Influence of seabird guano and camelid dung fertilization on the nitrogen isotopic composition of field-grown maize (Zea mays)

Organic fertilizers have the capacity to alter the nitrogen isotopic composition of plants. Camelid dung and seabird guano are two potentially important fertilizers in the agricultural systems of western South America, particularly Peru and Chile. This paper presents isotopic data (δ¹³C and δ¹⁵N) from field grown plants (maize, Zea mays) fertilized with the following four treatments: CO (control, no fertilizer applied), AS (ammonium sulfate, a chemical fertilizer), DU (camelid dung), and SG (seabird guano). Plants were grown in experimental plots in the Virú Valley, northern Peru. Plants fertilized with the chemical fertilizer presented very similar isotopic compositions compared to the control. Conversely, the camelid dung fertilized plants were characterized by higher δ¹⁵N values compared to the control plants (by 1.8 to 4.2‰ depending on the plant part). The seabird guano fertilized plants were greatly enriched in ¹⁵N in comparison to the control plants (by 11.3 to 20.0‰). The results of this study have important implications for the reconstruction of human diet using isotopic data derived from bone collagen and related tissues, particularly in the prehispanic Andes, but also in Europe and North America during the 19th century, when Peruvian seabird guano was used extensively. Specifically, the interpretation of the relative contributions of plant and animal protein to the diet on the basis of bulk isotopic compositions of bone collagen (or similar tissues) may be confounded by camelid dung fertilization if the carbon isotopic compositions of the two sources are similar. Likewise, the interpretation of the relative contributions of maize and marine protein may be confounded by seabird guano fertilization.     

Early Neolithic diets at Baijia,Wei River valley,China: stable carbon and nitrogen isotope analysis of human and faunal remains

Stable carbon and nitrogen isotope values are presented for faunal and human bone collagen from Baijia, in the Wei River valley region of Shaanxi Province, China. The remains have a calibrated age range of ca. 5709–5389 BC, and correspond with the early Neolithic Laoguantai Period. Stable isotopic results indicate that human diets included millet and probably aquatic foods such as fish and shellfish. Bovid samples are tentatively identified as water buffalo, and have a mean δ¹³C value of −14.6‰, which reflects some millet consumption. Whether bovids were grazing on wild millet, or had diets directly influenced by humans, is not known. The single Sus sample from Baijia had a diet dominated by C3 plants and is thus unlikely to have been a domesticated animal. Overall, the stable isotope results presented here conform to the current concept that the people of the Laoguantai culture were millet farmers, who had subsistence strategies that included hunted wild foods.     

Aquatic resources in human diet in the Late Mesolithic in Northern France and Luxembourg: insights from carbon,nitrogen and sulphur isotope ratios

We investigated the contribution of freshwater resources to the diet of seven Late Mesolithic hunter-gatherers (ca. 5300–7000 BC) from Northern France and Luxembourg using stable isotope ratios. In addition to the carbon and nitrogen stable isotope ratios (δ¹³C, δ¹⁵N), we explored the potential of the sulphur isotopic ratios (δ³⁴S) to detect and quantify the proportion of protein derived from aquatic foodstuff. In only two sites, animal remains from an associated settlement were available and subsequently examined to decipher the isotopic differential between terrestrial and freshwater resources. The quantification of their relative contribution was simulated using a Bayesian mixing model. The measurements revealed a significant overlap in δ¹³C values between freshwater and terrestrial resources and a large range of δ¹⁵N values for each food category. The δ³⁴S values of the aquatic and terrestrial animals were clearly distinct at the settlement in the Seine valley, while the results on fish from Belgium demonstrated a possible overlap in δ³⁴S values between freshwater and terrestrial resources. Local freshwater ecosystem likely contributed to ca. 30–40 % of the protein in the diet of the individuals found in the Seine settlement. Out of this context, the isotopic signature and thus contribution of the available aquatic foods was difficult to assess. Another potential source of dietary protein is wild boar. Depending on the local context, collagen δ³⁴S values may contribute to better assessment of the relative contribution of freshwater and terrestrial resources.     

