Mandibular shape and diet in extant primates: a 3D geometric morphometric analysis.

Establishing a link between mandibular morphology and diet in extant primates has long been a goal in biological anthropology because it should provide important insight into the diets of extinct primates, including fossil hominins. To date, efforts to explore this question have produced mixed results, largely perhaps due to a reliance on the use of 2D morphological data. Here, we report a study where we investigated whether 3D shape data would provide a clearer picture. We used geometric morphometrics to analyse 3D mandibular shape variation in a sample of > 200 primate specimens, representing individuals from 27 species and five families. Two sets of analyses investigated i) whether there was a relationship between mandibular shape and four standard dietary categories and ii) whether there was a relationship between mandibular shape and a well-known index of diet quality. We found an association between mandibular shape and the dietary categories when we employed raw Procrustes coordinates and allometry-free residuals, but the relationship was weak to non-existent when the effects of phylogeny were taken into account. We found no relationship between shape and the diet quality index, no matter whether the data were raw, corrected for the effects of allometry, corrected for the effects of phylogeny, or corrected for the effects of both allometry and phylogeny. Taken together, the results of the two sets of analyses suggest that there is a weak relationship between 3D mandibular shape and diet in extant primates. Allometry and phylogeny appear to be more important influences on the 3D shape of extant primate mandibles than is diet. We conclude from this that 3D analysis of mandibular shape is unlikely to further illuminate the diets of extinct primates, and research efforts should, therefore, be directed elsewhere.

An assessment of the impact of cross-cultural variation in plant macronutrients on the recommendations of the Paleo Diet.

BACKGROUND: One of the main recommendations of the Paleo Diet is that individuals replicate the whole-diet macronutrient ranges of hunter-gatherer diets. These are suggested to be 19%-35% protein, 22%-40% carbohydrate, and 28%-58% fat, by energy. However, the plant food contribution to these ranges was estimated exclusively from Australian data, which is a potential problem. OBJECTIVES: We investigated whether estimates of the contribution of protein, carbohydrate, and fat to hunter-gatherer diets are impacted by using plant data from other regions of the world. METHODS: The values that form the basis of the Paleo Diet' s recommended macronutrient ranges were generated with a set of equations. We combined these equations with published plant macronutrient data from a multi-region sample of 5 hunter-gatherer groups to generate new estimated macronutrient percentages and then statistically compared the old and new estimates. Subsequently, we collated plant macronutrient data for a sample of 10 hunter-gatherer groups from several regions and repeated the exercise. RESULTS: The whole-diet macronutrient percentages we calculated are significantly different from those that underpin the Paleo Diet's recommendations. Additionally, the ranges derived from our whole-diet macronutrient percentages (14%-35% protein, 21%-55% carbohydrate, 12%-58% fat) are markedly wider than those recommended by the Paleo Diet. CONCLUSIONS: The estimated whole-diet macronutrient percentages that form the basis of the Paleo Diet's macronutrient recommendations are not robust. Using plant data from multiple regions leads to significantly different estimates. Additionally, the macronutrient ranges derived from our whole-diet macronutrient percentages overlap with those recommended by the US Department of Agriculture and the WHO. This undercuts one of the main justifications for adopting the Paleo Diet-namely that because it is vastly different from Western diets, it can reduce the probability of experiencing noncommunicable diseases. There may still be reasons for adopting the Paleo Diet rather than a conventional diet, but healthier macronutrient percentages is not one of them.

Does the Locally-Adaptive Model of Archaeological Potential (LAMAP) work for hunter-gatherer sites? A test using data from the Tanana Valley, Alaska.

We report an assessment of the ability of the Locally-Adaptive Model of Archaeological Potential (LAMAP) to estimate archaeological potential in relation to hunter-gatherer sites. The sample comprised 182 known sites in the Tanana Valley, Alaska, which was occupied solely by hunter-gatherers for about 14,500 years. To estimate archaeological potential, we employed physiographic variables such as elevation and slope, rather than variables that are known to vary on short time scales, like vegetation cover. Two tests of LAMAP were carried out. In the first, we used the location of a random selection of 90 sites from all time periods to create a LAMAP model. We then evaluated the model with the remaining 92 sites. In the second test, we built a LAMAP model from 12 sites that pre-date 10,000 cal BP. This model was then tested with sites that post-date 10,000 cal BP. In both analyses, areas predicted to have higher archaeological potential contained higher frequencies of validation sites. The performance of LAMAP in the two tests was comparable to its performance in previous tests using archaeological sites occupied by agricultural societies. Thus, the study extends the use of LAMAP to the task of estimating archaeological potential of landscapes in relation to hunter-gatherer sites.

