In nature, many different types of complex system form hierarchical, self-similar or fractal-like structures that have evolved to maximize internal efficiency. In this paper, we ask whether hunter-gatherer societies show similar structural properties. We use fractal network theory to analyse the statistical structure of 1189 social groups in 339 hunter-gatherer societies from a published compilation of ethnographies. We show that population structure is indeed self-similar or fractal-like with the number of individuals or groups belonging to each successively higher level of organization exhibiting a constant ratio close to 4. Further, despite the wide ecological, cultural and historical diversity of hunter-gatherer societies, this remarkable self-similarity holds both within and across cultures and continents. We show that the branching ratio is related to density-dependent reproduction in complex environments and hypothesize that the general pattern of hierarchical organization reflects the self-similar properties of the networks and the underlying cohesive and disruptive forces that govern the flow of material resources, genes and non-genetic information within and between social groups. Our results offer insight into the energetics of human sociality and suggest that human social networks self-organize in response to similar optimization principles found behind the formation of many complex systems in nature.

Humans have a dual nature. We are subject to the same natural laws and forces as other species yet dominate global ecology and exhibit enormous variation in energy use, cultural diversity, and apparent social organization. We suggest scientists tackle these challenges with a macroecological approach-using comparative statistical techniques to identify deep patterns of variation in large datasets and to test for causal mechanisms. We show the power of a metabolic perspective for interpreting these patterns and suggesting possible underlying mechanisms, one that focuses on the exchange of energy and materials within and among human societies and with the biophysical environment. Examples on human foraging ecology, life history, space use, population structure, disease ecology, cultural and linguistic diversity patterns, and industrial and urban systems showcase the power and promise of this approach.

Life-history theory posits a fundamental trade-off between number and size of offspring that structures the variability in parental investment across and within species. We investigate this 'quantity-quality' trade-off across primates and present evidence that a similar trade-off is also found across natural-fertility human societies. Restating the classic Smith-Fretwell model in terms of allometric scaling of resource supply and offspring investment predicts an inverse scaling relation between birth rate and offspring size and a K 1 ⁄ 4 power scaling between birth rate and body size. We show that these theoretically predicted relationships, in particular the inverse scaling between number and size of offspring, tend to hold across increasingly finer scales of analyses (i.e. from mammals to primates to apes to humans). The advantage of this approach is that the quantity-quality trade-off in humans is placed into a general framework of parental investment that follows directly from first principles of energetic allocation.

Life expectancy is increasing in most countries and has exceeded 80 in several, as low-mortality nations continue to make progress in averting deaths. The health and economic implications of mortality reduction have been given substantial attention, but the observed malleability of human mortality has not been placed in a broad evolutionary context. We quantify the rate and amount of mortality reduction by comparing a variety of human populations to the evolved human mortality profile, here estimated as the average mortality pattern for ethnographically observed hunter-gatherers. We show that human mortality has decreased so substantially that the difference between hunter-gatherers and today's lowest mortality populations is greater than the difference between huntergatherers and wild chimpanzees. The bulk of this mortality reduction has occurred since 1900 and has been experienced by only about 4 of the roughly 8,000 human generations that have ever lived. Moreover, mortality improvement in humans is on par with or greater than the reductions in mortality in other species achieved by laboratory selection experiments and endocrine pathway mutations. This observed plasticity in age-specific risk of death is at odds with conventional theories of aging.biodemography | cross-species comparison | life history evolution | phenotypic plasticity | human lifespan

There persist two widely held but mutually inconsistent views on the evolution of post-fertile lifespan of human females. The first, prevalent within anthropology, sees post-fertile lifespan (PFLS) in the light of adaptive processes, focusing on the social and economic habits of humans that selected for a lengthy PFLS. This view rests on the assumption that human PFLS is distinct from that of other species, and focuses on quantifying the selective causes and consequences of that difference. The second view, prevalent within gerontology and comparative biology, emphasizes that PFLS is a phylogenetically widespread trait or that human PFLS is predictable based on life-history allometries. In this view, human PFLS is part of a broad cross-species pattern and its genesis cannot, therefore, rely on human-specific traits. Those who advocate the second view have questioned the "special pleading" for human specific explanations of PFLS, and have argued that human PFLS is quantitatively greater but not qualitatively different than PFLS in many other animals. Papers asking whether human PFLS is explained by the importance of mothers more than grandmothers, whether paternal or maternal grandparents have more of an effect on child survival, or who is providing the excess calories are associated with the first view that assumes the need to explain the existence of human PFLS on the basis of a uniquely human socioecology. Anthropologists largely see human PFLS as derived, while comparative gerontologists point to evidence that it is one instance of a ubiquitous cross-species pattern. The two groups generally occupy non-overlapping research circles, in terms of conferences and journals, and therefore interact little enough to largely avoid the need to reconcile their views, allowing the persistence of misconceptions in each field. Our goal is to identify and address the most important of these misconceptions and thereby make clear that both of these seemingly incongruent views contain valid points. We argue that two distinct but related traits have been lumped together under the same concept of "post-reproductive lifespan," one (post-fertile viability) that is tremendously widespread and another (a post-fertile life stage) that is derived to hominins, and that the differences and connections between these two traits are necessary for understanding human life-history evolution.

The biogeographic expansion of modern humans out of Africa began Ϸ50,000 years ago. This expansion resulted in the colonization of most of the land area and habitats throughout the globe and in the replacement of preexisting hominid species. However, such rapid population growth and geographic spread is somewhat unexpected for a large primate with a slow, density-dependent life history. Here, we suggest a mechanism for these outcomes by modifying a simple density-dependent population model to allow varying levels of intraspecific competition for finite resources. Reducing intraspecific competition increases carrying capacities, growth rates, and stability, including persistence times and speed of recovery from perturbations. Our model suggests that the energetic benefits of cooperation in modern humans may have outweighed the slow rate of human population growth, effectively ensuring that once modern humans colonized a region long-term population persistence was near inevitable. Our model also provides insight into the interplay of structural complexity and stability in social species.density dependence ͉ hunter-gatherers ͉ intraspecific competition ͉ population ecology ͉ range expansion A ppoximately 50,000 years ago, modern human populations expanded out of Africa and spread rapidly across the majority of the earth's land surface, colonizing all continents except Antarctica by the end of the late Pleistocene, reaching most major archipelagoes and even the most remote Pacific islands by the late Holocene (1-5). This global geographic range expansion involved both the successful colonization of previously unoccupied regions (i.e., Australia, the Americas, and Polynesia) and the replacement of archaic hominid species in Africa and Eurasia (6). Further, both archaeological (7-9) and genetic (10-17) evidence suggests that human populations established themselves quickly in newly colonized regions and grew rapidly, leading to near-continuous spread and persistence across all colonized continents, except in some parts of northern Eurasia, for example, where localized range contractions may have occurred during the late glacial maximum (18,19). The ability of modern human populations to colonize, compete, and persist in such diverse environments demonstrates a remarkable invasive ability.Life history theory suggests that the ability to colonize new habitat, replace competing species, and persist in the face of environmental perturbations is related to the speed of reproduction: Invasive species tend to have fast life histories that allow high rates of population growth (20). However, although humans have higher fertility rates than our nearest living relatives, the anthropoid apes, we resemble other primates in having slow life histories for a mammal of our body size (21,22). Human life histories are characterized by developmental and reproductive trade-offs, resulting in long life spans, extended parental care and juvenile dependency, male provisioning of females and their offspring, reduced interbirth intervals, and...

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