CHARACTERIZING THE DIET OF INDIVIDUALS AT THE NEOLITHIC CHAMBERED TOMB OF HAZLETON NORTH,GLOUCESTERSHIRE, ENGLAND,USING STABLE ISOTOPIC ANALYSIS*

Stable carbon and nitrogen isotope compositions were measured on human and faunal bones, sampled from the Neolithic chambered tomb of Hazleton North, Gloucestershire, UK. The values were used to characterize the diet of the burial community as a whole. Humans were higher in δ¹⁵N by 4.5–5.0‰ relative to animal δ¹⁵N, from which we conclude that, based on currently accepted interpretations of isotopic data, the humans consumed a diet that was very high in meat or animal products (75% by weight of protein). Comparison was also possible between cortical and cancellous femoral collagen, with the results showing no significant difference for the adult humans. The sample of human isotopic values showed little variability, in contrast to that found in the domestic and wild animals from the site (including cattle, pigs, sheep and deer). We suggest that this is due to local environmental differences, rather than to environmental change over time or physiological differences between individual animals, and that this pattern is likely to hold for many other archaeological sites when analysed with sufficient statistical weight.     

C4‐Plant Foraging in Northern Italy: Stable Isotopes,Sr/Ca and Ba/Ca Data of Human Osteological Samples from Roccapelago (16th–18th Centuries AD)

Human osteological samples (n = 23) taken from different anatomical parts of 11 individuals from the early modern (16th–18th century AD) site of Roccapelago (Modena, Italy) were systematically analysed for δ¹³C, δ¹⁵N and trace elements to investigate their diet. δ¹³C and δ¹⁵N correlate and show a high variability between individuals, attesting to the dietary contribution of C₄ plants. This is supported by pollen analysis of the burial site samples, which revealed the presence of maize. δ¹⁵N correlates with Sr/Ca, suggesting that the main protein source could have been milk and dairy. We therefore interpret the strong correlation between δ¹³C and δ¹⁵N as evidence for C₄‐plant foraging practice and the exploitation of livestock for meat and milk, combined with possible direct intake of C₄ plants. The Roccapelago site represents an important case study to track the evolution of the post‐medieval diet and the introduction of maize cultivation in southern Europe, as also attested by historical sources.     

Stable isotopes and human diet in central western Argentina

Central western Argentina is the southern boundary of prehispanic maize in the archaeological record. This paper explores the stable isotope information from human bone (δ¹³C, from collagen and δ¹³C from apatite) and tooth (δ¹³C from enamel) samples in order to characterize the temporal and spatial trends of these isotopes and to understand them in terms of human diet. We analyzed 104 human individuals from the last 6000 years. Using the Kellner and Schoeninger (2007) model the results shows a high variation in human diet with few human individuals with a notably high significance of C₄ plants (probably maize) as an energy source and these samples have a chronology around ca. 1000 years BP. On the other hand a high variation in an individual’s dietary life history is shown by differences between δ¹³C collagen, apatite and enamel.     

Age‐related variation in isotopic indicators of diet at medieval Kulubnarti,Sudanese Nubia

This study compares trends in dietary composition in two large cemetery populations from the site of Kulubnarti (AD 550–800) in Sudanese Nubia. Bone collagen and bone apatite carbonate were analysed to characterise stable carbon, nitrogen and oxygen isotopes. Previous research on these cemeteries has suggested marked differences in nutritional status and health between the populations. Contrary to expectations, there were no significant relationships between any isotopic indicators related to sex or cemetery of burial, suggesting no isotopically‐measurable differences in diet. However, collagen δ¹³C and δ¹⁵N were significantly related to age, suggesting age‐related differences in protein intake or other factors. Weaning trends are gradual and variable, with the range in δ¹⁵N values exceeding 4‰ among infants/young children (0–3 yrs) and standard deviations exceeding 1‰ in collagen δ¹³C and δ¹⁵N for both infants/young children and subadults (4–17 yrs). This suggests varied weaning strategies among both populations and variable diets prior to adulthood. Also observed was a distinct range of isotopic carbon and nitrogen values among individuals classified as subadults (4–17 yrs), who are depleted in collagen δ¹³C and δ¹⁵N relative to adults. However, both infants/young children and subadults are slightly enriched in δ¹⁸O relative to adults, which suggests the presence of non‐local individuals or age‐related variation in water sources. While most isotopic studies of age‐related dietary trends have focused on reconstructing the weaning process, this study presents findings that indicate tripartite isotopic trends distinguishing infancy, subadulthood and adulthood as separate dietary categories. Broad similarities are evident between the results presented here and those from several earlier studies of smaller populations and to nutritional studies of modern communities. These findings suggest that further research into health disparities at Kulubnarti should focus on non‐dietary causal factors, and more generally, that greater attention should be paid to subadulthood in palaeodiet studies. Copyright © 2006 John Wiley & Sons, Ltd.     

