Trophic ecology of a Late Pleistocene early modern human from tropical Southeast Asia inferred from zinc isotopes.

Tam Pà Ling, a cave site in northeastern Laos, has yielded the earliest skeletal evidence of Homo sapiens in mainland Southeast Asia. The reliance of Pleistocene humans in rainforest settings on plant or animal resources is still largely unstudied, mainly due to poor collagen preservation in fossils from tropical environments precluding stable nitrogen isotope analysis, the classical trophic level proxy. However, isotopic ratios of zinc (Zn) in bioapatite constitute a promising proxy to infer trophic and dietary information from fossil vertebrates, even under adverse tropical taphonomic conditions. Here, we analyzed the zinc isotope composition (66Zn/64Zn expressed as δ66Zn value) in the enamel of two teeth of the Late Pleistocene (63-46 ka) H. sapiens individual (TPL1) from Tam Pà Ling, as well as 76 mammal teeth from the same site and the nearby Nam Lot cave. The human individual exhibits relatively low enamel δ66Zn values (+0.24‰) consistent with an omnivorous diet, suggesting a dietary reliance on both plant and animal matter. These findings offer direct evidence of the broad utilization of resources from tropical rainforests by one of the earliest known anatomically modern humans in Southeast Asia.

Zinc isotopes in Late Pleistocene fossil teeth from a Southeast Asian cave setting preserve paleodietary information.

Stable carbon and nitrogen isotope ratios of collagen from bone and dentin have frequently been used for dietary reconstruction, but this method is limited by protein preservation. Isotopes of the trace element zinc (Zn) in bioapatite constitute a promising proxy to infer dietary information from extant and extinct vertebrates. The 66Zn/64Zn ratio (expressed as δ66Zn value) shows an enrichment of the heavy isotope in mammals along each trophic step. However, preservation of diet-related δ66Zn values in fossil teeth has not been assessed yet. Here, we analyzed enamel of fossil teeth from the Late Pleistocene (38.4-13.5 ka) mammalian assemblage of the Tam Hay Marklot (THM) cave in northeastern Laos, to reconstruct the food web and assess the preservation of original δ66Zn values. Distinct enamel δ66Zn values of the fossil taxa (δ66Zncarnivore < δ66Znomnivore < δ66Znherbivore) according to their expected feeding habits were observed, with a trophic carnivore-herbivore spacing of +0.60‰ and omnivores having intermediate values. Zn and trace element concentration profiles similar to those of modern teeth also indicate minimal impact of diagenesis on the enamel. While further work is needed to explore preservation for settings with different taphonomic conditions, the diet-related δ66Zn values in fossil enamel from THM cave suggest an excellent long-term preservation potential, even under tropical conditions that are well known to be adverse for collagen preservation. Zinc isotopes could thus provide a new tool to assess the diet of fossil hominins and associated fauna, as well as trophic relationships in past food webs.

Dental phenotypic shape variation supports a multiple dispersal model for anatomically modern humans in Southeast Asia.

The population history of anatomically modern humans (AMH) in Southeast Asia (SEA) is a highly debated topic. The impact of sea level variations related to the Last Glacial Maximum (LGM) and the Neolithic diffusion on past population dispersals are two key issues. We have investigated competing AMH dispersal hypotheses in SEA through the analysis of dental phenotype shape variation on the basis of very large archaeological samples employing two complementary approaches. We first explored the structure of between- and within-group shape variation of permanent human molar crowns. Second, we undertook a direct test of competing hypotheses through a modeling approach. Our results identify a significant LGM-mediated AMH expansion and a strong biological impact of the spread of Neolithic farmers into SEA during the Holocene. The present work thus favors a "multiple AMH dispersal" hypothesis for the population history of SEA, reconciling phenotypic and recent genomic data.

Anatomically modern human in Southeast Asia (Laos) by 46 ka.

Uncertainties surround the timing of modern human emergence and occupation in East and Southeast Asia. Although genetic and archeological data indicate a rapid migration out of Africa and into Southeast Asia by at least 60 ka, mainland Southeast Asia is notable for its absence of fossil evidence for early modern human occupation. Here we report on a modern human cranium from Tam Pa Ling, Laos, which was recovered from a secure stratigraphic context. Radiocarbon and luminescence dating of the surrounding sediments provide a minimum age of 51-46 ka, and direct U-dating of the bone indicates a maximum age of ~63 ka. The cranium has a derived modern human morphology in features of the frontal, occipital, maxillae, and dentition. It is also differentiated from western Eurasian archaic humans in aspects of its temporal, occipital, and dental morphology. In the context of an increasingly documented archaic-modern morphological mosaic among the earliest modern humans in western Eurasia, Tam Pa Ling establishes a definitively modern population in Southeast Asia at ~50 ka cal BP. As such, it provides the earliest skeletal evidence for fully modern humans in mainland Southeast Asia.

Estimating the distribution of probable age-at-death from dental remains of immature human fossils.

