The 10,000-year biocultural history of fallow deer and its implications for conservation policy.

Over the last 10,000 y, humans have manipulated fallow deer populations with varying outcomes. Persian fallow deer (Dama mesopotamica) are now endangered. European fallow deer (Dama dama) are globally widespread and are simultaneously considered wild, domestic, endangered, invasive and are even the national animal of Barbuda and Antigua. Despite their close association with people, there is no consensus regarding their natural ranges or the timing and circumstances of their human-mediated translocations and extirpations. Our mitochondrial analyses of modern and archaeological specimens revealed two distinct clades of European fallow deer present in Anatolia and the Balkans. Zooarchaeological evidence suggests these regions were their sole glacial refugia. By combining biomolecular analyses with archaeological and textual evidence, we chart the declining distribution of Persian fallow deer and demonstrate that humans repeatedly translocated European fallow deer, sourced from the most geographically distant populations. Deer taken to Neolithic Chios and Rhodes derived not from nearby Anatolia, but from the Balkans. Though fallow deer were translocated throughout the Mediterranean as part of their association with the Greco-Roman goddesses Artemis and Diana, deer taken to Roman Mallorca were not locally available Dama dama, but Dama mesopotamica. Romans also initially introduced fallow deer to Northern Europe but the species became extinct and was reintroduced in the medieval period, this time from Anatolia. European colonial powers then transported deer populations across the globe. The biocultural histories of fallow deer challenge preconceptions about the divisions between wild and domestic species and provide information that should underpin modern management strategies.

Wet feet: developing sulfur isotope provenance methods to identify wetland inhabitants.

The stable isotopes of sulfur provide a distinctive signature for marine proximity and interaction. Exploring coastal proximity has been the principal application of sulfur isotopes in archaeology and palaeoecology, but this deals only with high (greater than 14‰) isotope values, meaning little interpretation has been gained from lower values. Progress has been hindered by issues with biosphere mapping. Air pollution can impact modern landscapes, significantly lowering sulfur isotope baselines, leading to the assumption that modern vegetation-based sulfur maps are not reliable. This research explores the potential of previously undiagnostic low, and often, negative sulfur isotope values for identifying wetland dwellers. Impervious clays that support wetlands are distinctive ecosystems and this study tests the hypothesis that they will produce low isotope values owing to both the underlying substrate and to redox conditions. Primary mapping of targeted areas using modern plants highlights zones with natural negative sulfur values and demonstrates that this constitutes a distinctive wetland signature. Analysis of modern and archaeological fauna demonstrates that these distinctive isotope compositions are transferred into the food chain. These findings propel the interpretative potential of sulfur isotopes forward and add to the growing knowledge to provide means for identifying archaeological humans and animals raised in wetlands.

Diversity aboard a Tudor warship: investigating the origins of the Mary Rose crew using multi-isotope analysis.

The great Tudor warship, the Mary Rose, which sank tragically in the Solent in 1545 AD, presents a rare archaeological opportunity to research individuals for whom the precise timing and nature of death are known. A long-standing question surrounds the composition of the Tudor navy and whether the crew were largely British or had more diverse origins. This study takes a multi-isotope approach, combining strontium (87Sr/86Sr), oxygen (δ18O), sulfur (δ34S), carbon (δ13C) and nitrogen (δ15N) isotope analysis of dental samples to reconstruct the childhood diet and origins of eight of the Mary Rose crew. Forensic ancestry estimation was also employed on a subsample. Provenancing isotope data tentatively suggests as many as three of the crew may have originated from warmer, more southerly climates than Britain. Five have isotope values indicative of childhoods spent in western Britain, one of which had cranial morphology suggestive of African ancestry. The general trend of relatively high δ15N and low δ13C values suggests a broadly comparable diet to contemporaneous British and European communities. This multi-isotope approach and the nature of the archaeological context has allowed the reconstruction of the biographies of eight Tudor individuals to a higher resolution than is usually possible.

Feasting and Mobility in Iron Age Ireland: Multi-isotope analysis reveals the vast catchment of Navan Fort, Ulster.

