Proving communal warfare among hunter‐gatherers: The quasi‐rousseauan error

Was human fighting always there, as old as our species? Or is it a late cultural invention, emerging after the transition to agriculture and the rise of the state, which began, respectively, only around ten thousand and five thousand years ago? Viewed against the life span of our species, Homo sapiens, stretching back 150,000–200,000 years, let alone the roughly two million years of our genus Homo, this is the tip of the iceberg. We now have a temporal frame and plenty of empirical evidence for the “state of nature” that Thomas Hobbes and Jean‐Jacque Rousseau discussed in the abstract and described in diametrically opposed terms. All human populations during the Pleistocene, until about 12,000 years ago, were hunter‐gatherers, or foragers, of the simple, mobile sort that lacked accumulated resources. Studying such human populations that survived until recently or still survive in remote corners of the world, anthropology should have been uniquely positioned to answer the question of aboriginal human fighting or lack thereof. Yet access to, and the interpretation of, that information has been intrinsically problematic. The main problem has been the “contact paradox.” Prestate societies have no written records of their own. Therefore, documenting them requires contact with literate state societies that necessarily affects the former and potentially changes their behavior, including fighting.     

Ecological and cultural shifts of hunter‐gatherers of the Jomon period paralleled with environmental changes

Objectives

Holocene hunter‐gatherers adapted to climatic and environmental changes over time. Carbon and nitrogen stable isotope analysis of human skeletal remains from the Inariyama shell mound of the Final Jomon period have revealed large dietary variations in the population. This study analyzed radiocarbon dates of these individuals to test temporal changes in diet and its relationship with tooth ablation.

Materials and Methods

Twenty‐nine human skeletal remains from Inariyama were included in this study. Extracted bone collagen samples were purified to graphite. Then, radiocarbon dating of these samples was performed using the accelerator mass spectrometer.

Results

The radiocarbon ages of Inariyama ranged about, 3,230–2,140 cal BP and showed three peaks of occupation. In the early and late phases, terrestrial resource consumption and incisor extraction were observed, while marine resource consumption and canine extraction were observed in the middle phase.

Discussion

These temporal changes of diet and tooth ablation types occurred in parallel with climatic cooling and environmental change and help reveal how Holocene hunter‐gatherers adapted to the changing environments.

     

Activity‐induced dental modification in holocene siberian hunter‐fisher‐gatherers

The use of teeth as tools provides clues to past subsistence patterns and cultural practices. Five Holocene period hunter‐fisher‐gatherer mortuary sites from the south‐western region of Lake Baikal, Siberia, Russian Federation, are observed for activity‐induced dental modification (AIDM) to further characterize their adaptive regimes. Grooves on the occlusal surfaces of teeth are observed in 25 out of 123 individuals (20.3%) and were most likely produced during the processing of fibers from plants and animals, for making items such as nets and cordage. Regional variation in the frequency of individuals with occlusal grooves is found in riverine versus lakeshore sites. This variation suggests that production of material culture items differed, perhaps in relation to different fishing practices. There is also variation in the distribution of grooves by sex: grooves are found predominately in females, except at the Late Neolithic‐Bronze Age river site of Ust'‐Ida I where grooves are found exclusively in males. Occlusal grooves were cast using polyvinylsiloxane and maxillary canine impressions were examined by scanning electron microscopy (SEM) to determine striation patterns. Variation in striae orientation suggests that a variety of activities, and/or different manufacturing techniques, were involved in groove production. Overall, the variability in occlusal groove frequency, sex and regional distribution, and microscopic striae patterns, points to the multiplicity of activities and ways in which people used their mouths and teeth in cultural activities. Am J Phys Anthropol 143:266–278, 2010. © 2010 Wiley‐Liss, Inc.     

Sex‐biased weaning and early childhood diet among middle holocene hunter–gatherers in Central California

This article evaluates age of weaning and early childhood diets of eight males and nine females from a Middle Holocene (4300–3000 BP) site in Central California, CA‐CCO‐548. All individuals died as adults. δ¹⁵N values from serial sections of dentin collagen in first molars suggest females were fully weaned, on average, by 3.6 years of age, about 0.4 years later than males in the sample, suggesting possible greater parental investment in female offspring. However, throughout childhood females consumed lower trophic‐level foods than males. This could indicate greater investment in males through provisioning of higher quality foods, or alternatively, some degree of independent foraging by males starting as early as 2 to 3 years of age. Even as adults, these same males and females consumed a different range of foods as indicated by their bone collagen δ¹³C and δ¹⁵N values. Overall, the data suggest children were enculturated early into their respective gendered diets, with girls consuming greater amounts of plant foods and boys consuming greater amounts of higher‐trophic level fish and meat protein. Am J Phys Anthropol 152:471–483, 2013. © 2013 Wiley Periodicals, Inc.     