Fisherfolk and farmers: Carbon and nitrogen isotope evidence from Middle Horizon Ancón,Peru

This study investigates human dietary patterns and economic trends at the coastal site of Ancón, Peru during the Andean Middle Horizon (550AD –1000AD ) using stable isotopic data from 32 individuals buried at the site. δ¹³C and δ¹⁵N results from human bone collagen and δ¹³C from human tooth enamel and bone carbonate indicate that inhabitants consumed a mixed diet composed primarily of marine protein and C₄ resources, with only marginal reliance on C₃ foods. Over time, Ancóneros appear to have relied more heavily on C₄ resources, particularly maize, despite the fact that the crop could not have been grown locally. These results are notable given that C₃ rather than C₄ or marine foods dominate the site's archaeological record. These data suggest that Ancón's inhabitants either had access to more fertile land up‐valley where maize could be cultivated successfully or that they engaged in trade relationships with their valley neighbours. A third possibility is that increased maize consumption at Ancón during the Middle Horizon resulted from Wari imperial influence and interregional exchange. Comparisons of δ¹³C values in enamel and bone carbonate from Ancón individuals indicate that δ¹³C_{carb_enamel} values are significantly more positive than δ¹³C_{carb_bone} values. This suggests that the diets of young children were systematically enriched in ¹³C compared to that of adults, perhaps as a result of nursing activity and/or differential dietary practices among various age groups at the site. Copyright © 2009 John Wiley & Sons, Ltd.     

Horticulturists and oxygen ecozones in the tropical and subtropical forests of Southeast South America

We analyse the isotopic values (δ¹³C, δ¹⁵N) of the diet of pre-Columbian horticulturalist populations from tropical and subtropical areas of southeastern South America, belonging to the Guarani and Taquara archaeological units. The data indicate different trends in each one (T?=?4.21; P?=?0.0004), showing a mixed diet with maize consumption in the Guarani samples (δ¹³C_co?=??15.5?±?1.8‰; δ¹³C_ap ?10.4?±?0.8‰) and a depleted one in the Taquara ones (δ¹³C_co ?18.2?±?1.7‰; δ¹³C_ap ?11.9?±?0.9‰), with a significant internal dispersion in both populations. The first population has higher nitrogen values (δ¹⁵N 11.1?±?0.6‰) compared to the Taquara samples (δ¹⁵N 9.3?±?1‰), suggesting a more carnivorous diet. The recognition of these pre-Columbian mixed diets involves the identification of maize cultivation on the Atlantic side of the southernmost area of South America (Parana Delta, 34° SL). Through the analysis of δ¹⁸O we have identified two isotopic ecozones, the first along the Paraná River Valley, with an average value of δ¹⁸O ?3.7?±?0.5‰ (CV?=?13.5%; CI?=??3.83 / ?3.16), and the second one, located in the Planalto of southern Brazil (Araucaria Forest), with a mean value of δ¹⁸O ?1.5?±?0.3‰ (CV?=?16.5%; CI?=??1.69 / ?1.29). The isotopic data (δ¹³C, δ¹⁵N and δ¹⁸O) suggest human movements between these two ecozones.     