Navan Fort is an iconic prehistoric Irish ceremonial centre and the legendary capital of Ulster. The fort has produced an exceptional pig-dominated faunal assemblage that also contained a barbary macaque skull. Dating from the 4th to 1st century BC, it is likely to be a ceremonial feasting centre that may have drawn people and their animals from across Ulster and beyond. This study uses a multi-isotope (87Sr/86Sr, δ34S, δ13C, δ15N) approach to identify non-local animals and reconstruct site catchment. New biosphere mapping means that isotope data can be more confidently interpreted and the combination of strontium and sulphur analysis has the potential to estimate origins. In the absence of human remains, fauna provide the best proxy for human movement. Results for the 35 analysed animals are wide-ranging, especially in terms of strontium (0.707-0.715), which has the largest range for an Irish site. Sulphur values are more restricted (13.1‰-17.1‰) but are high in the context of British and Irish data. Results provide clear evidence for animals (and thus people) coming from across Ulster and beyond, demonstrating the site's wide catchment. Navan Fort was clearly a major ceremonial centre with far-reaching influence and hosted feasts that drew people and animals from afar.

Immune state is associated with natural dietary variation in wild mice Mus musculus domesticus.

The ability, propensity and need to mount an immune response vary both among individuals and within a single individual over time.A wide array of parameters has been found to influence immune state in carefully controlled experiments, but we understand much less about which of these parameters are important in determining immune state in wild populations.Diet can influence immune responses, for example when nutrient availability is limited. We therefore predict that natural dietary variation will play a role in modulating immune state, but this has never been tested.We measured carbon and nitrogen stable isotope ratios in an island population of house mice Mus musculus domesticus as an indication of dietary variation, and the expression of a range of immune-related genes to represent immune state.After accounting for potential confounding influences such as age, sex and helminth load, we found a significant association between carbon isotope ratio and levels of immune activity in the mesenteric lymph nodes, particularly in relation to the inflammatory response.This association demonstrates the important interplay between diet and an animal's response to immune challenges, and therefore potentially its susceptibility to disease. A plain language summary is available for this article.

The relationship between the phosphate and structural carbonate fractionation of fallow deer bioapatite in tooth enamel.

RATIONALE: The species-specific relationship between phosphate (δ18 OP values) and structural carbonate (δ18 OC values) oxygen isotope ratios has been established for several modern and fossil animal species but until now it has not been investigated in European fallow deer (Dama dama dama). This study describes the relationship between phosphate and structural carbonate bioapatite in tooth enamel of extant fallow deer, which will help us further understand the species' unique environmental and cultural history. METHODS: The oxygen isotope composition of phosphate (δ18 OP value) and structural carbonate (δ18 OC value) of hydroxylapatite was determined in 51 modern fallow deer tooth enamel samples from across Europe and West Asia. The δ18 OC values were measured on a GV IsoPrime dual-inlet mass spectrometer and the δ18 OP values on a temperature-controlled elemental analyser (TC/EA) coupled to a DeltaPlus XL isotope ratio mass spectrometer via a ConFlo III interface. RESULTS: This study establishes a direct and linear relationship between the δ18 OC and δ18 OP values from fallow deer tooth enamel (δ18 OC = +9.244(±0.216) + 0.958 * δ18 OP (±0.013)). Despite the successful regression, the variation in δ18 O values from samples collected in the same geographical area is greater than expected, although the results cluster in broad climatic groupings when Koppen-Geiger classifications are taken into account for the individuals' locations. CONCLUSIONS: This is the first comprehensive study of the relationship between ionic forms of oxygen (phosphate oxygen and structural carbonate) in fallow deer dental enamel. The new equation will allow direct comparison with other herbivore data. Variable δ18 O values within populations of fallow deer broadly reflect the ecological zones they are found in which may explain this pattern of results in other euryphagic species.

An early Cambrian greenhouse climate.