The influence of body proportions on femoral and tibial midshaft shape in hunter‐gatherers

Variation in femoral and tibial diaphyseal shape is used as an indicator of adaptation to patterns of terrestrial mobility. Recent experimentation has implied that lower limb diaphyseal shape may be primarily influenced by lower limb length, and less so by mobility patterns. If valid, this would, at most, render previous interpretations of mobility patterns based on analyses of diaphyseal shape questionable, and, at least, require additional standardization that considers the influence of limb length. Although the consequences could be profound, this implication has yet to be directly tested. Additionally, the influence of body breadth on tibial shape (and to a lesser extent femoral shape) remains uncertain. Tibial and femoral cross‐sectional midshaft shape measurements, taken from nine Pleistocene and Holocene skeletal populations, were compared against lower limb length, limb segment length, and bi‐iliac breadth. Generally, limb length and limb segment length do not significantly influence femoral or tibial midshaft shape. After controlling for body mass greater bi‐iliac breadth is associated with a relative mediolateral strengthening of the femoral midshaft, while the influence of a wider body shape (BIB/length) is associated with a relative M‐L strengthening of the tibia and femur of males, and the tibia of females. We conclude that; (1) mechanical interpretations of lower limb diaphyseal shape are most parsimonious due to the lack of evidence for a consistent relationship between segment length and shape; however, (2) further work is required to investigate the influence of bi‐iliac breadth on both femoral and tibial midshaft shape. Am J Phys Anthropol, 2010. © 2010 Wiley‐Liss, Inc.     

Oral microbiomes from hunter‐gatherers and traditional farmers reveal shifts in commensal balance and pathogen load linked to diet

Maladaptation to modern diets has been implicated in several chronic disorders. Given the higher prevalence of disease such as dental caries and chronic gum diseases in industrialized societies, we sought to investigate the impact of different subsistence strategies on oral health and physiology, as documented by the oral microbiome. To control for confounding variables such as environment and host genetics, we sampled saliva from three pairs of populations of hunter‐gatherers and traditional farmers living in close proximity in the Philippines. Deep shotgun sequencing of salivary DNA generated high‐coverage microbiomes along with human genomes. Comparing these microbiomes with publicly available data from individuals living on a Western diet revealed that abundance ratios of core species were significantly correlated with subsistence strategy, with hunter‐gatherers and Westerners occupying either end of a gradient of Neisseria against Haemophilus, and traditional farmers falling in between. Species found preferentially in hunter‐gatherers included microbes often considered as oral pathogens, despite their hosts' apparent good oral health. Discriminant analysis of gene functions revealed vitamin B5 autotrophy and urease‐mediated pH regulation as candidate adaptations of the microbiome to the hunter‐gatherer and Western diets, respectively. These results suggest that major transitions in diet selected for different communities of commensals and likely played a role in the emergence of modern oral pathogens.     

Hunter‐gatherers and human evolution

Although few hunter‐gatherers or foragers exist today, they are well documented in the ethnographic record. Anthropologists have been eager to study them since they assumed foragers represented a lifestyle that existed everywhere before 10,000 years ago and characterized our ancestors into some ill‐defined but remote past. In the past few decades, that assumption has been challenged on several grounds. Ethnographically described foragers may be a biased sample that only continued to exist because they occupied marginal habitats less coveted by agricultural people. 3 In addition, many foragers have been greatly influenced by their association with more powerful agricultural societies. 4 It has even been suggested that Holocene foragers represent a new niche that appeared only with the climatic changes and faunal depletion at the end of the last major glaciation. 5 Despite these issues, the ethnographic record of foragers provides the only direct observations of human behavior in the absence of agriculture, and as such is invaluable for testing hypotheses about human behavioral evolution. 6 .     

