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Accumulating isotopic evidence from fossil hominin tooth enamel has provided unexpected insights into early hominin dietary ecology. Among the South African australopiths, these data demonstrate significant contributions to the diet of carbon originally fixed by C(4) photosynthesis, consisting of C(4) tropical/savannah grasses and certain sedges, and/or animals eating C(4) foods. Moreover, high-resolution analysis of tooth enamel reveals strong intra-tooth variability in many cases, suggesting seasonal-scale dietary shifts. This pattern is quite unlike that seen in any great apes, even 'savannah' chimpanzees. The overall proportions of C(4) input persisted for well over a million years, even while environments shifted from relatively closed (ca 3 Ma) to open conditions after ca 1.8 Ma. Data from East Africa suggest a more extreme scenario, where results for Paranthropus boisei indicate a diet dominated (approx. 80%) by C(4) plants, in spite of indications from their powerful 'nutcracker' morphology for diets of hard objects. We argue that such evidence for engagement with C(4) food resources may mark a fundamental transition in the evolution of hominin lineages, and that the pattern had antecedents prior to the emergence of Australopithecus africanus. Since new isotopic evidence from Aramis suggests that it was not present in Ardipithecus ramidus at 4.4 Ma, we suggest that the origins lie in the period between 3 and 4 Myr ago.

The oxygen isotope ratio of CO(2) liberated from structural carbonate in tooth enamel apatite was measured at phosphoric acid reaction temperatures of 25 degrees C, 60 degrees C and 90 degrees C, and it was found that apparent acid fractionation factors for pristine enamel, fossilized enamel, and calcite follow different temperature relationships. Using sealed vessel reactions normalized to alpha(25) = 1.01025 (the fractionation factor for calcite at 25 degrees C), the apparent fractionation factor at 90 degrees C (alpha*(90)) for pristine enamel ranged between 1.00771 and 1.00820, and between 1.00695 and 1.00772 for fossilized enamel. Apparent fractionation factors for common acid bath reactions are similar to those for sealed vessel reactions. A significant correlation exists between alpha*(90) and F(-) content, suggesting that change in the acid fractionation factor may be related to the replacement of OH(-) with F(-) during fossilization of bioapatite. These results have important implications for making accurate comparisons between modern and fossil tooth enamel delta(18)O values, and for the uniformity of isotope data produced in different laboratories using different acid reaction temperatures.

Traditional methods of dietary reconstruction do not allow the investigation of dietary variability within the lifetimes of individual hominins. However, laser ablation stable isotope analysis reveals that the delta13C values of Paranthropus robustus individuals often changed seasonally and interannually. These data suggest that Paranthropus was not a dietary specialist and that by about 1.8 million years ago, savanna-based foods such as grasses or sedges or animals eating these foods made up an important but highly variable part of its diet.

We use the oxygen isotopic composition of tooth enamel from multiple mammalian taxa across eastern Africa to present a proxy for aridity. Here we report tooth enamel delta(18)O values of 14 species from 18 locations and classify them according to their isotopic sensitivity to environmental aridity. The species are placed into two groups, evaporation sensitive (ES) and evaporation insensitive (EI). Tooth enamel delta(18)O values of ES animals increase with aridity, whereas the tooth enamel delta(18)O values of EI animals track local meteoric water delta(18)O values, demonstrating that bioapatite delta(18)O values of animals with different behaviors and physiologies record different aspects of the same environment. The enrichment between tooth enamel delta(18)O values of ES and EI animals records the degree of (18)O enrichment between evaporated water (ingested water or body water) and source water, which increases with environmental aridity. Recognition of the ES-EI distinction creates the opportunity to use the (18)O composition of bioapatite as an index of terrestrial aridity.

Laser ablation of tooth enamel was used to analyze stable carbon isotope compositions of teeth of hominins, red deer, and bears from middle Pleistocene sites in the Sierra de Atapuerca in northern Spain, to investigate the possibility that this technique could be used as an additional tool to identify periods of physiological change that are not detectable as changes in tooth morphology. Most of the specimens were found to have minimal intra-tooth variation in carbon isotopes (< 2.3‰), suggesting isotopically uniform diets through time and revealing no obvious periods of physiological change. However, one of the two sampled hominin teeth displayed a temporal carbon isotope shift (3.2‰) that was significantly greater than observed for co-occurring specimens. The δ13C value of this individual averaged about -16‰ early in life, and -13‰ later in life. This isotopic change occurred on the canine crown about 4.2 mm from the root, which corresponds to an approximate age of two to four years old in modern humans. Our dataset is perforce small owing to the precious nature of hominid teeth, but it demonstrates the potential utility of the intra-tooth isotope profile method for extracting ontogenetic histories of human ancestors. [ABSTRACT FROM AUTHOR]

