Animal Management at the Ancient Metropolis of Teotihuacan, Mexico: Stable Isotope Analysis of Leporid (Cottontail and Jackrabbit) Bone Mineral.

Human-animal interactions have played crucial roles in the development of complex societies across the globe. This study examines the human-leporid (cottontail and jackrabbit) relationship at the pre-Hispanic (AD 1-550) city of Teotihuacan in the Basin of Mexico and tests the hypothesis that leporids were managed or bred for food and secondary products within the urban core. We use stable isotope analysis (δ13Capatite and δ18Oapatite) of 134 leporid specimens from five archaeological contexts within the city and 13 modern specimens from across central Mexico to quantify aspects of leporid diet and ecology. The results demonstrate that leporids from Oztoyahualco, a residential complex associated with a unique rabbit sculpture and archaeological traces of animal butchering, exhibit the highest δ13Capatite values of the sample. These results imply greater consumption of human-cultivated foods, such as maize (Zea mays), by cottontails and jackrabbits at this complex and suggest practices of human provisioning. A lack of significant differences in δ18Oapatite values between ancient and modern leporids and between Oztoyahualco and other locations within Teotihuacan indicates generally similar relative humidity from sampled contexts. Results of this study support the notion that residents provisioned, managed, or bred leporids during the height of the city, and provide new evidence for mammalian animal husbandry in the ancient New World.

Environmental variables across Pan troglodytes study sites correspond with the carbon, but not the nitrogen, stable isotope ratios of chimpanzee hair.

Diet influences the stable isotope ratios of carbon and nitrogen (δ(13) C and δ(15) N values) in animal tissue; but here we explore the influences of particular aspects of the local environment on those values in chimpanzees (Pan troglodytes). In this article we present new δ(13) C and δ(15) N values in Gombe chimpanzees using hairs collected from night nests in 1989. Then, we explore the influence of environmental factors by comparing our Gombe data to those from eight additional Pan study sites with previously published stable isotope data. We compare chimpanzee δ(13) Chair and δ(15) Nhar values to specific characteristics of local site ecology (biome and ecoregion) and to local Mean Annual Precipitation (MAP) to test hypotheses based on known effects of these variables on the δ(13) C and δ(15) N values in plant tissues. The comparison shows that hair from chimpanzees living in savanna sites with lower MAP have higher δ(13) Chair values than do chimpanzees living in woodland and forested sites with higher MAP. These results demonstrate the potential of using δ(13) C values in primate tissue to indicate aspects of their local ecology in cases where the ecology is uncertain, such as samples collected early in the last century and in fossil hominins. In contrast to expectations, however, chimpanzee δ(15) Nhair values from some savanna sites with lower MAP are lower, not higher, than those living in more forested areas with higher MAP. It is likely that diet selectivity by chimpanzees affects δ(15) Nhair values to a greater extent than does the influence of precipitation on plants. Am. J. Primatol. 78:1055-1069, 2016. © 2015 Wiley Periodicals, Inc.

Stable Isotopes and Zooarchaeology at Teotihuacan, Mexico Reveal Earliest Evidence of Wild Carnivore Management in Mesoamerica.

From Roman gladiatorial combat to Egyptian animal mummies, the capture and manipulation of carnivores was instrumental in helping to shape social hierarchies throughout the ancient world. This paper investigates the historical inflection point when humans began to control animals not only as alimental resources but as ritual symbols and social actors in the New World. At Teotihuacan (A.D. 1-550), one of the largest pre-Hispanic cities, animal remains were integral components of ritual caches expressing state ideology and militarism during the construction of the Moon and the Sun Pyramids. The caches contain the remains of nearly 200 carnivorous animals, human sacrificial victims and other symbolic artifacts. This paper argues the presence of skeletal pathologies of infectious disease and injuries manifest on the carnivore remains show direct evidence of captivity. Stable isotope analysis (δ13C and δ15N) of bones and teeth confirms that some of these carnivores were consuming high levels of C4 foods, likely reflecting a maize-based anthropocentric food chain. These results push back the antiquity of keeping captive carnivores for ritualistic purposes nearly 1000 years before the Spanish conquistadors described Moctezuma's zoo at the Aztec capital. Mirroring these documents the results indicate a select group of carnivores at Teotihuacan may have been fed maize-eating omnivores, such as dogs and humans. Unlike historical records, the present study provides the earliest and direct archaeological evidence for this practice in Mesoamerica. It also represents the first systematic isotopic exploration of a population of archaeological eagles (n = 24) and felids (n = 29).