A comparison of antemortem tooth loss in human hunter‐gatherers and non‐human catarrhines: Implications for the identification of behavioral evolution in the human fossil record

Middle and Late Pleistocene fossil hominin specimens with severe antemortem tooth loss are often regarded as evidence for the precocious evolution of human‐like behaviors, such as conspecific care or cooking, in ancient hominin species. The goal of this project was to ask whether the theoretical association between antemortem tooth loss and uniquely human behaviors is supported empirically in a large skeletal sample of human hunter‐gatherers, chimpanzees, orangutans, and baboons. Binomial regression modeling in a Bayesian framework allows for the investigation of the effects of tooth class, genus, age, and sex on the likelihood of tooth loss. The results strongly suggest that modern humans experience more antemortem tooth loss than non‐human primates and identify age in years as an important predictor. Once age is accounted for, the difference between the humans and the closest non‐human genus (chimpanzees) is less pronounced; humans are still more likely on average to experience antemortem tooth loss though 95% uncertainty envelopes around the average prediction for each genus show some overlap. These analyses support theoretical links between antemortem tooth loss and modern human characteristics; humans' significantly longer life history and a positive correlation between age and antemortem tooth loss explain, in part, the reason why humans are more likely to experience tooth loss than non‐human primates, but the results do not exclude behavioral differences as a contributing factor. Am J Phys Anthropol 151:252–264, 2013. © 2013 Wiley Periodicals, Inc.     

How two become one: HJURP dimerization drives CENP‐A assembly

EMBO J 32 15, 2113–2124 doi:10.1038/emboj.2013.142; published online June142013Curr Biol 23 9, 764–769 doi:10.1016/j.cub.2013.03.037; published online May062013Curr Biol 23 9, 770–774 doi:10.1016/j.cub.2013.03.042; published online May062013CENP-A containing nucleosomes epigenetically specify centromere position on chromosomes. Deposition of CENP-A into chromatin is mediated by HJURP, a specific CENP-A chaperone. Paradoxically, HJURP binding sterically prevents dimerization of CENP-A, which is critical to form functional centromeric nucleosomes. A recent publication in The EMBO Journal (Zasadzińska et al, 2013) demonstrates that HJURP itself dimerizes through a C-terminal repeat region, which is essential for centromeric assembly of nascent CENP-A.CENP-A containing nucleosomes have a well-established role in the epigenetic specification of centromere position. However, the composition of the CENP-A nucleosome has been the subject of intense investigation and debate (as has been extensively reviewed, e.g., in Black and Cleveland, 2011). X-ray crystallography data, biochemical interaction experiments and in vivo mutational analysis provide strong evidence that CENP-A nucleosomes are octameric (CENP-A/H4/H2A/H2B)₂, analogous to their histone H3-containing counterparts (Tachiwana et al, 2011; Bassett et al, 2012). Alternatively, based primarily on AFM data and nucleosome crosslinking assays, a tetrameric CENP-A/H4/H2A/H2B ‘hemisome'' has been proposed to be present at centromeres, at least during part of the cell cycle (Dalal et al, 2007; Bui et al, 2012). Whether both nucleosome types exist under specific conditions remains an unresolved question. However, recent studies by the Maddox and Black labs have reported single-molecule fluorescence measurements of CENP-A nucleosomes and high-resolution DNA protection assays of centromeric chromatin, respectively, both of which indicate that octamers are the predominant species of CENP-A in vivo (Hasson et al, 2013; Padeganeh et al, 2013).HJURP is the centromeric histone chaperone that is responsible for timely assembly of CENP-A nucleosomes. HJURP binds to soluble CENP-A and is recruited to centromeric chromatin in early G1 phase, concurrently with nascent CENP-A (Stellfox et al, 2013). Importantly, HJURP facilitates CENP-A nucleosome formation in vitro and its transient targeting to non-centromeric chromatin is sufficient to stably deposit CENP-A at these sites in vivo (Barnhart et al, 2011). Together, these observations identify HJURP as a bona fide centromeric CENP-A histone assembly factor.However, there is an apparent discrepancy between the role of HJURP in CENP-A assembly and the octameric nature of CENP-A nucleosomes. The crystal structure of the human prenucleosomal complex clearly shows that HJURP binds to CENP-A/H4 dimers in a manner that precludes CENP-A/H4 hetero-tetramerization (Hu et al, 2011). Interestingly, however, mutational analysis of CENP-A has shown that tetramerization is crucial for centromere assembly (Bassett et al, 2012). Thus, a mechanism must exist to allow for two trimeric HJURP/CENP-A/H4 complexes to coordinately assemble a tetrameric (CENP-A/H4)₂ particle.In this issue, a study by the Foltz lab sheds light on these paradoxical observations (Zasadzińska et al, 2013). Human HJURP contains two C-terminal repeat regions (HJURP C-terminal domains; HCTD). Expression of short fragments of HJURP containing either of these was sufficient to allow for centromere targeting. However, depletion of endogenous HJURP abolished centromere targeting of the C-terminally located HCTD2 fragment, without affecting the localization of the fragment containing HCTD1. These observations suggest that HCTD1 is required for centromere targeting, while HCTD2 allows for HJURP dimerization. Indeed, the authors go on to show that the latter fragment is both necessary and sufficient to form functional dimers of HJURP. RNAi replacement experiments show that HJURP lacking the HCTD2 dimerization domain is incapable of loading nascent CENP-A into centromeres. Importantly, Zasadzińska et al (2013) demonstrate that the defect in CENP-A loading can be directly attributed to a lack of HJURP dimerization. In an elegant experiment where the HCTD2 containing domain is replaced by an unrelated dimerization domain (that of bacterial LacI), CENP-A assembly is rescued to wild-type levels (Figure 1). This indicates that dimerization of HJURP is an essential step in centromeric chromatin assembly and provides a potential mechanism for the assembly of tetrameric (CENP-A/H4)₂ structures into chromatin as precursors to octameric nucleosomes.Open in a separate windowFigure 1Human HJURP contains separate protein domains that are responsible for CENP-A/H4 binding (blue), centromere targeting (brown) and dimerization (red). Full-length HJURP containing all these domains is capable of assembling CENP-A nucleosomes at centromeres (left). Zasadzińska et al (2013) now show that HJURP lacking the dimerization domain is still able to localize to centromeres, but is unable to assemble CENP-A nucleosomes (middle). However, replacement of the HJURP dimerization domain by an exogenous dimerization domain fully rescues the capability to form CENP-A nucleosomes at centromeres (right). These findings show that HJURP dimerization is an essential feature in the process of nucleosome formation, and explain how (CENP-A/H4)₂ tetramers can be formed by a chaperone that exclusively binds to CENP-A/H4 dimers.While the composition of the HJURP complex suggests a likely mechanism for the formation of octameric nucleosomes, this poses a new challenge to the field. Future studies will be needed to dissect how the shielded HJURP-bound state of CENP-A/H4 can transition to a tetramer on DNA. Interestingly, HJURP is not the only histone chaperone that exclusively binds to histone dimers. Crystal structures of trimeric complexes of both Asf1a/H3.1/H4 (English et al, 2006) as well as DAXX/H3.3/H4 (Elsässer et al, 2012) clearly show sterical incompatibility between chaperone binding and histone tetramerization. It follows that efficient chromatin assembly requires a mode for two histone chaperones to deposit histone dimers in a coordinated fashion, e.g., through dimerization as has been shown for Nap1 (McBryant and Peersen, 2004) and now for HJURP. However, dimerization does not appear to be a universal feature for histone chaperones, as a single CAF1 chaperone is able to bind two H3/H4 dimers as well as (H3/H4)₂ tetramers (Winkler et al, 2012). Thus, while deposition of H3.1/H4 at the replication fork may be driven by the high density of pre-assembly complexes, assembly of nucleosomes containing the replacement variant H3.3, H3.1 nucleosomes at DNA damage sites, and CENP-A at the centromere would require a more active form of coordination. Histone chaperone dimerization may therefore be a common feature in the pipeline to chromatin formation.In summary, Zasadzińska et al (2013) propose a solution to a paradox in the assembly pathway of CENP-A. They show that while each HJURP molecule can exclusively bind a single CENP-A/H4 dimer, HJURP itself dimerizes, ultimately allowing for the formation of tetrameric (CENP-A/H4)₂ structures in chromatin. Interestingly, exclusive dimer binding has been observed for a number of histone chaperones, suggesting that chaperone dimerization may be a more general process in the nucleosome assembly pathway.     