ABSTRACT Objective: Archaeological remains strongly suggest that the Holocene Japanese hunter-gatherers, the Jomon people, utilized terrestrial plants as their primary food source. However, carbon and nitrogen isotope analysis of bone collagen indicates that they primarily exploited marine resources. We hypothesize that this inconsistency stems from the route of protein synthesis and the different proportions of protein-derived carbon in tooth enamel versus bone collagen. Carbon isotope ratios from bone collagen reflect that of dietary protein and may provide a biased signal of diet, whereas isotope ratios from tooth enamel reflect the integrated diet from all macronutrients (carbohydrates, lipids, and proteins). Methods: In order to evaluate the differences in inferred diet between the archaeological evidence and bone collagen isotope data, this study investigated carbon isotopes in Jomon tooth enamel from four coastal sites of the Middle to Late-Final Jomon period (5,000-2,300 years BP). Results: Carbon isotope ratios of human teeth are as depleted as coeval terrestrial mammals, suggesting that C3 plants and terrestrial mammals were major dietary resources for the Jomon people. Dietary dependence on marine resources calculated from enamel was significantly lower than that calculated from bone collagen. The discrepancy in isotopic ratios between enamel and collagen and the nitrogen isotope ratio in collagen shows a negative correlation on individual and population levels, suggesting diets with variable proportions of terrestrial and marine resources. Conclusion: This study highlights the usefulness of coupling tooth enamel and bone collagen in carbon isotopic studies to reconstruct prehistoric human diet. Am J Phys Anthropol 158:300-311, 2015. © 2015 Wiley Periodicals, Inc. [ABSTRACT FROM AUTHOR]

We present dental enamel stable carbon and oxygen isotope data, histological analyses of daily cross striations and perikymata, and microwear data of Late Miocene primates Indopithecus and Sivaladapis nagrii and an early Pleistocene primate Theropithecus delsoni, known from the Indian Siwaliks. The results indicate that the Late Miocene giant ape Indopithecus had a C3 diet, likely including nuts, seeds and fruits with hard cover, and most probably lived in a seasonal forest that experienced long dry and wet seasons. While the seasons changed as depicted by intra-annual variation in δ18O values, the C3 component of diet was similar throughout the year, as indicated by consistent δ13C values. Sivaladapis nagrii dental enamel δ13C and δ18O values, striations and pit percentages indicate that this small primate (3 kg) was a C3 mixed folivore-frugivore. Theropithecus delsoni dental enamel δ13C and δ18O values and microwear pattern indicate a diet comprising mainly C4 grasses or sedges. [ABSTRACT FROM AUTHOR]

Stable carbon isotope analysis in tooth enamel is a well-established approach to infer C3 and C4 dietary composition in fossil mammals. The bulk of past work has been conducted on large herbivorous mammals. One important finding is that their dietary habits of fossil large mammals track the late Miocene ecological shift from C3 forest and woodland to C4 savannah. However, few studies on carbon isotopes of fossil small mammals exist due to limitations imposed by the size of rodent teeth, and the isotopic ecological and dietary behaviors of small mammals to climate change remain unknown. Here we evaluate the impact of ecological change on small mammals by fine-scale comparisons of carbon isotope ratios (δ13C) with dental morphology of murine rodents, spanning 13.8 to ∼2.0 Ma, across the C3 to C4 vegetation shift in the Miocene Siwalik sequence of Pakistan. We applied in-situ laser ablation GC-IRMS to lower first molars and measured two grazing indices on upper first molars. Murine rodents yield a distinct, but related, record of past ecological conditions from large herbivorous mammals, reflecting available foods in their much smaller home ranges. In general, larger murine species show more positive δ13C values and have higher grazing indices than smaller species inhabiting the same area at any given age. Two clades of murine rodents experienced different rates of morphological change. In the faster-evolving clade, the timing and trend of morphological innovations are closely tied to consumption of C4 diet during the vegetation shift. This study provides quantitative evidence of linkages among diet, niche partitioning, and dental morphology at a more detailed level than previously possible. [ABSTRACT FROM AUTHOR]

