Paleolithic public goods games: why human culture and cooperation did not evolve in one step

It is widely agreed that humans have specific abilities for cooperation and culture that evolved since their split with their last common ancestor with chimpanzees. Many uncertainties remain, however, about the exact moment in the human lineage when these abilities evolved. This article argues that cooperation and culture did not evolve in one step in the human lineage and that the capacity to stick to long-term and risky cooperative arrangements evolved before properly modern culture. I present evidence that Homo heidelbergensis became increasingly able to secure contributions form others in two demanding Paleolithic public good games (PPGGs): cooperative feeding and cooperative breeding. I argue that the temptation to defect is high in these PPGGs and that the evolution of human cooperation in Homo heidelberngensis is best explained by the emergence of modern-like abilities for inhibitory control and goal maintenance. These executive functions are localized in the prefrontal cortex and allow humans to stick to social norms in the face of competing motivations. This scenario is consistent with data on brain evolution that indicate that the largest growth of the prefrontal cortex in human evolution occurred in Homo heidelbergensis and was followed by relative stasis in this part of the brain. One implication of this argument is that subsequent behavioral innovations, including the evolution of symbolism, art, and properly cumulative culture in modern Homo sapiens, are unlikely to be related to a reorganization of the prefrontal cortex, despite frequent claims to the contrary in the literature on the evolution of human culture and cognition.     

The Ceprano Skull and the Earliest Peopling of Europe: Overview and Perspectives

This paper deals with the cranial remains belonging to the earliest hominids found in Europe over the past 15 years. Morphological and metric traits were scored and compared with the data on the pleistocenic remains found in the Old World. The cladistical and multivariate analyses – carried out respectively on the morphological and metric traits – seem to suggest that the Asian remains attributed to Homo erectus differ significantly from those found in the western territories (southern Africa and northern Europe). Among such reports, further differences between the earliest African forms (H. habilis and H. ergaster) and the most recent European and African H. heidelbergenis are worth mentioning. Relying upon this research, the Ceprano calvaria seems to represent a new species (H. cepranensis) from which the later forms – specifically H. heidelbergenis – originated. The presence of some heidelbergensis traits observed in nuce on the Ceprano specimen seems to support this scenario.     

Who Killed the Neanderthals?

Homo neanderthalensis, evolved from the European populations of H. heidelbergensis, and shows some special morphological traits, probably due to an adaptation to particular climatic conditions. It also appears that H. neanderthals had a specialized diet which was mostly carnivorous. Anatomically modern humans of the European Upper Palaeolithic seem to differ from the alimentary behaviour that characterized the Neanderthals; their diet was more varied, with a greater contribution from freshwater alimentary resources (molluscs and fishes). Comparison between the various strategies of subsistence adopted by the two species allows us to propose a hypothesis about the extinction of H. neanderthalensis.     

Progressive changes in brain size and musculo-skeletal traits in seven hominoid populations

Neurological complexity has increased over evolutionary time for invertebrates and vertebrates alike, with the hominid brain tripling in size over the last 3 million years. Since magnetic resonance imaging (MRI) studies among humans indicate a significant correlation (meanr>0.40) between individual differences in brain size and general cognitive ability, it is reasonable to hypothesize that increasing brain size confers greater intelligence. However, larger brains have associated costs, taking longer to build and requiring more energy to run. Sufficient advantages must have accrued for them to override these trade-offs. The present paper documents that in hominoids, as brain size increased from 380 to 1364 cm³ over seven hominoid groups (chimpanzees to australopithecines toHomo habilis toHomo erectus to differences amongHomo sapiens), it was accompanied by changes in 74 musculo-skeletal traits (rs=0.90). These occurred on both cranial traits (temporalis fossae, post-orbital constrictions, mandibles, dentition, nuchal muscle attachments) and on post-cranial traits (pelvic widths, femoral heads, tibial plateaus). It is concluded that in the evolutionary competition to find and fill new niches, there was “room at the top” for greater behavioral complexity and larger brain size, leading to cascading effects on other traits.     

Middle pleistocene humans from africa

Patterns of human evolution in the Middle Pleistocene remain poorly understood. There is general consensus that by the onset of this time period, populations ofHomo erectus were dispersed from Africa into Eurasia, including the Far East. In the western part of this range (perhaps in Africa),Homo erectus then produced a daughter lineage exhibiting more advanced characters of the face, braincase and cranial base. How this new species should be defined is currently debated. In my view, fossils from sites such as Bodo and Broken Hill in Africa may be lumped with material from earlier Middle Pleistocene localities in Europe. Such a taxon is appropriately namedHomo heidelbergensis. Whether the hypodigm should be extended to include fossils from China is another question. In any case, this group of hominids is plausibly ancestral to both the specialized Neanderthals of Europe and more modern humans of the later Middle Pleistocene.     

