; Thornton and Emery 2014). but the degree of isotopic separation may vary according to rearing and foddering techniques. Variation in turkey stable carbon isotope values may therefore be useful for comparing spatial and temporal diversity in turkey husbandry practices within Mesoamerica, and between Mesoamerica and the American Southwest. Other isotopic ratios can be used to answer additional questions such as whether the earliest examples of turkeys found outside their natural geographic ranges represent isolated trade items or on-site, local rearing. Combined strontium ( 87 Sr/ 86 Sr) and oxygen (δ 18 O) isotope ratios in archaeological remains record the local isotopic signatures found in a region s underlying bedrock, and hydrological systems (Faure and Powell 1972; Longinelli 1984). As an animal feeds and drinks, the local strontium and oxygen values are thus recorded in its skeletal tissues. These isotopic measures have been previously used in Mesoamerica to study human migration (e.g.,white et al. 2007; Wright 2005) and animal trade (Somerville et al. 2010; Thornton 2011). When applied to archaeological turkey remains found outside the natural geographic range of wild M. gallopavo, strontium and oxygen isotope ratios can thus determine whether the nonlocal turkeys represent traded animals killed before or shortly after their arrival, or established domestic/captive populations. This information will allow archaeologists to reconstruct the diffusion of turkey husbandry throughout Mesoamerica. On a more conceptual level, researchers need to further explore the role that captive, managed, and domestic resources played in Mesoamerican subsistence, ritual, and political economies. The culture-specific motivations for animal management and rearing often cannot be explained in terms of simple, single-force causality (Zeder 2006) but greater attention to how wild and domestic turkeys were used by ancient