Diet and social status on Taumako,a Polynesian outlier in the Southeastern Solomon Islands

Stable isotopes (δ¹³C, δ¹⁵N, and δ³⁴S) are used to characterize the diet of the adult individuals (n = 99) interred in the Namu burial ground located on the Polynesian outlier of Taumako (～300–750 BP). Polynesian outliers are islands on the fringe of Remote Oceania that were inhabited by a back migration of populations from Polynesia during prehistory. As a result of admixture with nearby island communities, little is known about the social structure and social diversity of the prehistoric inhabitants of Taumako. The distribution of prestige grave goods within the Namu cemetery has been used as evidence to support the premise that Taumakoan social structure was stratified like Polynesian societies. Here we test the hypothesis that “wealthy” individuals and males will display isotopic ratios indicative of the consumption of “high status” foods in the Pacific islands such as pork, chicken, sea turtle, and pelagic fish. The isotope results suggest the δ³⁴S values were diagenetically altered, possibly an effect of volcanism. The carbon and nitrogen stable isotope ratios indicate that the diet of all the individuals included a mixture of C₃ terrestrial plant foods (likely starchy staples such as yam, taro, and breadfruit, in addition to nuts) and a variety of marine resources, including reef and pelagic fish. The stable isotope results indicate that wealthy individuals and males were eating more foods from higher trophic levels, interpreted as being high status animal foods. The socially differentiated food consumption patterns are discussed within a Pacific island context. Am J Phys Anthropol 151:589–603, 2013. © 2013 Wiley Periodicals, Inc.     

Differences in δ15N and δ13C between embryonic and maternal tissues of the ovoviviparous bluntnose sixgill shark Hexanchus griseus

Stable nitrogen (δ¹⁵N) and carbon (δ¹³C) isotope ratios from muscle, liver and yolk were analysed from the mother and embryos of an ovoviviparous shark, Hexanchus griseus. Embryonic liver and muscle had similar δ¹⁵N and δ¹³C ratios or were depleted in heavy isotopes, compared to the same maternal somatic and reproductive yolk tissues, but no relationship existed between δ¹⁵N or δ¹³C and embryo length, as expected, because a switch to placental nourishment is lacking in this species. This study expands the understanding of maternal nourishment and embryonic stable isotope differences in ovoviviparous sharks.     

A skew in poo: Biases in primate fecal isotope analysis and recommendations for standardized sample preparation

Feces are a treasure trove in the study of animal behavior and ecology. Stable carbon and nitrogen isotope analysis allows to assess the dietary niches of elusive primate species and primate breastfeeding behavior. However, some fecal isotope data may unwillingly be biased toward the isotope ratios of undigested plant matter, requiring more consistent sample preparation protocols. We assess the impact of this potential data skew in 114 fecal samples of wild bonobos (Pan paniscus) by measuring the isotope differences (Δ¹³C, Δ¹⁵N) between bulk fecal samples containing larger particles (>1 mm) and filtered samples containing only small particles (<1 mm). We assess the influence of fecal carbon and nitrogen content (ΔC:N) and sample donor age (subadult, adult) on the resulting Δ¹³C, Δ¹⁵N values (n = 228). Additionally, we measure the isotope ratios in three systematically sieved fecal samples of chimpanzees (Pan troglodytes verus), with particle sizes ranging from 20 μm to 8 mm (n = 30). We found differences in fecal carbon and nitrogen content, with the smaller fecal fraction containing more nitrogen on average. While the Δ¹³C values were small and not affected by age or ΔC:N, the Δ¹⁵N values were significantly influenced by fecal ΔC:N, possibly resulting from the differing proportions of undigested plant macroparticles. Significant relationships between carbon stable isotope ratios (δ¹³C) values and %C in large fecal fractions of both age groups corroborated this assessment. Δ¹⁵N values were significantly larger in adults than subadults, which should be of concern in isotope studies comparing adult females with infants to assess breastfeeding. We found a random variation of up to 3.0‰ in δ¹³C and 2.0‰ in nitrogen stable isotope ratios within the chimpanzee fecal samples separated by particle sizes. We show that particle size influences isotope ratios and propose a simple, cost-effective filtration method for primate feces to exclude larger undigested food particles from the analysis, which can easily be adopted by labs worldwide.     