The oceans of the early Cambrian (~541 to 509 million years ago) were the setting for a marked diversification of animal life. However, sea temperatures-a key component of the early Cambrian marine environment-remain unconstrained, in part because of a substantial time gap in the stable oxygen isotope (δ18O) record before the evolution of euconodonts. We show that previously overlooked sources of fossil biogenic phosphate have the potential to fill this gap. Pristine phosphatic microfossils from the Comley Limestones, UK, yield a robust δ18O signature, suggesting sea surface temperatures of 20° to 25°C at high southern paleolatitudes (~65°S to 70°S) between ~514 and 509 million years ago. These sea temperatures are consistent with the distribution of coeval evaporite and calcrete deposits, peak continental weathering rates, and also our climate model simulations for this interval. Our results support an early Cambrian greenhouse climate comparable to those of the late Mesozoic and early Cenozoic, offering a framework for exploring the interplay between biotic and environmental controls on Cambrian animal diversification.

A multi-isotope investigation of diet and subsistence amongst island and mainland populations from early medieval western Britain.

OBJECTIVES: This is the first investigation of dietary practices amongst multiple early medieval populations (AD 500-1000) from Wales and the Isle of Man using carbon, nitrogen, and sulphur isotope analysis. The analysis will illuminate similarities or differences between the diets and subsistence strategies of populations occupying different geographical regions, specifically those living in marginal coastal regions in comparison to inland populations well-connected to ecclesiastical centres and high-status settlements. MATERIALS AND METHODS: One hundred and two human skeletons were sampled for carbon and nitrogen isotope analysis, and 69 human skeletons were sampled for sulphur isotope analysis from nine cemetery sites from western Britain (Isle of Man = 3, southwest Wales = 4, southeast Wales = 2). Thirteen faunal skeletons from St Patrick's Chapel (southwest Wales) were sampled for carbon, nitrogen, and sulphur isotope analysis. RESULTS: Human δ13 C values range from -19.4‰ to -21.2‰ (δ13 C mean=-20.4 ±0.4‰, 1σ, n = 86), and δ15 N values range from 9.1‰ to 13.8‰ (δ15 N mean = 10.8 ± 0.9‰, 1σ, n = 86). δ34 S values range from 1.2‰ to 18.4‰ (δ34 S mean = 11.6 ± 4.5‰, 1σ, n = 66). Significant differences were noted between the mean δ13 C, δ15 N and δ34 S values according to geographic region: Isle of Man (δ13 C = -20.7 ± 0.4‰, δ15 N = 11.4 ±0.6‰, n = 13/86; δ34 S mean = 17.1 ±0.6, n = 4/66), southwest Wales (δ13 C = -20.5 ± 0.4‰, δ15 N = 11.0 ±1‰, n = 32/86; δ34 S = 16.1 ± 2.1, n = 21/66), and southeast Wales (δ13 C =-20.3 ±0.4‰, δ15 N = 10.4 ±0.7‰, n = 41/86; δ34 S= 8.8 ±3‰, n = 41/66). Faunal δ13 C values range from -23.1‰ to -21.2‰ (δ13 C mean= -22.1 ±0.5‰, 1σ, n = 13), and δ15 N values range from 6.3‰ to 9.8‰ (δ15 N mean = 7.3 ± 1.1‰, 1σ, n = 13). δ34 S values range from 4.7‰ to 18.4‰ (δ34 S mean= 16.3 ± 3.6‰, 1σ, n = 13). CONCLUSIONS: The data reveal a reliance on terrestrial protein, however differences are observed between the resource consumption of populations from southwest Wales and the Isle of Man in comparison to the populations from southeast Wales. Populations from the west coast have a marine sulphur signature that reflects their coastal proximity and may also include a reliance on seaweed as a fertiliser/food source. Populations in the southeast were connected to ecclesiastical centres and high-status settlements and had access to inland-grown produce. The data add support to the suggestion that δ34 S can be used as a mobility indicator.

The oxygen isotope relationship between the phosphate and structural carbonate fractions of human bioapatite.