Rainfall, temperature, and Classic Maya conflict: A comparison of hypotheses using Bayesian time-series analysis.

Studies published over the last decade have reached contrasting conclusions regarding the impact of climate change on conflict among the Classic Maya (ca. 250-900 CE). Some researchers have argued that rainfall declines exacerbated conflict in this civilisation. However, other researchers have found that the relevant climate variable was increasing summer temperatures and not decreasing rainfall. The goal of the study reported here was to test between these two hypotheses. To do so, we collated annually-resolved conflict and climate data, and then subjected them to a recently developed Bayesian method for analysing count-based times-series. The results indicated that increasing summer temperature exacerbated conflict while annual rainfall variation had no effect. This finding not only has important implications for our understanding of conflict in the Maya region during the Classic Period. It also contributes to the ongoing discussion about the likely impact of contemporary climate change on conflict levels. Specifically, when our finding is placed alongside the results of other studies that have examined temperature and conflict over the long term, it is clear that the impact of climate change on conflict is context dependent.

A 3D basicranial shape-based assessment of local and continental northwest European ancestry among 5th to 9th century CE Anglo-Saxons.

The settlement of Great Britain by Germanic-speaking people from continental northwest Europe in the Early Medieval period (early 5th to mid 11th centuries CE) has long been recognised as an important event, but uncertainty remains about the number of settlers and the nature of their relationship with the preexisting inhabitants of the island. In the study reported here, we sought to shed light on these issues by using 3D shape analysis techniques to compare the cranial bases of Anglo-Saxon skeletons to those of skeletons from Great Britain that pre-date the Early Medieval period and skeletons from Denmark that date to the Iron Age. Analyses that focused on Early Anglo-Saxon skeletons indicated that between two-thirds and three-quarters of Anglo-Saxon individuals were of continental northwest Europe ancestry, while between a quarter and one-third were of local ancestry. In contrast, analyses that focused on Middle Anglo-Saxon skeletons suggested that 50-70% were of local ancestry, while 30-50% were of continental northwest Europe ancestry. Our study suggests, therefore, that ancestry in Early Medieval Britain was similar to what it is today-mixed and mutable.

The composition of the founding population of Iceland: A new perspective from 3D analyses of basicranial shape.

The settlement of Iceland in the Viking Age has been the focus of much research, but the composition of the founding population remains the subject of debate. Some lines of evidence suggest that almost all the founding population were Scandinavian, while others indicate a mix of Scandinavians and people of Scottish and Irish ancestry. To explore this issue further, we used three-dimensional techniques to compare the basicrania of skeletons from archaeological sites in Iceland, Scandinavia, and the British Isles. Our analyses yielded two main results. One was that the founding population likely consisted of roughly equal numbers of Scandinavians and people from the British Isles. The other was that the immigrants who originated from the British Isles included individuals of southern British ancestry as well as individuals of Scottish and Irish ancestry. The first of these findings is consistent with the results of recent analyses of modern and ancient DNA, while the second is novel. Our study, therefore, strengthens the idea that the founding population was a mix of Scandinavians and people from the British Isles, but also raises a new possibility regarding the regions from which the settlers originated.

Spondylolysis and spinal adaptations for bipedalism: The overshoot hypothesis.