Isotopic Comparison of Hair,Nail and Bone: Modern Analyses

This paper presents a comprison of the isotopic values of eight pairings of hair keratin and bone collagen and 12 pairings of hair keratin and nail keratin taken from living humans resident in the U.K., with the aim of examining whether modern human isotopic data can be directly compared to archaeological isotopic data.Results showed that bone collagen was enriched relative to hair keratin from the same individual by +1·4‰ in δ¹³C and +0·86‰ in δ¹⁵N, with some small degree of variability. Isotopic comparison of hair keratin and nail keratin from the same individual showed that there is no significant difference between hair and nail keratin in δ¹³C, but that nail keratin is on average +0·65‰ more enriched in δ¹⁵N than hair.Differences in amino acid composition between hair keratin and bone collagen may account for the carbon isotopic differences between the two proteins, and there is no significant overall carbon isotopic difference between hair and nail. However there are significant isotopic differences for nitrogen in the two pairings, that differences in amino acid composition and turnover times cannot explain. We suggest that these results indicate that constancy of isotopic values between tissues, even for similar proteins, cannot be taken for granted.     

Carbon and nitrogen isotope composition of natural pastures in the dry Puna of Argentina: a baseline for the study of prehistoric herd management strategies

This study represents an isotopic survey of modern plants developed to establish baseline isotopic values in order to explore prehistoric herd management strategies employed by the South American camelid herders that occupied the southern Andean highlands during the past 3000 years. We present carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopic compositions of natural pastures collected from different plant communities along an altitudinal gradient in the dry Puna of Argentina. Our results show that C₃ plants are the most abundant along the whole altitudinal gradient and that C₃ and C₄ plants exhibit a differential distribution, the latter being less abundant in those sites located above 3900?meters above sea level (masl). At the same time, plants growing at low-altitude sites with low water availability exhibit higher δ¹⁵N values than plants growing at high-altitude sites with higher water availability. These results explain the negative correlation found between altitude and South American camelid bone collagen δ¹³C and δ¹⁵N values published in previous studies. This work represents a fundamental step towards the building of an isotopic ecology for the dry Puna area with the ultimate goal to explore herd management strategies employed by human groups in the past. In this sense, modern plant and South American camelid tissue isotopic compositions would provide a frame of reference to interpret isotopic compositions measured on archaeofaunal remains recovered at pastoral sites, with the aim to explore mobility and pastureland use by prehistoric herders.     

The stable and radio-isotope chemistry of southeastern Utah Basketmaker II burials: dietary analysis using the linear mixing model SISUS,age and sex patterning,geolocation and temporal patterning

The Bayesian linear mixing model SISUS is used to reconstruct reliance on maize and intake of animal protein from the stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope values of 149 directly dated Basketmaker II burials. Maize is the primary dietary staple. δ¹³C values are also enriched by intake of yucca and wild C₄ plant foods but these resources are of minor importance in sampled diets. Animal protein intake is low and cottontail rabbits make the greatest contribution to female diets. Adolescent diets are indistinguishable from the diets of adult females and both groups are less reliant on artiodactyls than males. The δ¹³C and δ¹⁸O values of bone hydroxyapatite distinguish nursing infants and oxygen isotope values identify outsider adults relative to those local to the region where their deaths occurred. Most Utah Basketmaker II burials are contemporary with Black Mesa Lolomai phase occupations, postdating the Marsh Pass, AZ, White Dog phase. All burials in both studies were recovered from sheltered rather than open sites and reliance on maize does not increase over the temporal span of the Utah study, covering the two sigma range 415 BC–AD 322.     