In two historic longitudinal growth studies, Moorrees et al. (Am J Phys Anthropol 21 (1963) 99-108; J Dent Res 42 (1963) 1490-1502) presented the "mean attainment age" for stages of tooth development for 10 permanent tooth types and three deciduous tooth types. These findings were presented graphically to assess the rate of tooth formation in living children and to age immature skeletal remains. Despite being widely cited, these graphical data are difficult to implement because there are no accompanying numerical values for the parameters underlying the growth data. This analysis generates numerical parameters from the data reported by Moorrees et al. by digitizing 358 points from these tooth formation graphs using DataThief III, version 1.5. Following the original methods, the digitized points for each age transition were conception-corrected and converted to the logarithmic scale to determine a median attainment age for each dental formation stage. These values are subsequently used to estimate age-at-death distributions for immature individuals using a single tooth or multiple teeth, including estimates for 41 immature early modern humans and 25 immature Neandertals. Within-tooth variance is calculated for each age estimate based on a single tooth, and a between-tooth component of variance is calculated for age estimates based on two or more teeth to account for the increase in precision that comes from using additional teeth. Finally, we calculate the relative probability of observing a particular dental formation sequence given known-age reference information and demonstrate its value in estimating age for immature fossil specimens.

Regional variation in the postcranial robusticity of late Upper Paleolithic humans.

Early modern humans from the European Upper Paleolithic (UP) demonstrate trends in postcranial biomechanical features that coincide with the last glacial maximum (LGM). These features have been interpreted as evidence that ecological changes of the LGM played a critical role in cultural and biological adaptation in European UP populations. In areas outside of Europe, similar environmental changes occurred with the LGM. This analysis introduces postcranial material from the Late Upper Paleolithic (LUP) of North Africa and Southeast Asia and tests two related hypotheses: 1) LUP samples across the Old World had similar patterns of postcranial robusticity and 2) relative to an available Early Upper Paleolithic (EUP) sample, regional LUP samples demonstrate similar trends in robusticity that may be attributable to climatic effects of the LGM. Cross-sectional geometric data of the humeri and femora were obtained for 26 EUP and 100 LUP humans from Europe, Africa, and Asia. Despite regional differences, LUP samples are similar relative to the EUP sample. In the humerus, bilateral asymmetry decreases in all LUP samples relative to the EUP sample. In the femur, LUP samples demonstrate increasingly circular femoral midshaft sections, reflecting reduced anteroposterior bending strength relative to the EUP sample. These patterns suggest changes in subsistence behavior and mobility after the LGM across the Old World that are most consistent with reduced mobility and broad-spectrum resource exploitation.

Morphological variation and airflow dynamics in the human nose.

Airflow dynamics are recognized as being important to the functioning of the human nose in conditioning and filtering inspired air, yet these dynamics are poorly understood. Despite considerable research on airflow dynamics by otolaryngologists, respiratory physiologists, and toxicologists, major disagreements remain about the nature of airflow in the human nose. Specifically, there is little consensus about the character of nasal airflow regimes (laminar or turbulent) and about the major pathways of airflow through the internal chamber. Additionally, a number of features in the human nose have been argued to enhance airflow turbulence, thus increasing the exposure of moving air to the nasal mucosa and facilitating heat and moisture exchange in cold and/or dry climates. These features include: an inferior orientation of the nares; a nasal sill that is high relative to the floor of the internal nasal chamber; a nasal valve that is small in cross-sectional area relative to that of the internal chamber; and large, projecting conchae. The claim that these features affect airflow dynamics has never been tested. To clarify the nature of human nasal airflow and to test these claims of functional significance to nasal variation, we studied airflow across physiological flow rates using water and dye flowing through anatomically accurate acrylic models of human nasal air passageways (with adjustment of water flow rates to maintain dynamic similarity). The models were derived from direct casting of the nasal passageways of 10 Caucasian ("leptorrhine") cadavers (six male, four female). Measures of naris angle, nasal sill height, nasal valve area relative to internal chamber cross-sectional area, and relative projection of the inferior and middle turbinates were taken directly on the resulting casts. The relationships between aspects of nasal morphology and turbulent air flow were evaluated by examining the flow regimes (laminar, semiturbulent, or turbulent) at varying flow rates, with the expectation that the greater the development of the proposed turbulence-enhancing features the slower the flow rate at which flow would shift from one regime to another. Flow characteristics (both flow regimes and principal pathways) were highly variable within our sample. The relative projection of the inferior turbinate was the only variable that significantly affected the flow rate at which flow became turbulent. However, more projecting turbinates appear to laminate flow rather than to induce turbulence. Nostril orientation was moderately correlated with flow dynamics (with more inferiorly directed nares producing turbulence at slower flow rates), but this correlation was not statistically significant. Relative nasal valve area and nasal sill height were unrelated to turbulence in our models.

Late pleistocene human femoral diaphyseal curvature.

Anterior femoral curvature is a consistent characteristic of Pleistocene and recent humans, although variation exists in the degree of curvature among individuals and across populations. In particular, one group, the Neandertals, has been characterized for a century as having marked femoral curvature. To evaluate the degree of anterior femoral curvature in both Neandertals and other Late Pleistocene humans, their curvature subtenses and proximodistal positions were evaluated in the context of recent human variation. Recent human comparisons show little relationship between subtense (absolute curvature) and femoral length, suggesting that an index that incorporates subtense relative to the length of the femur is inappropriate for between-group assessments. Neandertals were statistically indistinguishable from Middle or earlier Upper Paleolithic modern humans in the degree of absolute curvature, all of whom had greater curvature on average than all later humans. Additionally, Neandertals and Qafzeh-Skhul early modern humans had a more distal point of maximum curvature than any other group. Curvature was not strongly correlated with functional considerations including body mass estimates, surrogate variables for body size, proximal femoral articular orientation, or knee anteroposterior dimensions. The functional role of femoral anterior curvature is unknown; however, the general decrease in curvature subtense closely parallels the between-group changes in inferred levels of mobility from femoral diaphyseal robusticity and shape, suggesting that femoral curvature may reflect mobility levels and patterns among Late Pleistocene and recent humans.