BACKGROUND AND OBJECTIVES: The study reported here focused on the aetiology of spondylolysis, a vertebral pathology usually caused by a fatigue fracture. The goal was to test the Overshoot Hypothesis, which proposes that people develop spondylolysis because their vertebral shape is at the highly derived end of the range of variation within Homo sapiens. METHODOLOGY: We recorded 3D data on the final lumbar vertebrae of H. sapiens and three great ape species, and performed three analyses. First, we compared H. sapiens vertebrae with and without spondylolysis. Second, we compared H. sapiens vertebrae with and without spondylolysis to great ape vertebrae. Lastly, we compared H. sapiens vertebrae with and without spondylolysis to great ape vertebrae and to vertebrae of H. sapiens with Schmorl's nodes, which previous studies have shown tend to be located at the ancestral end of the range of H. sapiens shape variation. RESULTS: We found that H. sapiens vertebrae with spondylolysis are significantly different in shape from healthy H. sapiens vertebrae. We also found that H. sapiens vertebrae with spondylolysis are more distant from great ape vertebrae than are healthy H. sapiens vertebrae. Lastly, we found that H. sapiens vertebrae with spondylolysis are at the opposite end of the range of shape variation than vertebrae with Schmorl's nodes. CONCLUSIONS: Our findings indicate that H. sapiens vertebrae with spondylolysis tend to exhibit highly derived traits and therefore support the Overshoot Hypothesis. Spondylolysis, it appears, is linked to our lineage's evolutionary history, especially its shift from quadrupedalism to bipedalism.Lay summary: Spondylolysis is a relatively common vertebral pathology usually caused by a fatigue fracture. There is reason to think that it might be connected with our lineage's evolutionary shift from walking on all fours to walking on two legs. We tested this idea by comparing human vertebrae with and without spondylolysis to the vertebrae of great apes. Our results support the hypothesis. They suggest that people who experience spondylolysis have vertebrae with what are effectively exaggerated adaptations for bipedalism.

Breastfeeding Duration and the Social Learning of Infant Feeding Knowledge in Two Maya Communities.

Variation in the durations of exclusive breastfeeding (exBF) and any breastfeeding (anyBF) is associated with socioecological factors. This plasticity in breastfeeding behavior appears adaptive, but the mechanisms involved are unclear. With this concept in mind, we investigated whether durations of exBF and anyBF in a rural Maya population covary with markers of a form of socioecological change-market integration-and whether individual factors (individual learning, physiological plasticity) and/or learning from others in the community (social learning, norm adherence) mediate these changes. Using data from 419 mother-child pairs from two Guatemalan Maya villages, we fit a bivariate linear mixed model. The model compared exBF and anyBF among children from households of varying degrees of market integration whose mothers follow what we inferred to be local infant-feeding norms. It controlled for other factors expected to affect breastfeeding durations. We found evidence that exBF is associated with whether mothers follow their population's infant feeding norms, but no evidence that exBF is associated with the household's level of market integration. Conversely, anyBF is significantly associated with the household's market integration, but not with the villages' inferred norms. Because deviations from exBF norms are likely to result in infant mortality and reduced fitness, we hypothesize that the incentive to conform is relatively strong. Relatively greater individual plasticity in anyBF allows mother-child pairs to tailor it to socioecological conditions. Deviations from anyBF norms may be tolerated because they may provide later-life health/fitness payoffs, while posing few risks to infant survival.

3D shape analyses of extant primate and fossil hominin vertebrae support the ancestral shape hypothesis for intervertebral disc herniation.

BACKGROUND: Recently we proposed an evolutionary explanation for a spinal pathology that afflicts many people, intervertebral disc herniation (Plomp et al. [2015] BMC Evolutionary Biology 15, 68). Using 2D data, we found that the bodies and pedicles of lower vertebrae of pathological humans were more similar in shape to those of chimpanzees than were those of healthy humans. Based on this, we hypothesized that some individuals are more prone to intervertebral disc herniation because their vertebrae exhibit ancestral traits and therefore are less well adapted for the stresses associated with bipedalism. Here, we report a study in which we tested this "Ancestral Shape Hypothesis" with 3D data from the last two thoracic and first lumbar vertebrae of pathological Homo sapiens, healthy H. sapiens, Pan troglodytes, and several extinct hominins. RESULTS: We found that the pathological and healthy H. sapiens vertebrae differed significantly in shape, and that the pathological H. sapiens vertebrae were closer in shape to the P. troglodytes vertebrae than were the healthy H. sapiens vertebrae. Additionally, we found that the pathological human vertebrae were generally more similar in shape to the vertebrae of the extinct hominins than were the healthy H. sapiens vertebrae. These results are consistent with the predictions of the Ancestral Shape Hypothesis. Several vertebral traits were associated with disc herniation, including a vertebral body that is both more circular and more ventrally wedged, relatively short pedicles and laminae, relatively long, more cranio-laterally projecting transverse processes, and relatively long, cranially-oriented spinous processes. We found that there are biomechanical and comparative anatomical reasons for suspecting that all of these traits are capable of predisposing individuals to intervertebral disc herniation. CONCLUSIONS: The results of the present study add weight to the hypothesis that intervertebral disc herniation in H. sapiens is connected with vertebral shape. Specifically, they suggest that individuals whose vertebrae are towards the ancestral end of the range of shape variation within H. sapiens have a greater propensity to develop the condition than other individuals. More generally, the study shows that evolutionary thinking has the potential to shed new light on human skeletal pathologies.