Carbon Isotope Variability in the Bone Collagen of Red Kangaroos (Macropus rufus) is Age Dependent: Implications for Palaeodietary Studies

Isotopic assessment of bone collagen is often used as an environmental tracer in both contemporary and palaeoenvironmental studies. However, variability in the isotopic composition of this tissue remains poorly understood for naturally occurring and wild populations of animals. In this study the stable carbon isotope composition of both diet and bone collagen was assessed for a population of red kangaroos (Macropus rufus). Animals sampled ranged in age from approximately 10 months to 15 years. The diet of this population, estimated from faeces collected in the field, varied from predominantly C₄grasses in late summer (δ¹³C⋍−16·5‰) to mostly C₃herbage in late winter (δ¹³C⋍−22·5‰), with a long-term average δ¹³C of between −19 and −20‰. Bone collagen was enriched in¹³C by 3 to 4‰ in older animals relative to pouch young. Isotopic analysis of hair, used to assess more recent diet in individuals, indicated that diet selection was similar in all animals that had been weaned. We suggest that the most likely explanation for the age-dependent relationship in the δ¹³C of bone collagen occurs because milk (the only source of nutrition in suckling kangaroos) is not fractionated in the same manner as plant-derived carbon during its assimilation into skeletal tissue. If this is the case, then such a relationship should be most predominant in mammals that have low birth weights (relative to the adult mother) and gain significant weight from milk. Whatever the precise mechanism(s) for the observed fractionation, bone collagen of kangaroos seems to retain an isotopic memory of the carbon laid down prior to weaning, which takes several years to be diluted and replaced with carbon derived from an “adult” herbaceous diet. These results have implications for palaeoecological research where fossil skeletal tissue is used as dietary or environmental tracers particularly if the relative age of the animals sampled is unknown.     

Stable isotopic analysis of human bones from Jiahu site,Henan, China: implications for the transition to agriculture

We have investigated change in subsistence during the transition to agriculture in the site of Jiahu, Henan Province, China, using stable isotopic analysis of collagen and apatite in human bones. Millet agriculture is well documented at drier high latitudes of the Yellow River Valley, while rice agriculture predominated at wetter lower latitudes of the Yangtze Valley region. The early Neolithic site of Jiahu lies near the boundary between the drier north and wetter south. Archaeobotanical evidence shows that rice was a significant component of diet at Jiahu, but its δ¹³C value is similar to that of other foods, and therefore cannot be conclusively identified by carbon isotope analysis. Foxtail and broomcorn millets are the only C₄ crops known for the Chinese Neolithic. Because of their high δ¹³C values, their consumption can be evaluated by stable carbon isotope analysis of human bone. Collagen reflects mainly the δ¹³C value of dietary protein, and apatite accurately records that of the whole diet. Isotopic analysis of 15 well-preserved samples from three periods shows that collagen δ¹³C values were very low for almost all individuals, suggesting C₃-based foods dominated their diets. However, apatite carbonate δ¹³C values and δ¹³C spacing between collagen and apatite (Δ¹³C_ap-co) indicate that millet may have been a minor component of the diet in this region. Individuals, who consumed the smallest amounts of animal protein, as indicated by low δ¹⁵N, generally had the highest apatite δ¹³C values. Archaeobotanical evidence for millet at Jiahu is needed to support this interpretation.     

Isotopic palaeodiet studies of Ancient Egyptian fauna and humans

We report here stable carbon and nitrogen isotope results from human and faunal bone collagen from samples taken from sites in the Egyptian Nile Valley and surrounding region, dating from the Predynastic (c. 5500 BC) through to the Dynastic (c. 343 BC) periods. Isotopic values for the human population cluster together across this broad time range, with high δ¹⁵N ratios, and δ¹³C values indicating a largely C₃ based diet. The human data is not easily explained through comparison with our associated faunal data, and so may be explained by the consumption of protein from an ecosystem we did not adequately sample, such as freshwater fish or plants and fauna with unusually high δ¹⁵N values due to the extreme aridity of this region. The faunal isotopic data we did produce shows a great range in values between and within species, especially in cattle, and reflects the close proximity of three ecozones across a relatively narrow geographical area; the river, the immediate flood-plain area around the Nile and the desert surroundings.     