When species become generalists: on‐going large‐scale changes in bird habitat specialization

Aim Species specialization is often considered as a stable species characteristic over the short term. However, this assumption has hardly been tested, even though it may impair our ability to track the impoverishment of biodiversity induced by the rapid replacement of specialists by generalists. We tested whether species specialization in birds varied over a short period of time, and assessed whether variations in species specialization influence community‐level metrics of biotic homogenization. Location France. Methods We studied the variations in specialization to habitat closure of 94 bird species over the period 2002–08, accounting for species variations in mean density, habitat preference and migratory status. We then quantified the temporal changes in a community specialization index, which measures functional homogenization. Results Specialization decreased over time for 35 species (37%), while 46 (49%) showed non‐significant negative trends and 13 (14%) had null or non‐significant positive trends. The more a species was specialized at the beginning of the study, the more it generalized. We additionally found that changes in the specialization level were density dependent: 34 species (36%) became more generalist in years of higher densities while only one became more specialized. At the community level, accounting for this inter‐annual variability in species specialization accentuated the trend in the functional homogenization of bird communities. Main conclusions Habitat specialization is a labile ecological trait, which may change in the short term following habitat degradation, density dependence and source–sink dynamics. Accounting for short‐term temporal variations in observed habitat specialization of species can increase our understanding of the effects of global changes on species strategies and community dynamics.     