Stable nitrogen isotope analysis of dentine serial sections elucidate sex differences in weaning patterns of wild chimpanzees (Pan troglodytes)

Offspring provisioning is one of the most energetically demanding aspects of reproduction for female mammals. Variation in lactation length and weaning strategies between chimpanzees (Pan troglodytes), our closest living relative, and modern human societies have been reported. When and why these changes occurred is frequently debated. Our study used stable nitrogen isotope data of tooth root dentine from wild Western chimpanzees (Pan troglodytes verus) in Taï National Park, Côte d'Ivoire, to quantify weaning in these chimpanzees and explore if infant sex plays a role in maternal investment. We analyzed serial sections of deciduous lateral incisor root dentine from four Taï chimpanzees to establish the δ¹⁵N signal of nursing infants; we then analyzed serial sections of first permanent mandibular molar root dentine from 12 Taï chimpanzees to provide quantitative δ¹⁵N data on weaning in this population. Up to 2 years of age both sexes exhibited dentine δ¹⁵N values ≈2–3‰ higher than adult female Taï chimpanzees, consistent with a nursing signal. Thereafter a steady decrease in δ¹⁵N values consistent with the onset, and progression, of weaning, was visible. Sex differences were also evident, where male δ¹⁵N values decreased at a significantly slower rate compared to females. Confirmation of sex differences in maternal investment among Taï chimpanzees, demonstrates the viability of using isotope analysis to investigate weaning in non‐human primates. Additionally, assuming that behaviors observed in the Taï chimpanzees are illustrative of the ancestral pattern, our results provide a platform to enable the trajectory of weaning in human evolution to be further explored. Am J Phys Anthropol 153:635–642, 2014. © 2013 Wiley Periodicals, Inc.     

Within-lake variability in carbon and nitrogen stable isotope signatures

1. We assessed spatial and temporal variation in carbon and nitrogen isotopic signatures in different compartments of a single lake ecosystem. Stable isotope analyses were made on samples of particulate organic matter (POM), zooplankton, periphyton, macrophytes, macroinvertebrates and fish collected from several locations throughout the ice‐free period. 2. No spatial variation in δ¹³C or δ¹⁵N values was found for pelagic samples of POM and zooplankton. However, pelagic δ¹⁵N signatures increased steadily through the summer resulting in an almost 6‰ average increase in POM and zooplankton. A concurrent decrease in epilimnetic nitrate concentrations suggested that the increase in δ¹⁵N of POM and zooplankton could have resulted from a progressive ¹⁵N‐enrichment of the available inorganic nitrogen pool as the size of this pool was reduced. 3. Significant spatial variation in isotopic ratios was observed within littoral and profundal communities. Some spatial differences were likely related to lake‐specific characteristics, such as a major inlet and a small harbour area and some were interconnected with temporal events. 4. Marked differences between spring and autumn δ¹⁵N and δ¹³C values of fish at one site probably reflected a spring spawning immigration from a larger downstream lake and also indicated limited dispersal of these immigrants. 5. Our results indicate that restricted sampling of ecosystem components from lakes may provide misleading single values for the isotope end members needed for quantitative uses of stable isotopes in mixing models and for estimating trophic position. Hence we strongly advise that studies of individual lakes, or multiple lake comparisons, that utilise stable isotope analyses should pay more attention to potential within lake spatial and temporal variability of isotope ratios.     

Temporal variability in stable isotope ratios of C and N in the vibrissa of captive and wild adult South American sea lions Otaria byronia: More than just diet shifts

We analyzed the δ¹³C and δ¹⁵N values in the vibrissae of captive adult breeding South American sea lions (Otaria byronia) fed at a constant diet and then used this information to analyze the change in stable isotope values along the vibrissae from wild individuals. The overall diet‐to‐vibrissa discrimination factor of the captive animals was 3.0‰ ± 0.1‰ for δ¹³C and 3.6‰ ± 0.1‰ for δ¹⁵N, but the stable isotope ratios fluctuated periodically despite constant diet. The δ¹³C and δ¹⁵N values of the captive male declined at the end of the breeding season, whereas the δ¹³C values of the female increased during the central part of pregnancy and the δ¹⁵N values peaked during lactation. The δ¹³C and δ¹⁵N values of adult wild specimens also fluctuated periodically and vibrissae growth rate (0.15 mm/d in both sexes) was slightly lower than in captivity (0.17 mm/d), assuming an annual periodicity for oscillations. Similarities in the amplitude of the cycles of captive and wild males suggested that fasting was probably the main source of periodic variability in the δ¹⁵N of wild males, whereas pregnancy and lactation were probably the main source of periodic variability for the δ¹³C of wild females.     