Stable isotope analysis of dog,fox, and human diets at a Late Holocene Chumash village (CA-SRI-2) on Santa Rosa Island,California

Stable carbon (δ¹³C) and nitrogen (δ¹⁵N) isotope analyses of dog (Canis familiaris), island fox (Urocyon littoralis), and human bone collagen from CA-SRI-2 (AD 130–1830) on Santa Rosa Island, California provide a proxy of diet and the relationships between humans and these animals. Carbon isotopic signatures indicate that Native Americans and their dogs at CA-SRI-2 subsisted almost exclusively on marine resources, while the island fox ate primarily terrestrial foods. Nitrogen isotopes and archaeofaunal remains indicate that humans and dogs also ate higher trophic level foods, including finfishes, marine mammals, and seabirds with smaller amounts of shellfish. The CA-SRI-2 island foxes appear to have eaten higher amounts of terrestrial foods, similar to the diets observed in modern fox populations. These data generally confirm the commensal relationship assumed to exist between domesticated dogs and people, but the carbon isotopic composition of dogs is enriched ∼2‰ compared to humans. We hypothesize that the difference in carbon isotopes between dogs and humans may have resulted from a higher consumption of C3 plants with lower δ¹³C values by humans, or less likely from the ingestion by dogs of significant amounts of bone collagen, which is enriched by ∼4‰ over associated muscle.     

Stable isotope investigations of charred barley (Hordeum vulgare) and wheat (Triticum spelta) grains from Danebury Hillfort: implications for palaeodietary reconstructions

Palaeodietary studies typically focus on the analysis of bone collagen due to the limited availability of plant remains. Isotopic analysis of plant remains, however, allow for a more extensive consideration of the contribution of plants to the human diet and can potentially provide information about the environment in which the crops were grown. This paper reports the results of carbon and nitrogen isotope analyses performed on charred barley and wheat grains recovered from pits within Danebury Iron Age hillfort. To the best of our knowledge, this is the first Iron Age site in Britain from which charred grains have been isotopically analysed. Our results suggest that cereals found at the hillfort were grown in several different environmental contexts. The isotope data demonstrate that the herbivores were not consuming a diet primarily based on grains as the δ¹⁵N values of the grains are very similar to those of the herbivores. Palaeodietary investigations typically assume that humans eating plant protein only would have the same δ¹⁵N value as the local herbivores. This assumption is clearly invalid at Danebury, where the humans and animals appear to have consumed either different parts of the same plants or different plants. Researchers typically interpret high differences between human and animal δ¹⁵N values as indicative of diets high in animal protein, however where major plant resources have δ¹⁵N values similar to those of the herbivores our ability to distinguish between plant and animal sources of protein in the diet is limited. Our research has demonstrated that whenever possible it is desirable to measure the isotopic signatures of potential major plant resources in order to understand past subsistence strategies.     

Foddering by Mongolian pastoralists is recorded in the stable carbon (δC) and nitrogen (δN) isotopes of caprine dentinal collagen

Direct identification of fodder provisioning in individual animals is a previously unexplored area that has important implications for determining animal husbandry strategies in prehistory. In this study, foddering practices are examined in modern domestic and wild caprines from Mongolia (Baga Gazar'in Chuluu) through carbon and nitrogen analyses of bulk sampled dentin collagen in molar teeth. Foddered caprines display significantly different δ¹³C values compared to non-provisioned caprines. This pattern reflects the consumption of C₄ plants year-round by provisioned caprines while non-provisioned caprines ingest C₄ plants only during the summer months. Provisioned caprines also display significantly different nitrogen isotopic values relative to non-provisioned animals. The difference in nitrogen isotopes between wild and foddered animals may be due to increased access in quality graze through foddering by domestic caprines, which results in improved nutrition and extended lactation periods, or enriched δ¹⁵N of fodder plants relative to background flora. Differences in tooth wear between sampled wild and foddered animals may also contribute to the observed δ¹⁵N differences.     