RATIONALE: Oxygen isotope analysis of archaeological human dental enamel is widely used as a proxy for the drinking water composition (δ(18)O(DW)) of the individual and thus can be used as an indicator of their childhood place of origin. In this paper we demonstrate the robustness of structural carbonate oxygen isotope values (δ(18)O(C)) in bioapatite to preserve the life signal of human tooth enamel by comparing it with phosphate oxygen isotope values (δ(18)O(P)) derived from the same archaeological human tooth enamel samples. METHODS: δ(18)O(C) analysis was undertaken on 51 archaeological tooth enamel samples previously analysed for δ(18)O(P) values and strontium isotopes. δ(18)O(C) values were determined on a GV IsoPrime dual inlet mass spectrometer, following a series of methodological tests to assess: (1) The reaction time needed to ensure complete release of CO(2) from structural carbonate in the enamel; (2) The effect of an early pre-treatment with dilute acetic acid to remove diagenetic carbonate; (3) Analytical error; (4) Intra-tooth variation; and (5) Diagenetic alteration. RESULTS: This study establishes a direct relationship between δ(18)O(C) and δ(18)O(P) values from human tooth enamel (δ(18)O(P) = 1.0322 × Î´(18)O(C) - 9.6849). We have combined this equation with the drinking water equation of Daux et al. (J. Hum. Evol. 2008, 55, 1138) to allow direct calculation of δ(18)O(DW) values from human bioapatite δ(18)O(C) (δ(18)O(DW) = 1.590 × Î´(18)O(C) - 48.634). CONCLUSIONS: This is the first comprehensive study of the relationship between the ionic forms of oxygen (phosphate oxygen and structural carbonate) in archaeological human dental enamel. The new equation will allow direct comparison of data produced by the different methods and allow drinking water values to be calculated from structural carbonate data with confidence.

Multi-tissue analysis of oxygen isotopes in wild rhesus macaques (Macaca mulatta).

Oxygen isotopes in animal tissues are directly related to body water composition and thus the environment. Accurate measurement of animal tissue δ(18)O provides information about local climate, an animal's geographical origin and subsequent movements, with wide applications in palaeobiology and forensic science. The genesis and evolution of tissue-based oxygen isotopes within species and within individuals are complex. We present the first data, for non-human primates, rhesus macaques (Macaca mulatta), on the relationship between oxygen isotope sources in bio-apatite (PO(4) and PCO(3)) and hair taken from six sample sites in Asia, ranging from western India to northern Vietnam. The range of values is similar within each tissue type, with good correlation between tissues (r = 0.791 to 0.908), allowing cross-tissue extrapolations. This is important when the availability of suitable tissues is limited. Biological interpretation of the small data set is difficult: macaque diets are eclectic, and the samples are from various locations. However, factors such as overall climate, precipitation quantity and source, and altitude are clearly influencing the results for each discrete geographical grouping. Future work could be aimed at assessing δ(18)O tissue associations for other species as the relationships appear to be species-specific.

Modern macaque dietary heterogeneity assessed using stable isotope analysis of hair and bone.

Dietary variability might have been a major factor in the dispersal and subsequent persistence of the genus Macaca in both tropical and temperate areas. Macaques are found from northern Africa to Japan, yet there have been few systematic attempts to compare diets between different modern populations. Here we have taken a direct approach and sampled museum-curated tissues (hair and bone) of Macaca mulatta (rhesus macaques) for carbon and nitrogen stable isotope dietary analyses. Samples from India, Vietnam, and Burma (Myanmar) were taken, representing both tropical and temperate populations. The delta(13)C values obtained from hair show that the temperate macaques, particularly those from Uttar Pradesh, have a delta(13)C signature that indicates at least some use of C(4) resources, while the tropical individuals have a C(3)-based diet. However, delta(13)C values from bone bioapatite indicate a C(3)-based diet for all specimens and they do not show the C(4) usage seen in the hair of some animals, possibly because bone represents a much longer turnover period than that of hair. The results of delta(15)N analyses grouped animals by geographic region of origin, which may be related to local soil nitrogen values. The greatest variation in delta(15)N values was seen in the specimens from Burma, which may be partly due to seasonality, as specimens were collected at different times of year. We also investigated the relationship between the hair, bone collagen, and bone bioapatite delta(13)C results, and found that they are highly correlated, and that one tissue can be used to extrapolate results for another. However, our results also suggest that hair may pick up discrete feeding traces (such as seasonal usage), which are lost when only bone collagen and bioapatite are examined. This has important implications for dietary reconstructions of archaeological and paleontological populations.