Theropithecus was a common large-bodied primate that co-occurred with hominins in many Plio-Pleistocene deposits in East and South Africa. Stable isotope analyses of tooth enamel from T. brumpti (4.0-2.5 Ma) and T. oswaldi (2.0-1.0 Ma) in Kenya show that the earliest Theropithecus at 4 Ma had a diet dominated by C4 resources. Progressively, this genus increased the proportion of C4-derived resources in its diet and by 1.0 Ma, had a diet that was nearly 100% C4-derived. It is likely that this diet was comprised of grasses or sedges; stable isotopes cannot, by themselves, give an indication of the relative importance of leaves, seeds, or underground storage organs to the diet of this primate. Theropithecus throughout the 4- to 1-Ma time range has a diet that is more C4-based than contemporaneous hominins of the genera Australopithecus, Kenyanthropus, and Homo; however, Theropithecus and Paranthropus have similar proportions of C4-based resources in their respective diets. [ABSTRACT FROM AUTHOR]

Hominin fossil evidence in the Turkana Basin in Kenya from ca. 4.1 to 1.4 Ma samples two archaic early hominin genera and records some of the early evolutionary history of Paranthropus and Homo. Stable carbon isotopes in fossil tooth enamel are used to estimate the fraction of diet derived from C3 or C4 resources in these hominin taxa. The earliest hominin species in the Turkana Basin, Australopithecus anamensis, derived nearly all of its diet from C3 resources. Subsequently, by ca. 3.3 Ma, the later Kenyanthropus platyops had a very wide dietary range—from virtually a purely C3 resource-based diet to one dominated by C4 resources. By ca. 2 Ma, hominins in the Turkana Basin had split into two distinct groups: specimens attributable to the genus Homo provide evidence for a diet with a ca. 65/35 ratio of C3- to C4-based resources, whereas P. boisei had a higher fraction of C4-based diet (ca. 25/75 ratio). Homo sp. increased the fraction of C4-based resources in the diet through ca. 1.5 Ma, whereas P. boisei maintained its high dependency on C4-derived resources. [ABSTRACT FROM AUTHOR]

Carbon isotope studies of early hominins from southern Africa showed that their diets differed markedly from the diets of extant apes. Only recently, however, has a major influx of isotopic data from eastern Africa allowed for broad taxonomic, temporal, and regional comparisons among hominins. Before 4 Ma, hominins had diets that were dominated by C3 resources and were, in that sense, similar to extant chimpanzees. By about 3.5 Ma, multiple hominin taxa began incorporating 13C-enriched [C4 or crassulacean acid metabolism (CAM)] foods in their diets and had highly variable carbon isotope compositions which are atypical for African mammals. By about 2.5 Ma, Paranthropus in eastern Africa diverged toward C4/CAM specialization and occupied an isotopic niche unknown in catarrhine primates, except in the fossil relations of grass-eating geladas (Theropithecus gelada). At the same time, other taxa (e.g., Australopithecus africanus) continued to have highly mixed and varied C3/C4 diets. Overall, there is a trend toward greater consumption of 13C-enriched foods in early hominins over time, although this trend varies by region. Hominin carbon isotope ratios also increase with postcanine tooth area and mandibular cross-sectional area, which could indicate that these foods played a role in the evolution of australopith masticatory robusticity. The 13C-enriched resources that hominins ate remain unknown and must await additional integration of existing paleodietary proxy data and new research on the distribution, abundance, nutrition, and mechanical properties of C4 (and CAM) plants. [ABSTRACT FROM AUTHOR]

Stable isotope and molecular data suggest that C4 grasses first appeared globally in the Oligocene. In East Africa. stable isotope data from pedogenic carbonate and fossil tooth enamel suggest a first appearance between 15-10 Ma and subsequent expansion during the Plio-Pleistocene. The fossil enamel record has the potential to provide detailed information about the rates of dietary adaptation to this new resource among different herbivore lineages. We present carbon isotope data from 452 fossil teeth that record differential rates of diet change from C3 to mixed C3/C4 or C4 diets among East African herbivore families at seven different time periods during the Late Miocene to the Pliocene (9.9-3.2 Ma). Significant amounts of C4 grasses were present in equid diets beginning at 9.9 Ma and in rhinocerotid diets by 9.6 Ma, although there is no isotopic evidence for expansive C4 grasslands in this part of the Late Miocene. Bovids and hippopotamids followed suit with individuals that had C4-dominated (>65%) diets by 7.4 Ma. Suids adopted C4-dominated diets between 6.5 and 4.2 Ma. Gomphotheriids and elephantids had mostly C3-dominated diets through 9.3 Ma. but became dedicated C4 grazers by 6.5 Ma. Dein otheriids and giraffids maintained a predominantly C3 diet throughout the record. The sequence of differential diet change among herbivore lineages provides ecological insight into a key period of hominid evolution and valuable information for future studies that focus on morphological changes associated with diet change. [ABSTRACT FROM AUTHOR]