Potential adaptations for bipedalism in the thoracic and lumbar vertebrae of Homo sapiens: A 3D comparative analysis.

A number of putative adaptations for bipedalism have been identified in the hominin spine. However, it is possible that some have been overlooked because only a few studies have used 3D and these studies have focused on cervical vertebrae. With this in mind, we used geometric morphometric techniques to compare the 3D shapes of three thoracic and two lumbar vertebrae of Homo sapiens, Pan troglodytes, Gorilla gorilla, and Pongo pygmaeus. The study had two goals. One was to confirm the existence of traits previously reported to distinguish the thoracic and lumbar vertebrae of H. sapiens from those of the great apes. The other was to, if possible, identify hitherto undescribed traits that differentiate H. sapiens thoracic and lumbar vertebrae from those of the great apes. Both goals were accomplished. Our analyses not only substantiated a number of traits that have previously been discussed in the literature but also identified four traits that have not been described before: (1) dorsoventrally shorter pedicles in the upper thoracic vertebrae; (2) dorsoventrally longer laminae in all five of the vertebrae examined; (3) longer transverse processes in the upper thoracic vertebrae; and (4) craniocaudally 'pinched' spinous process tips in all of the vertebrae examined. A review of the biomechanical literature suggests that most of the traits highlighted in our analyses can be plausibly linked to bipedalism, including three of the four new ones. As such, the present study not only sheds further light on the differences between the spines of H. sapiens and great apes but also enhances our understanding of how the shift to bipedalism affected the hominin vertebral column.

Radiocarbon dating uncertainty and the reliability of the PEWMA method of time-series analysis for research on long-term human-environment interaction.

Statistical time-series analysis has the potential to improve our understanding of human-environment interaction in deep time. However, radiocarbon dating-the most common chronometric technique in archaeological and palaeoenvironmental research-creates challenges for established statistical methods. The methods assume that observations in a time-series are precisely dated, but this assumption is often violated when calibrated radiocarbon dates are used because they usually have highly irregular uncertainties. As a result, it is unclear whether the methods can be reliably used on radiocarbon-dated time-series. With this in mind, we conducted a large simulation study to investigate the impact of chronological uncertainty on a potentially useful time-series method. The method is a type of regression involving a prediction algorithm called the Poisson Exponentially Weighted Moving Average (PEMWA). It is designed for use with count time-series data, which makes it applicable to a wide range of questions about human-environment interaction in deep time. Our simulations suggest that the PEWMA method can often correctly identify relationships between time-series despite chronological uncertainty. When two time-series are correlated with a coefficient of 0.25, the method is able to identify that relationship correctly 20-30% of the time, providing the time-series contain low noise levels. With correlations of around 0.5, it is capable of correctly identifying correlations despite chronological uncertainty more than 90% of the time. While further testing is desirable, these findings indicate that the method can be used to test hypotheses about long-term human-environment interaction with a reasonable degree of confidence.

The ex-pat effect: presence of recent Western immigrants is associated with changes in age at first birth and birth rate in a Maya population from rural Guatemala.