Infant feeding and weaning practices in Roman Egypt.

Current knowledge of infant feeding and weaning practices during the Roman period in Egypt is limited to scanty documentary and iconographic evidence. Stable nitrogen and carbon isotope analysis provides another avenue to explore this question. A sample of 49 infant and juvenile human skeletal remains from the Kellis 2 cemetery in the Dakhleh Oasis, Egypt, was used to determine patterns of infant feeding and weaning. delta(15)N values indicate that supplementary foods were introduced at around 6 months of age, and that weaning was complete by 3 years of age. By 6 months of age, delta(13)C values become increasingly enriched over adult values, and reach peak enrichment at approximately 1.5 years of age. Beyond this age, delta(13)C gradually declines to approach adult values. This enrichment in infant delta(13)C values is indicative of consumption of (13)C-enriched supplementary foods. Based on isotopic study of faunal and botanical remains from the ancient village of Kellis, we conclude that at approximately 6 months of age, infants were fed milk of goat and/or cow.     

Stable nitrogen and carbon isotope ratios in wild native honeybees: the influence of land use and climate

The eastern hive bee Apis cerana is a major honeybee species in Asia providing numerous ecosystem services. Understanding how much the honeybees depend on natural and human-influenced plants and landscapes in different climates is important could contribute to evaluate how wild honeybees use food resources and to measure the ecosystem services. We investigated the effects of land use and climate changes on stable nitrogen and carbon isotope ratios in wild populations of A. cerana. In populations from 139 individual sites throughout Japan, we measured nitrogen (δ¹⁵N) and carbon (δ¹³C) stable isotope ratios and analyzed the effects of land use and climate. Our results showed that forested areas and annual precipitation had significant effects on δ¹⁵N, and that paddy fields and urban areas had significant effects on δ¹³C. These results suggest that A. cerana sensibly uses available food resources in the various environments and that stable nitrogen and carbon isotope ratios clearly reflect the effects of land use and climate changes on the populations of A. cerana. Thus, stable nitrogen and carbon isotope ratios in A. cerana, which widely distributes in Asia, can be used as indicators of the environments, such as land use and climate, of an area within its foraging range.     

Intergroup variation in stable isotope ratios reflects anthropogenic impact on the Barbary macaques (Macaca sylvanus) of Gibraltar

Interactions with humans impact many aspects of behavior and ecology in nonhuman primates. Because of the complexities of the human–nonhuman primate interface, methods are needed to quantify the effects of anthropogenic interactions, including their intensity and differential impacts between nonhuman primate groups. Stable isotopes can be used to quickly and economically assess intergroup dietary variation, and provide a framework for the development of specific hypotheses about anthropogenic impact. This study uses stable carbon and nitrogen isotope analysis to examine intraspecific variation in diet between five groups of Barbary macaques, Macaca sylvanus, in the Upper Rock Nature Reserve, Gibraltar. Analysis of hair from 135 macaques showed significant differences in δ¹³C and δ¹⁵N values between a group with minimal tourist contact and groups that were main tourist attractions. Because we observed no overt physiological or substantial behavioral differences between the groups, feeding ecology is the most likely cause of any differences in stable isotope ratios. Haphazard provisioning by tourists and Gibraltarians is a likely source of dietary variation between groups. Stable isotope analysis and observational data facilitate a deeper understanding of the feeding ecology of the Barbary macaques relevant to the role of an anthropogenic ecology for the species.     

Investigating the weaning process in past populations

The 19th century St. Thomas' Anglican churchyard in Belleville, Ontario, Canada is associated with a large and well-preserved infant skeletal collection (n = 149) and good-quality parish records that document interments in the graveyard (1821–1874). By using a combination of historical demographic and stable nitrogen isotope analyses on the parish records and skeletal remains, respectively, a general pattern of extended nursing for about 14 months, introduction of foods other than breast milk by around 5 months of age, and variation in breast-feeding and weaning behaviours were detected for St. Thomas' infants. The results demonstrate that it is possible to go beyond the concept of weaning age to explore the weaning process in past populations when appropriate and large samples of documentary and skeletal data are available. Am J Phys Anthropol 105:425–439, 1998. © 1998 Wiley-Liss, Inc.     