Reconstructing Diet of the Early Qin (ca. 700–400 BC) at Xishan,Gansu Province,China

Here we report the bone collagen carbon and nitrogen isotopic results of humans (n = 33) and animals (n = 58) to reconstruct the dietary practices of an early Qin population dating to the Zhou Dynasty (Late Western–Early Eastern period ca. 700–400 BC ) at the Xishan site in Gansu Province, China. The humans have a very large range of δ¹³C (−23.3‰ to −7.1‰) and δ¹⁵N (4.3‰ to 10.9‰) values which reflects extraordinarily diverse diets and included individuals with predominately C₃ as well as those with exclusive C₄ diets. This wide span of isotopic results produced a subtle linear trend (R² = 0.62) in the human data, which paralleled the animals across the C₃ and C₄ environmental gradient. However, the majority of the individuals had a predominately C₄ diet based on millet with δ¹⁵N results only slightly elevated above the animals, except for the pigs and cattle. This is evidence that many of the animals were likely used for their secondary products, labour or as sacrificial offerings and that pork and beef were the main sources of animal protein for the population. High status individuals had elevated δ¹⁵N values (10.2 ± 0.6‰) compared to medium (8.9 ± 0.3‰) and lower status (8.8 ± 0.8‰) individuals, possibly related to increased animal protein in the diet. Differences related to gender were also found with females having elevated δ¹³C (−11.2 ± 1.9‰) and δ¹⁵N (9.4 ± 0.8‰) values compared to the males (δ¹³C = −14.1 ± 4.2‰; δ¹⁵N = 7.9 ± 1.9‰), but these results necessitate caution given the large number of individuals that could not be sexed. The results of this study support the view that the early Qin people were a more sedentary society focused on millet agriculture and animal husbandry, and that they were influenced by the pre‐existing populations of the central Gansu region. Copyright © 2015 John Wiley & Sons, Ltd.     

Patterns of Resource Use and Isotopic Niche Overlap Among Guanaco (Lama guanicoe), Pampas Deer (Ozotoceros bezoarticus) and Marsh Deer (Blastocerus dichotomus) in the Pampas. Ecological,Paleoenvironmental and Archaeological Implications

ABSTRACT

In this paper, we analyze the isotopic niche and resource use of the guanaco (Lama guanicoe), pampas deer (Ozotoceros bezoarticus) and marsh deer (Blastocerus dichotomus) on the temperate plain of the northern Pampa region in the southeast of South America. The measured stable isotope compositions in bone tissue are δ¹³C, δ¹⁵N and δ¹⁸O. All the analyzed samples were recovered from different archaeological sites from the late Holocene. According to the results, the guanaco was strictly confined to the Pampa plain environment, developing a broad niche based on C₃ plants and a variable contribution of C₄ grasses within a grazing trend. The pampas deer preferentially used the Pampa plain, but also the prairies on the borders of the wetland, showing eurioic characteristics and a narrower niche based on C₃ plants. In turn, the marsh deer was strictly confined to wetland environments, developing a C₃ diet–base, within a narrow isotopic niche and stenoic characteristics. The three mammals showed a wide range of intraspecific variability, which was a key factor in their adaptability to spatial and temporal changes in the vegetation coverage. In fact, the temporal trends of their isotopic values were concurrent with the major climatic variations of the Holocene. Differences in the correlations between the values of both carbon sources in both deer species compared with the guanaco suggest a distinct chemical composition of their diet and/or differences in the allocation of nutrients. The isotopic values of nitrogen and spacing of the carbon sources in guanaco (pseudoruminant) and both deer species (ruminants) show no significant differences between them, thus establishing the values for local large herbivores. Significant correlations between δ¹³C and δ¹⁵N were found in the guanaco and marsh deer. The regional and extra–regional variability in the guanaco's δ¹³C_collagen and δ¹⁵N probably reflect the clinal variations in the vegetation coverage and the amount of rainfall. The collagen isotope values in the guanaco throughout the entire Holocene show that the humid Pampa would have shifted between being a more recurrent mesic and temperate plain with minor phases of dry–mesic conditions like during the Little Ice Age, and a humid and warm one at the peaks of the Holocene Thermal Maximum and the Medieval Climatic Anomaly. The guanaco and the pampas deer adapted to all the climatic changes that happened on the humid Pampa until the biological invasions of large European mammals changed the herbivores’ guild composition of this vast plain, pushing them into peripheral habitats due to competition. The within–species variability in isotopic signals through time and space make it necessary to carry out adequate sampling before reconstructing the diets of local past populations.