BACKGROUND: Economic transitions expose indigenous populations to a variety of ecological and cultural challenges, especially regarding diet and stress. These kinds of challenges are predicted by evolutionary ecological theory to have fitness consequences (differential reproduction) and, indeed, are often associated with changes in fertility dynamics. It is currently unclear whether international immigration might impact the nature of such an economic transition or its consequences for fertility. AIM: To examine measures of fertility, diet and stress in two economically transitioning Maya villages in Guatemala that have been differentially exposed to immigration by Westerners. SUBJECTS AND METHODS: This study compared Maya women's ages at first birth and birth rates between villages and investigated whether these fertility indicators changed through time. It also explored whether the villages differed in relation to diet and/or a proxy of stress. RESULTS: It was found that, in the village directly impacted by immigration, first births occurred earlier, but birth rate was slower. In both villages, over the sampled time window, age at first birth increased, while birth rate decreased. The villages do not differ significantly in dietary indicators, but the immigration-affected village scored higher on the stress proxy. CONCLUSION: Immigration can affect fertility in host communities. This relationship between immigration and fertility dynamics may be partly attributable to stress, but this possibility should be evaluated prospectively in future research.

Energy-related influences on variation in breastfeeding duration among indigenous Maya women from Guatemala.

OBJECTIVES: The causes of variation in breastfeeding duration in humans are poorly understood, but life history factors related to maternal energetics drive much of the variation in lactation duration in nonhuman animals. With this in mind, we investigated whether four energy-related factors influence variation in breastfeeding duration in a non-industrial human population: (1) mortality risk during mother's development (assessed via mother's adult height), (2) reliance on nutrient-dense weaning foods, (3) access to and need for help with infant feeding and care ("allomaternal care"), and (4) maternal tradeoffs between current and future reproduction (measured via child's birth order). MATERIALS AND METHODS: The data pertain to 51 Kakchiquel-speaking Maya mothers and 283 children from a village in rural Guatemala. We developed a linear mixed model to evaluate the relationships between breastfeeding duration and the energy-related factors. RESULTS: Duration of breastfeeding was associated with two of the energy-related factors in the ways we predicted but not with the other two. Contrary to predictions, taller mothers breastfed for shorter periods and we found no evidence that weanling diet quality impacts breastfeeding duration. As predicted, women who had more help with infants breastfed for shorter periods, and later-born infants breastfed longer than earlier-born ones. DISCUSSION: The results regarding allomaternal care suggest that help reduces mothers' lactation demands. The energy saved may be redirected to increasing fecundity or investment in other children. The birth order result suggests that children born to mothers nearing reproductive senescence receive higher levels of investment, which likely impacts children's fitness.

The evolutionary relationships and age of Homo naledi: An assessment using dated Bayesian phylogenetic methods.

Homo naledi is a recently discovered species of fossil hominin from South Africa. A considerable amount is already known about H. naledi but some important questions remain unanswered. Here we report a study that addressed two of them: "Where does H. naledi fit in the hominin evolutionary tree?" and "How old is it?" We used a large supermatrix of craniodental characters for both early and late hominin species and Bayesian phylogenetic techniques to carry out three analyses. First, we performed a dated Bayesian analysis to generate estimates of the evolutionary relationships of fossil hominins including H. naledi. Then we employed Bayes factor tests to compare the strength of support for hypotheses about the relationships of H. naledi suggested by the best-estimate trees. Lastly, we carried out a resampling analysis to assess the accuracy of the age estimate for H. naledi yielded by the dated Bayesian analysis. The analyses strongly supported the hypothesis that H. naledi forms a clade with the other Homo species and Australopithecus sediba. The analyses were more ambiguous regarding the position of H. naledi within the (Homo, Au. sediba) clade. A number of hypotheses were rejected, but several others were not. Based on the available craniodental data, Homo antecessor, Asian Homo erectus, Homo habilis, Homo floresiensis, Homo sapiens, and Au. sediba could all be the sister taxon of H. naledi. According to the dated Bayesian analysis, the most likely age for H. naledi is 912 ka. This age estimate was supported by the resampling analysis. Our findings have a number of implications. Most notably, they support the assignment of the new specimens to Homo, cast doubt on the claim that H. naledi is simply a variant of H. erectus, and suggest H. naledi is younger than has been previously proposed.

The empirical case against the 'demographic turn' in Palaeolithic archaeology.