Lapita diet and subsistence strategies on Watom Island,Papua New Guinea: New stable isotope evidence from humans and animals

Stable isotope ratios (δ¹³C and δ¹⁵N) were analyzed from the bone collagen of individuals (n = 8) from a Lapita burial ground (ca. 2800–2350 BP) on Watom Island, located off northeast New Britain in the Bismarck Archipelago. The aim of this study was to assess the diet and subsistence strategies of humans that lived during the later Lapita period in Near Oceania. To aid in the interpretation of the human diet we analyzed the stable isotope ratios of faunal material from the site (n = 27). We also aim to assess methods of animal husbandry at the site over time from an analysis of the stable isotope ratios (δ¹³C and δ¹⁵N) of pig bones (n = 22) from different temporal periods (Lapita, post‐Lapita, and late prehistoric). The protein diet of the humans consisted of marine organisms from the inshore environment and some deep‐water species, most likely marine turtle, in addition to higher trophic level terrestrial foods, likely pig and native animals (e.g., fruit bat, Cuscus and bandicoot). Although the sample sizes were small, females (n = 4) displayed more variable δ¹³C and δ¹⁵N values compared with males (n = 4), which may be associated with the movement of adult females to the island. The stable isotope analysis of the pig bones indicated that there were few differences between the diets of the pigs from the Lapita and post‐Lapita layers, suggesting that the method of pig husbandry was similar between these two periods and was likely relatively free‐range. Am J Phys Anthropol 157:30–41, 2015. © 2015 Wiley Periodicals, Inc.     

Conformity in diversity? Isotopic investigations of infant feeding practices in two iron age populations from Southern Öland,Sweden

This article presents the results of a study of infant diet at two Iron Age sites on the island of Öland, Sweden. The cemetery at Bjärby contained a large number of subadults who had survived the earliest years of life, whereas most individuals at Triberga had died by 6 months of age. To investigate whether differences in infant feeding could explain the different mortality rates, the carbon, nitrogen, and sulfur stable isotope ratios of bone and tooth dentin collagen from the two sites were analyzed. Twenty‐two samples from Triberga and 102 from Bjärby yielded data that could be included in the carbon and nitrogen analysis. Twelve samples from Triberga and 42 from Bjärby were included in the sulfur analysis. The results for carbon (δ¹³C: Triberga X = –18.8, s.d. = 1.1; Bjärby X = –19.8, s.d. = 0.4), nitrogen (δ¹⁵N: Triberga X = 12.9, s.d. = 1.5; Bjärby X = 13.4, s.d. = 1.4), and sulfur (δ³⁴S: Triberga X = 8.1, s.d. = 1.1; Bjärby X = 5.8, s.d. = 1.3) suggest that diet was broadly similar at both sites and based on terrestrial resources. At Bjärby, females and high‐status individuals consumed higher‐trophic level protein than other males from early childhood onward. There was some indication that the contribution of marine resources to the diet may also have differed between the sexes at Triberga. No consistent differences in breast milk intake were observed between the two sites, but there was substantial variation at each. This variation may reflect an influence of gender and social status on infant feeding decisions. Am J Phys Anthropol 149:217–230, 2012. © 2012 Wiley Periodicals, Inc.     