Abstract: Mammalian teeth are invaluable archives of ancient seasonality because they record along their growth axes an isotopic record of temporal change in environment, plant diet, and animal behavior. A major problem with the intra-tooth method is that intra-tooth isotope profiles can be extremely time-averaged compared to the actual pattern of isotopic variation experienced by the animal during tooth formation. This time-averaging is a result of the temporal and spatial characteristics of amelogenesis (tooth enamel formation), and also results from laboratory sampling. This paper develops and evaluates an inverse method for reconstructing original input signals from time-averaged intra-tooth isotope profiles. The method requires that the temporal and spatial patterns of amelogenesis are known for the specific tooth and uses a minimum length solution of the linear system Am = d, where d is the measured isotopic profile, A is a matrix describing temporal and spatial averaging during amelogenesis and sampling, and m is the input vector that is sought. Accuracy is dependent on several factors, including the total measurement error and the isotopic structure of the measured profile. The method is shown to accurately reconstruct known input signals for synthetic tooth enamel profiles and the known input signal for a rabbit that underwent controlled dietary changes. Application to carbon isotope profiles of modern hippopotamus canines reveals detailed dietary histories that are not apparent from the measured data alone. Inverse methods show promise as an effective means of dealing with the time-averaging problem in studies of intra-tooth isotopic variation. [Copyright &y& Elsevier]

Temporal changes in the carbon and oxygen isotopic composition of an animal are an environmental and behavioral input signal that is recorded into the enamel of developing teeth. In this paper, we evaluate changes in phosphorus content and density along the axial lengths of three developing ungulate teeth to illustrate the protracted nature of mineral accumulation in a volume of developing enamel. The least mature enamel in these teeth contains by volume about 25% of the mineral mass of mature enamel, and the remaining 75% of the mineral accumulates during maturation. Using data from one of these teeth (a Hippopotamus amphibius canine), we develop a model for teeth growing at constant rate that describes how an input signal is recorded into tooth enamel. The model accounts for both the temporal and spatial patterns of amelogenesis (enamel formation) and the sampling geometry. The model shows that input signal attenuation occurs as a result of time-averaging during amelogenesis when the maturation interval is long compared to the duration of features in the input signal. Sampling does not induce significant attenuation, provided that the sampling interval is several times shorter than the maturation interval. We present a detailed δ13C and δ18O record for the H. amphibius canine and suggest possible input isotope signals that may have given rise to the measured isotope signal. [Copyright &y& Elsevier]

Reconstructing water availability in terrestrial ecosystems is key to understanding past climate and landscapes, but there are few proxies for aridity that are available for use at terrestrial sites across the Cenozoic. The isotopic composition of tooth enamel is widely used as a paleoenvironmental indicator and recent work suggests the potential for using the triple oxygen isotopic composition of the carbonate component of mammalian tooth enamel (Δ′17O enamel) as an indicator of aridity. However, the extent to which Δ′17O enamel values vary across environments is unknown and there is no framework for evaluating past aridity using Δ′17O enamel data. Here we present Δ′17O enamel and δ18O enamel values from 50 extant mammalian herbivores that vary in physiology, behavior, diet, and water-use strategy. Teeth are from sites in Africa, Europe, and North America and represent a range of environments (humid to arid) and latitudes (34°S to 69°N), where mean annual δ18O values of meteoric water range from –26.0‰ to 2.2‰ (VSMOW). Δ′17O enamel values from these sites span 162 per meg (–252 to –90 per meg), where 1 per meg = 0.001‰). The observed variation in Δ′17O enamel values increases with aridity, forming a wedge-shaped pattern in a plot of aridity index vs. Δ′17O enamel that persists regardless of geographic region. In contrast, the plot of aridity index vs. δ18O enamel for these same samples does not yield a distinct pattern. We use these new Δ′17O enamel data from extant teeth to provide guidelines for using Δ′17O enamel data from fossil teeth to assess and classify the aridity of past environments. Δ′17O enamel values from the fossil record have the potential to be a widely used proxy for aridity without the limitations inherent to approaches that use δ18O enamel values alone. In addition, the data presented here have implications for how Δ′17O enamel values of large mammalian herbivores can be used in evaluations of diagenesis and past p CO 2 and past gross primary productivity. [ABSTRACT FROM AUTHOR]