Diet and gender in the Tiwanaku colonies: Stable isotope analysis of human bone collagen and apatite from Moquegua, Peru.

OBJECTIVES: Gender and other facets of social identity play important roles in the organization of complex societies. This study reconstructs dietary practices within the Middle Horizon (AD 500-1000) Tiwanaku colonies in southern Peru to increase our knowledge of gendered patterns of consumption within this early expansive state. METHODS: We use stable isotope analysis of 43 human bone samples representing 14 females, 20 males, 8 juveniles, and 1 indeterminate individual recovered from burial excavations at the sites of Rio Muerto and Omo in the Moquegua Valley. Data are contextualized by comparisons with previously published Tiwanaku isotope data from the period. RESULTS: Our results find mean values of δ(13) Capatite = -7.3 ± 1.6% (N = 36, 1SD), δ(13) Ccollagen = -12.3 ± 1.5% (N = 43, 1SD), and δ(15) Ncollagen = 8.4 ± 1.6% (N = 43, 1SD). Between the sexes, Mann-Whitney U tests demonstrate significant differences in δ(13) Ccollagen (U = 74, P = 0.021), but no differences in δ(13) Capatite (U = 58, P = 0.095) or δ(15) Ncollagen (U = 116, P = 0.755) values. CONCLUSIONS: These data indicate relatively high C4 plant consumption among the Tiwanaku colonies, and support paleobotanical and archaeological evidence that maize (Zea mays) was the staple crop. Dietary values are similar overall between the sexes, but significantly higher δ(13) Ccollagen values in males is consistent with a model of gendered norms of consumption similar to that of the later Inca (AD 1438-1533), where males consumed more maize than females, often in the form of beer (chicha). Results provide new insights on social dynamics within the Tiwanaku colonies and suggest the increased importance maize consumption for males during the Tiwanaku expansion.

Paleomobility in the Tiwanaku diaspora: biogeochemical analyses at Rio Muerto, Moquegua, Peru.

Paleomobility has been a key element in the study of the expansion of ancient states and empires, including the Tiwanaku polity of the South Central Andes (AD 500-1000). We present radiogenic strontium and oxygen isotope data from human burials from three cemeteries in the Tiwanaku-affiliated Middle Horizon archaeological site complex of Rio Muerto in the Moquegua Valley of southern Peru. At Rio Muerto, archaeological human enamel and bone values range from (87) Sr/(86) Sr = 0.70657-0.72018, with a mean of (87) Sr/(86) Sr = 0.70804 ± 0.00207 (1σ, n = 55). For the subset of samples analyzed for oxygen isotope values (n = 48), the data ranges from δ(18) Ocarbonate(VSMOW) = +18.1 to +27.0‰. When contextualized with other lines of archaeological evidence, we interpret these data as evidence for an archaeological population in which the majority of individuals had "local" origins, and were likely second-generation, or more, immigrants from the Tiwanaku heartland in the altiplano. Based on detailed life history data, we argue a smaller number of individuals came at different ages from various regions within the Tiwanaku polity. We consider whether these individuals with isotopic values consistent with "nonlocal" geographic origins could represent first-generation migrants, marriage exchange partners, or occupationally mobile herders, traders or other travelers. By combining isotopic life history studies with mortuary treatment data, we use a person-centered migration history approach to state integration and expansion. Isotopic analyses of paleomobility at the Rio Muerto site complex contribute to the role of diversity in ancient states by demonstrating the range of geographic origins rather than simply colonists from the Lake Titicaca Basin.

Old stones' song: use-wear experiments and analysis of the Oldowan quartz and quartzite assemblage from Kanjera South (Kenya).

Evidence of Oldowan tools by ∼2.6 million years ago (Ma) may signal a major adaptive shift in hominin evolution. While tool-dependent butchery of large mammals was important by at least 2.0 Ma, the use of artifacts for tasks other than faunal processing has been difficult to diagnose. Here we report on use-wear analysis of ∼2.0 Ma quartz and quartzite artifacts from Kanjera South, Kenya. A use-wear framework that links processing of specific materials and tool motions to their resultant use-wear patterns was developed. A blind test was then carried out to assess and improve the efficacy of this experimental use-wear framework, which was then applied to the analysis of 62 Oldowan artifacts from Kanjera South. Use-wear on a total of 23 artifact edges was attributed to the processing of specific materials. Use-wear on seven edges (30%) was attributed to animal tissue processing, corroborating zooarchaeological evidence for butchery at the site. Use-wear on 16 edges (70%) was attributed to the processing of plant tissues, including wood, grit-covered plant tissues that we interpret as underground storage organs (USOs), and stems of grass or sedges. These results expand our knowledge of the suite of behaviours carried out in the vicinity of Kanjera South to include the processing of materials that would be 'invisible' using standard archaeological methods. Wood cutting and scraping may represent the production and/or maintenance of wooden tools. Use-wear related to USO processing extends the archaeological evidence for hominin acquisition and consumption of this resource by over 1.5 Ma. Cutting of grasses, sedges or reeds may be related to a subsistence task (e.g., grass seed harvesting, cutting out papyrus culm for consumption) and/or a non-subsistence related task (e.g., production of 'twine,' simple carrying devices, or bedding). These results highlight the adaptive significance of lithic technology for hominins at Kanjera.