Isotopic fractionation in a large herbivorous insect, the Auckland tree weta

Determining diet and trophic position of species with stable isotopes requires appropriate trophic enrichment estimates between an animal and its potential foods. These estimates are particularly important for cryptic foragers where there is little comparative dietary information. Nonetheless, many trophic enrichment estimates are based on related taxa, without confirmation of accuracy using laboratory trials. We used stable isotope analysis to investigate diet and to resolve trophic relationships in a large endemic insect, the Auckland tree weta (Hemideina thoracica White). Comparisons of isotopes in plant foods fed to captive wetas with isotope ratios in their frass provided variable results, so frass isotope values had limited usefulness as a proxy indicator of trophic level. Isotopic values varied between different tissues, with trophic depletion of ¹⁵N highest in body fat and testes. Tissue fractionation was consistent in captive and wild caught wetas, and isotopic values were not significantly different between the two groups, suggesting that this weta species is primarily herbivorous. Whole-body values in captive wetas demonstrated trophic depletion (Δδ) for δ¹⁵N of about −0.77‰ and trophic enrichment of 4.28‰ for δ¹³C. These values differ from commonly estimated trophic enrichments for both insects and herbivores and indicate the importance of laboratory trials to determine trophic enrichment. Isotopic values for femur muscles from a number of local wild weta populations did not vary consistently with body weight or size, suggesting that juveniles eat the same foods as adults. Considerable variation among individuals within and between populations suggests that isotopic values are strongly influenced by food availability and individual foraging traits.     

Differentiation of western and eastern Baltic Sea cod stocks (Gadus morhua) by means of stable isotope ratios in muscles and otoliths

Stable isotope analysis of Baltic Sea cod from the Bornholm Sea (ICES‐SD 25) and western Baltic (Belt Sea, ICES‐SD 22) revealed significant differences in the δ¹⁵N and δ¹³C values of the dorsal muscle, as well as in the δ¹⁸O values of otoliths. The method pledges to become especially appropriate to the Baltic due to the high variability in oxygen isotope ratios associated with its estuarine nature.     

Stable carbon and nitrogen isotopes in multiple tissues of wild and captive harbor seals (Phoca vitulina) off the California coast

Stable carbon and nitrogen isotope ratios (δ¹³C and δ¹⁵N) of serum, red blood cells (RBC), muscle, and blubber were measured in captive and wild northeast Pacific harbor seals (Phoca vitulina richardii) at three coastal California sites (San Francisco Bay, Tomales Bay, and Channel Islands). Trophic discrimination factors (Δ_{Tissue‐Diet}) were calculated for captive seals and then applied in wild counterparts in each habitat to estimate trophic position and feeding behavior. Trophic discrimination factors for δ¹⁵N of serum (+3.8‰), lipid‐extracted muscle (+1.6‰), and lipid‐blubber (+6.5‰) are proposed to determine trophic position. An offset between RBC and serum of +0.3‰ for δ¹³C and ?0.6‰ for δ¹⁵N was observed, which is consistent with previous research. Specifically, weaner seals (<1 yr) had large offsets, suggesting strong trophic position shifts during this life stage. Isotopic values indicated an average trophic position of 3.6 at both San Francisco Bay and Tomales Bay and 4.2 at Channel Islands. Isotopic means were strongly dependent on age class and also suggested that mean diet composition varies considerably between all locations. Together, these data indicate that isotopic composition of blood fractions can be an effective approach to estimate trophic position and dietary behavior in wild pinnipeds.     

Nitrogen stable isotope ratios in surface sediments, epilithon and macrophytes from upland lakes with differing nutrient status

1. Thirty small upland lakes in Cumbria, Wales, Scotland and Northern Ireland were each visited once during June and July 2000. From each lake, samples of surface sediment epilithon, macrophytes and total dissolved nitrogen (TDN) were collected for nitrogen stable isotope analysis. As part of a wider programme, samples were also collected for chemical analysis and bioassays. 2. Considerable variation was found in δ¹⁵N values in all measured nitrogen compartments. Some regional variation was evident but was generally weak. Sediment and epilithon δ¹⁵N were positively correlated with δ¹⁵N of TDN, suggesting that baseline nitrogen isotope ratios influence those in some organic matter compartments in the lakes. 3. Sediment δ¹⁵N was higher when inorganic nitrogen concentration in the water was low, possibly reflecting reduced isotope fractionation under these conditions. However, this was not the case for epilithon or macrophytes. Sediment δ¹⁵N values were also negatively related to annual nitrogen deposition. 4. Sediment, epilithon and macrophyte δ¹⁵N values all showed significant relations to nutrient limitation in the lakes as determined by algal bioassays. We suggest that sediment δ¹⁵N might be developed as a simple integrating measure of the degree of nitrogen limitation in lakes.     