Recently, it has become commonplace to interpret major transitions and other patterns in the Palaeolithic archaeological record in terms of population size. Increases in cultural complexity are claimed to result from increases in population size; decreases in cultural complexity are suggested to be due to decreases in population size; and periods of no change are attributed to low numbers or frequent extirpation. In this paper, we argue that this approach is not defensible. We show that the available empirical evidence does not support the idea that cultural complexity in hunter-gatherers is governed by population size. Instead, ethnographic and archaeological data suggest that hunter-gatherer cultural complexity is most strongly influenced by environmental factors. Because all hominins were hunter-gatherers until the Holocene, this means using population size to interpret patterns in the Palaeolithic archaeological record is problematic. In future, the population size hypothesis should be viewed as one of several competing hypotheses and its predictions formally tested alongside those of its competitors.This article is part of the themed issue 'Major transitions in human evolution'.

Food Aversions and Cravings during Pregnancy on Yasawa Island, Fiji.

Women often experience novel food aversions and cravings during pregnancy. These appetite changes have been hypothesized to work alongside cultural strategies as adaptive responses to the challenges posed by pregnancy (e.g., maternal immune suppression). Here, we report a study that assessed whether data from an indigenous population in Fiji are consistent with the predictions of this hypothesis. We found that aversions focus predominantly on foods expected to exacerbate the challenges of pregnancy. Cravings focus on foods that provide calories and micronutrients while posing few threats to mothers and fetuses. We also found that women who experience aversions to specific foods are more likely to crave foods that meet nutritional needs similar to those provided by the aversive foods. These findings are in line with the predictions of the hypothesis. This adds further weight to the argument that appetite changes may function in parallel with cultural mechanisms to solve pregnancy challenges.

Population size does not explain past changes in cultural complexity.

Demography is increasingly being invoked to account for features of the archaeological record, such as the technological conservatism of the Lower and Middle Pleistocene, the Middle to Upper Paleolithic transition, and cultural loss in Holocene Tasmania. Such explanations are commonly justified in relation to population dynamic models developed by Henrich [Henrich J (2004)Am Antiq69:197-214] and Powell et al. [Powell A, et al. (2009)Science324(5932):1298-1301], which appear to demonstrate that population size is the crucial determinant of cultural complexity. Here, we show that these models fail in two important respects. First, they only support a relationship between demography and culture in implausible conditions. Second, their predictions conflict with the available archaeological and ethnographic evidence. We conclude that new theoretical and empirical research is required to identify the factors that drove the changes in cultural complexity that are documented by the archaeological record.

The Acheulean handaxe: More like a bird's song than a beatles' tune?

The goal of this paper is to provoke debate about the nature of an iconic artifact-the Acheulean handaxe. Specifically, we want to initiate a conversation about whether or not they are cultural objects. The vast majority of archeologists assume that the behaviors involved in the production of handaxes were acquired by social learning and that handaxes are therefore cultural. We will argue that this assumption is not warranted on the basis of the available evidence and that an alternative hypothesis should be given serious consideration. This alternative hypothesis is that the form of Acheulean handaxes was at least partly under genetic control.

The Nature of Culture: an eight-grade model for the evolution and expansion of cultural capacities in hominins and other animals.

Tracing the evolution of human culture through time is arguably one of the most controversial and complex scholarly endeavors, and a broad evolutionary analysis of how symbolic, linguistic, and cultural capacities emerged and developed in our species is lacking. Here we present a model that, in broad terms, aims to explain the evolution and portray the expansion of human cultural capacities (the EECC model), that can be used as a point of departure for further multidisciplinary discussion and more detailed investigation. The EECC model is designed to be flexible, and can be refined to accommodate future archaeological, paleoanthropological, genetic or evolutionary psychology/behavioral analyses and discoveries. Our proposed concept of cultural behavior differentiates between empirically traceable behavioral performances and behavioral capacities that are theoretical constructs. Based largely on archaeological data (the 'black box' that most directly opens up hominin cultural evolution), and on the extension of observable problem-solution distances, we identify eight grades of cultural capacity. Each of these grades is considered within evolutionary-biological and historical-social trajectories. Importantly, the model does not imply an inevitable progression, but focuses on expansion of cultural capacities based on the integration of earlier achievements. We conclude that there is not a single cultural capacity or a single set of abilities that enabled human culture; rather, several grades of cultural capacity in animals and hominins expanded during our evolution to shape who we are today.