Moderate to high levels of exercise are associated with higher resting energy expenditure in community-dwelling postmenopausal women.

Postmenopausal women experience an age-related decline in resting energy expenditure (REE), which is a risk factor for energy imbalance and metabolic disease. Exercise, because of its association with greater lean tissue mass and other factors, has the potential to mediate REE decline, but the relation between exercise and REE in postmenopausal women is not well characterized. This study tests the hypothesis that exercise energy expenditure (EEE) is positively associated with REE and can counter the effects of age and menopause. It involves a cross-sectional sample of 31 healthy postmenopausal women (aged 49-72 years) with habitual exercise volumes at or above levels consistent with current clinical recommendations. Subjects kept exercise diaries for 4 weeks that quantified exercise activity and were measured for body composition, maximal oxygen uptake, and REE. Multiple regression analysis was used to test for associations between EEE, age, body composition, and REE. There was a significant positive relation between EEE and lean tissue mass (fat-free mass and fat-free mass index). The relation between REE and EEE remained significant even after controlling for lean tissue mass. These results support the hypothesis that exercise is positively associated with REE and can counter the negative effects of age and menopause. They also indicate a continuous relation between exercise and REE across ranges of exercise, from moderate to high. Exercise at levels that are at or above current clinical guidelines might, in part, ameliorate the risk for energy imbalance and metabolic disease because of its positive relation with REE.

Paleoclimate during Neandertal and anatomically modern human occupation at Amud and Qafzeh, Israel: the stable isotope data.

The δ(13)C(en) and δ(18)O(en) values of goat and gazelle enamel carbonate indicate that Neandertals at Amud Cave, Israel (53-70 ka) lived under different ecological conditions than did anatomically modern humans at Qafzeh Cave, Israel (approximately 92 ka). During the Last Glacial Period, Neandertals at Amud Cave lived under wetter conditions than those in the region today. Neither faunal species ate arid-adapted C(4) plants or drought-stressed C(3) plants. The variation in gazelle δ(18)O(en) values suggests multiple birth seasons, which today occur under wetter than normal conditions. The magnitude and pattern of intra-tooth variation in goat δ(18)O(en) values indicate that rain fell throughout the year unlike today. Anatomically modern humans encountered a Qafzeh Cave region that was more open and arid than Glacial Period Amud Cave, and more open than today's Upper Galilee region. Goat δ(13)C(en) values indicate feeding on varying amounts of C(4) plants throughout the year. The climate apparently ameliorated higher in the sequence; but habitats remained more open than at Amud Cave. Both gazelles and goats fed on C(3) plants in brushy habitats without any inclusion of C(4) plants. The magnitude of intra-tooth variation in goat δ(18)O(en) values, however, suggest that some rain fell throughout the year, and the relative representation of woodland dwelling species indicates the occurrence of woodlands in the region. Climate differences affecting the distribution of plants and animals appear to be the significant factor contributing to behavioral differences previously documented between Neandertals and anatomically modern humans in the region. Climate forcing probably affected the early appearances of anatomically modern humans, although not the disappearance of Neandertals from the Levant.

Stable isotope evidence for the adoption of maize agriculture.

Over the past three decades, dozens of studies have produced carbon and nitrogen stable isotope data on early human bones and teeth from North America. Because these data record individual diets, they provide one way to test various hypotheses about the uptake and continued dependence on maize agriculture that is complementary to floral, paleopathological, and demographic approaches to the same problem. The delta13C values in the organic fraction of bone (bone collagen) plotted against those in the mineral fraction of bone (bone apatite) from several regions across North America reveal that not all human groups responded in the same way to maize agriculture. Rather there was variation between and within geographic regions. Similarly, the delta15N values in bone collagen from sites from across the southeastern United States show variation in dependence on maize and on marine foods that cannot be fully explained by geography or technical expertise. In combination, the data emphasize the need to consider social forces, both internal and external, to the group under study as well as environmental and technological constraints.