Inferring origins of migrating insects using isoscapes: a case study using the true armyworm,Mythimna unipuncta,in North America

1. Many important insect pests undertake seasonal migrations at continental scales in response to changes in resource quality and availability. The frequency and timing of these events could be influenced by the impact of climate change on the suitability of the different sites exploited throughout the year, yet, in many cases, little is known about the origin of seasonal populations, as tracking insect movements is extremely challenging due to their small body size. 2. The use of stable isotope measurements in insect tissues combined with the development of tissue‐specific ‘isoscapes’ of modelled geographic isotope patterns presents a potentially valuable but rarely used approach for obtaining such information on important pest species. In this paper it is illustrated how stable hydrogen isotope analyses (δ²H) in wing chitin of the true armyworm (Mythimna (Psuedaletia) unipuncta Haworth), a seasonal migrant, clearly delineated between 2016 spring immigrants and later locally produced moths in southern Ontario, Canada. 3. It is shown that adult moths captured in early fall in Texas were immigrants from farther north, the first direct confirmation of a southward return migration of this species. Stable carbon isotope (δ¹³C) measurements indicated that spring immigrants in Ontario and autumn immigrants in Texas were from exclusively C3 biomes. Stable nitrogen isotope (δ¹⁵N) measurements also provided information on the probability of individuals coming from agriculturally intensive (i.e. higher δ¹⁵N) sites. Finally, several recommendations are provided regarding future research that could improve the Bayesian assignment models and thus improve assignment accuracy.     

Stable Isotope Analysis in Primatology: A Critical Review

Stable isotope analysis has become an important tool in ecology over the last 25 years. A wealth of ecological information is stored in animal tissues in the relative abundances of the stable isotopes of several elements, particularly carbon and nitrogen, because these isotopes navigate through ecological processes in predictable ways. Stable carbon and nitrogen isotopes have been measured in most primate taxonomic groups and have yielded information about dietary content, dietary variability, and habitat use. Stable isotopes have recently proven useful for addressing more fine‐grained questions about niche dynamics and anthropogenic effects on feeding ecology. Here, we discuss stable carbon and nitrogen isotope systematics and critically review the published stable carbon and nitrogen isotope data for modern primates with a focus on the problems and prospects for future stable isotope applications in primatology. Am. J. Primatol. 74:969‐989, 2012. © 2012 Wiley Periodicals, Inc.     

Equations for Lipid Normalization of Carbon Stable Isotope Ratios in Aquatic Bird Eggs

Stable isotope ratios are biogeochemical tracers that can be used to determine the source of nutrients and contaminants in avian eggs. However, the interpretation of stable carbon ratios in lipid-rich eggs is complicated because ¹³C is depleted in lipids. Variation in ¹³C abundance can therefore be obscured by variation in percent lipids. Past attempts to establish an algebraic equation to correct carbon isotope ratios for lipid content in eggs have been unsuccessful, possibly because they relied partly on data from coastal or migratory species that may obtain egg lipids from different habitats than egg protein. We measured carbon, nitrogen and sulphur stable isotope ratios in 175 eggs from eight species of aquatic birds. Carbon, nitrogen and sulphur isotopes were enriched in lipid-extracted egg samples compared with non extracted egg samples. A logarithmic equation using the C∶N ratio and carbon isotope ratio from the non extracted egg tissue calculated 90% of the lipid-extracted carbon isotope ratios within ±0.5‰. Calculating separate equations for eggs laid by species in different habitats (pelagic, offshore and terrestrial-influenced) improved the fit. A logarithmic equation, rather than a linear equation as often used for muscle, was necessary to accurately correct for lipid content because the relatively high lipid content of eggs compared with muscle meant that a linear relationship did not accurately approximate the relationship between percent lipids and the C∶N ratio. Because lipid extraction alters sulphur and nitrogen isotope ratios (and cannot be corrected algebraically), we suggest that isotopic measurement on bulk tissue followed by algebraic lipid normalization of carbon stable isotope ratio is often a good solution for homogenated eggs, at least when it is not possible to complete separate chemical analyses for each isotope.     

Insights into Strata Use of Forest Animals Using the ‘Canopy Effect’

Stable carbon isotope ratios (δ¹³C) of plants increase linearly from ground to canopy. Accordingly, I used δ¹³C for estimating strata use of fig-eating bats. Data suggest that, overall, bats commuted at lower but fed at higher forest strata, and that small bats foraged at lower forest strata than large bats.