The emergence and intensification of early hunter‐gatherer niche construction

Hunter‐gatherers, especially Pleistocene examples, are not well‐represented in archeological studies of niche construction. However, as the role of humans in shaping environments over long time scales becomes increasingly apparent, it is critical to develop archeological proxies and testable hypotheses about early hunter‐gatherer impacts. Modern foragers engage in niche constructive behaviors aimed at maintaining or increasing the productivity of their environments, and these may have had significant ecological consequences over later human evolution. In some cases, they may also represent behaviors unique to modern Homo sapiens. Archeological and paleoenvironmental data show that African hunter‐gatherers were niche constructors in diverse environments, which have legacies in how ecosystems function today. These can be conceptualized as behaviorally mediated trophic cascades, and tested using archeological and paleoenvironmental proxies. Thus, large‐scale niche construction behavior is possible to identify at deeper time scales, and may be key to understanding the emergence of modern humans.     

The ecological and evolutionary energetics of hunter‐gatherer residential mobility

Residential mobility is a key aspect of hunter‐gatherer foraging economies and therefore is an issue of central importance in hunter‐gatherer studies. 1 - 7 Hunter‐gatherers vary widely in annual rates of residential mobility. Understanding the sources of this variation has long been of interest to anthropologists and archeologists. The vast majority of hunter‐gatherers who are dependent on terrestrial plants and animals move camp multiple times a year because local foraging patches become depleted and food, material, and social resources are heterogeneously distributed through time and space. In some environments, particularly along coasts, where resources are abundant and predictable, hunter‐gatherers often become effectively sedentary. But even in these special cases, a central question is how these societies have maintained viable foraging economies while reducing residential mobility to near zero.     

Age‐ratio bias among hunter‐based surveys of Eurasian Wigeon Anas penelope based on wing vs. field samples

We compared age and sex ratios among Eurasian Wigeon Anas penelope derived from Danish field observations and hunter‐based shot samples throughout an entire winter. Sex ratios did not differ significantly between the two samples. Overall, first‐year males were more than three times more likely to be represented than adult males in the hunter sample compared with field samples and were 7–20 times overrepresented in the hunting sample at the beginning of the season. These results confirm the need to account for such bias and its temporal variation when using the results of hunting surveys to model population parameters. Hunter‐shot age ratios may provide a long‐term measure of reproductive success of dabbling duck flyway populations given an understanding of such bias.     

Costly signaling and the handicap principle in hunter‐gatherer research: A critical review

It has been argued that men's hunting in many forager groups is not, primarily, a means of family provisioning but is a costly way of signaling otherwise cryptic qualities related to hunting ability. Much literature concerning the signaling value of hunting draws links to Zahavi's handicap principle and the costly signaling literature in zoology. However, although nominally grounded in the same theoretical paradigm, these literatures have evolved separately. Here I review honest signaling theory in both hunter‐gatherer studies and zoology and highlight three issues with the costly signaling literature in hunter‐gather studies: (a) an overemphasis on the demonstration of realized costs, which are neither necessary nor sufficient to diagnose costly signaling; (b) a lack of clear predictions about what specific qualities hunting actually signals; and (c) an insufficient focus on the broadcast effectiveness of hunting and its value as a heuristics for signal recipients. Rather than signaling hunting prowess, hunting might instead facilitate reputation‐building.     

The politics of Eros: ritual dialogue and egalitarianism in three Central African hunter‐gatherer societies

It has become standard practice within feminist anthropology to repudiate any essential relationship between the biological body and cultural identity. In recent years, the ongoing deconstruction of the body has come to seem the only ‘natural’ fact. By contrast, this article seeks to reconnect sex, power, and culture in a positive sense, by identifying a political system in which power is kept in motion through the body. Literally dancing it out, organized Mbuti or Yaka gender groups perform a recurrent ritual repartee where power is continually churned up and funnelled back and forth between coalitions. The graphic somatic language that emerges through these dances suggests an alternative power‐principle: kinetic, erotic, and fundamentally non‐coercive. Here, the drawing back of the collective eye to the anatomical nature of power, with the simultaneous ritual de‐privatization of ‘biology’, explodes the body out into a collective political force. The cultural visibility of the female procreative body in such contexts is striking. Using the core theme of dialogism, I rethink the creative potential of sexual duality, and work towards a new understanding of gender, power, and the body.