A simple carbon isotope model for reconstructing prehistoric human diet.

A compilation of experimental animal data shows that neither delta13C(collagen) nor delta13C(apatite) nor Delta13C(CO-AP) indicate diagnostic reconstructions of diet, diet energy and diet protein. In contrast, plots of delta13C(collagen) against delta13C(apatite) provide a model of three regression lines (C3, C4, and marine diet protein) where position on each line indicates the energy source (C3, C4, or mixed). Neither body size nor trophic position appears to affect these relationships. Modern free-ranging, terrestrial fauna do not fit the model perhaps because they, unlike the experimental fauna, mainly use fermentation rather than digestion during energy metabolism. Archaeological humans fall as expected based on associated floral and faunal evidence. Foraging people plot at positions expected from associated C3 fauna and plants. Those from Cahokia plot, as expected, from associated deer, nuts, and maize whereas people from nearby smaller sites plot in positions consistent with eating more fish. Agriculturists from Ontario and Grasshopper Pueblo plot consistent with dependence on fish by the former and on turkeys by the latter. In Tierra del Fuego, people from interior regions ate more terrestrial fauna, as suggested by ethnohistoric reports, than did people from the coast. In the Southwestern Cape in South Africa individuals late in the sequence have pure C3 diets whereas ones early in the sequence ate marine protein as suggested by independent archaeological evidence. People on San Nicolas Island depended on C4 plants in contrast to other islands off California's coast. This simple model provides more detailed and precise dietary information than do individual isotopic measures.

Intraspecies variation in BMR does not affect estimates of early hominin total daily energy expenditure.

We conducted a meta-analysis of 45 studies reporting basal metabolic rate (BMR) data for Homo sapiens and Pan troglodytes to determine the effects of sex, age, and latitude (a proxy for climate, in humans only). BMR was normalized for body size using fat-free mass in humans and body mass in chimpanzees. We found no effect of sex in either species and no age effect in chimpanzees. In humans, juveniles differed significantly from adults (ANCOVA: P < 0.001), and senescent adults differed significantly from adults younger than 50 years (P < 0.001). Europeans differed significantly from tropical populations (P < 0.001). On the basis of these observations, we derived new equations describing the relationship between BMR and body size, and used them to predict total daily energy expenditure (TEE) in four early hominin species. Our predictions concur with previous TEE estimates (i.e. Leonard and Robertson: Am J Phys Anthropol 102 (1997) 265-281), and support the conclusion that TEE increased greatly with H. erectus. Our results show that intraspecific variation in BMR does not affect TEE estimates for interspecific comparisons. Comparisons of more closely related groups such as humans and Neandertals, however, may benefit from consideration of this variation.

Ecological attributes recorded in stable isotope ratios of arboreal prosimian hair.

Carbon and nitrogen stable isotope ratios were measured in hair samples from two species of Galago from Gedi Ruins National Monument in eastern Kenya and from Lepilemur leucopus from Beza Mahafaly Special Reserve in southern Madagascar. Forest structure was generally similar in the two areas but average rainfall was lower in Madagascar. Species average 13C values varied with feeding height in the forest canopy and with average rainfall level as expected from reported variation in plant 13C values. G. garnettii, which feeds higher in the forest canopy, had less negative 13C values than G. zanzibaricus, which spends more time below 5 m. L. leucopus, from a drought-afflicted forest, had less negative hair 13C values than the two galago species. The values within the Lepilemur sample showed a positive linear relation with percent dependence on a CAM tree species and with xeric conditions within the species reserve. Nitrogen stable isotope ratios varied with trophic level of feeding and with time spent feeding on leguminous plants. The insectivorous galagos had significantly more positive 15N values than the folivorous L. leucopus. Within the Lepilemur sample, 15N values varied inversely with the percent of feeding time spent on leguminous plants. The range of 15N and 13C values in each of the prosimian species is larger than reported for animals fed monotonous diets and for New World monkey species. The monkey species feed as groups of individuals whereas the prosimians have solitary feeding habits. The ranges in the prosimian species apparently reflect the greater variation in diet among individual prosimians compared to individual monkeys. The isotope data reported here are equivalent, on average, to those reported for other arboreal species from similar forest habitats and with similar dietary habits. This supports the use of such data for paleoecological reconstruction of forest and woodland systems and diet reconstruction of extinct primate populations and species.