Intratooth stable isotope profiles in enamel provide time series of dietary and environmental information that if correctly interpreted, serve as archives of seasonal variability in past environments. A major challenge in interpreting these profiles arises from time averaging imparted by enamel mineralization and developmental geometry, whereby the primary (δ13C or δ18O) input signal is attenuated and shifted, which can potentially lead to incorrect interpretations of the magnitude or frequency of seasonal variability. Several forward and inverse models have been developed to reconstruct the primary input signal from intratooth profiles in continuously growing teeth. Here the models developed by Passey and Cerling (2002) and Passey et al. (2005) are extended to molars of Elephantinae, which grow over a long but finite interval of time. Proboscidean molars are particularly attractive for intratooth profiles because they may contain a decade or more of information and they are often well preserved in the fossil record because of their thick enamel and large size. Forward model parameters are established using histological analysis of molar thin sections of extant African elephants (Loxodonta africana) and a mammoth (Mammuthus columbi) and by micro-CT analysis of L. africana molar plates. The density of immature enamel is about 65% of the final density of mature enamel. The appositional length varies from approximately 35 to 55 mm, and the maturation length is about 70 mm. Histological methods are used to determine crown formation time (CFT) in elephant and mammoth molar plates. CFT for the elephant and mammoth molar plates studied in thin section are about 5–6 years and 11 years, which translate to mean growth rates of about 21 mm/year and 16 mm/year, respectively. Coeval molar and tusk profiles from a zoo elephant are compared. The tusk isotope profile serves as a proxy for the primary input signal, and thus provides an opportunity to evaluate the forward and inverse models. The results from the zoo elephant profiles demonstrate that the inverse model accurately reconstructs the amplitude and overall structure of the primary input signal. Inverse model results of mammoth molar profiles show double the range of δ13C in measured enamel profiles. Inversion model results illustrate that improved reconstruction of the primary input signal can lead to more accurate interpretations of the seasonal variability of diet and body water and by extension, vegetation and precipitation in past environments. [ABSTRACT FROM AUTHOR]

Intra-tooth stable isotope variations have been used to interpret seasonality and aridity in paleoenvironmental reconstructions of paleontological and archeological sites. However, most intra-tooth datasets only permit qualitative interpretations of seasonality, because the measured signal is attenuated due to the duration of enamel mineralization process and sampling geometry. The common warthog (Phacochoerus africanus) is an ideal organism to investigate stable isotope variation in enamel. Their canines grow continuously through the life of the individual and are therefore excellent candidates for mathematical modeling of seasonal signals and of signal attenuation; further, their isotope profiles (a series of isotope measurements) can be compared to isotope profiles of third molars (M3) to provide insights into environmental reconstructions. We first obtained paired intra-tooth enamel samples from ever-growing canines and hypsodont M3s of two extant common warthog specimens from Laikipia, Kenya. Second, from a different set of specimens, we collected data on enamel growth patterns and geometry using histological thin sections and transmitted light microscopy, and enamel mineralization parameters using micro-CT scans in each tooth type. Third, we reconstructed the timeline of unattenuated seasonal δ18O signal from canine enamel using growth rate estimates and the inverse model of Passey et al. (2005). Our results demonstrate that canines, which capture ~1.5 years of time, exhibit near-constant growth rates and simple enamel maturation geometry, whereas M3s, which also represent ~1.5 years of time, exhibit linearly decreasing growth rates and more complex maturation patterns. We compare the timelines of unattenuated seasonal δ18O signal and measured M3 profiles and find an average signal reduction of ~50% in the M3s, providing interpretations of the duration of seasonal cycles that are consistent 75% of the time. We conclude that warthog canines are well suited for the inverse model approach, and we established the model parameters for the forward and inverse methods. Timeline reconstructions based on M3 histology are promising for investigating the pattern of rainfall seasonality in the past. Finally, we found an unexpected carbon isotopic spacing of ~2‰ between canine and M3 enamel, which suggests caution in interpreting δ13C results from suid canine or molar enamel alone. • Warthog canines and M3s exhibit different growth and mineralization patterns. • Inverse modeling is used in canines to recover unattenuated seasonal δ18O signals. • A 50% reduction in the original signal is found in the δ18O profiles of M3s. • Timeline reconstruction based on M3s permits estimation of seasonal cycle lengths. • There is a carbon isotope spacing of ~2‰ between canine and M3 enamel. [ABSTRACT FROM AUTHOR]