Reconstitution 3D par Computerized-tomography (CT) et endocrâne virtuel du crâne 5 du site de la Sima de Los Huesos (Atapuerca)

The 3d Ct reconstruction and virtual brain endocast of Cranium 5 from the site of “La Sima de los Huesos” (Atapuerca), allows us to get new information to increase the study and knowledge of Homo heidelbergensis population, and to compare this specimen with others in the fossil record, in order to understand the evolutionary process of the brain, focusing on the middle Pleistocene period. Furthermore, we can observe the changes this species (Homo heidelbergensis) has undergone, at least in Sima de los Huesos population making comparative studies with African and Asian middle Pleistocene specimens. We have used the new data to compare European Homo heidelbergensis represented by SH5 with Kabwe, a controversial specimen considered by some authors like the African Homo heidelbergensis representative, in order to establish the similarities and differences between both specimens.     

Towards a unified theory of cooperative breeding: the role of ecology and life history re-examined

We present quantitative models that unify several adaptive hypotheses for the evolution of cooperative breeding in a single framework: the ecological constraints hypothesis, the life-history hypothesis and the benefits-of-philopatry hypothesis. Our goal is to explain interspecific variation in the occurrence of cooperative breeding in terms of interspecific variation in life-history traits and ecological conditions. We analyse two models, according to whether or not helpers can inherit their parents' territory. Major results are (i) territory inheritance always promotes cooperative breeding; (ii) if territories are not inherited, neither ecological constraints nor variation in life-history traits predict interspecific variation in cooperative breeding; and (iii) if territories are inherited, the mechanism of density regulation is crucial in determining which factors promote cooperative breeding. If density dependence acts on the probability to obtain a free territory or on the survival of dispersers, variation in ecological constraints cannot explain variation in cooperative breeding. Lower adult mortality favours helping, not because it reduces the availability of free territories, but because it enhances the direct benefits of helpers. If density dependence acts on fecundity, lower probability of obtaining a free territory and lower survival of dispersers promote cooperative breeding. In this case, lower adult mortality works against the evolution of helping. We suggest that the difference between birds and social insects in the covariance between cooperative breeding and life-history traits is due to different mechanisms of density regulation that operate in these taxa, and we explain how natural selection on habitat choice might have caused these different mechanisms to operate.     

Fission-fusion and the evolution of hominin social systems

The course of hominin evolution has involved successive migrations towards higher absolute latitudes over the past three million years. Poorer habitat quality further from the equator has led to the necessity for groups occupying higher latitudes to live at lower population densities. Coupled with a trend towards increasing group size over this time period, this tendency towards expansion has led to exponential increases in the area requirements of hominin groups, and a concomitant need to adjust foraging patterns. The current analyses suggest that the development of increasingly complex, multi-level fission-fusion social systems could have freed hominins of the foraging constraints imposed by large group sizes and low population densities. Analyses of the fossil record suggest latitudinally-driven differences in area requirements of the australopithecines from East and South Africa, and African and Asian Homo erectus. In contrast, chronologically-driven differences appear between H. erectus as a whole and Homo heidelbergensis, and between H. heidelbergensis and the Neanderthals. These results are discussed in relation to studies of the foraging patterns of primates and hunter-gatherers.     

Exploratory behavior,but not aggressiveness,is correlated with breeding dispersal propensity in the highly philopatric thorn-tailed rayadito

Studies on the relationship between behavioral traits and dispersal are necessary to understand the evolution of dispersal syndromes. Empirical studies have mainly focused on natal dispersal, even though behavioral differences between dispersers and philopatric individuals are suspected to hold through the whole life cycle, potentially affecting breeding dispersal propensity. Using capture–mark–recapture data and behavioral trials in a forest passerine, the thorn-tailed rayadito Aphrastura spinicauda, we describe inter-individual differences in exploratory behavior and aggressiveness, and investigate the relationship between those traits and breeding dispersal. Our study took place in Fray Jorge National Park, north-central Chile, where a relatively isolated population of rayaditos inhabits a naturally fragmented environment. We found that scores for behavioral traits were consistent between years. Exploratory behavior was similar between sexes, while males showed higher levels of aggression towards a conspecific male intruder. Only exploratory behavior was related to breeding dispersal propensity, with fast-exploring rayaditos being more likely to have dispersed between seasons. This finding provides indirect evidence for the existence of a dispersal strategy that could reduce dispersal costs in the fragmented landscape of Fray Jorge. To our knowledge, this is the first study documenting an association between breeding dispersal and exploratory behavior in a wild bird population. A longitudinal individual-based study will help determining whether this association constitutes a behavioral syndrome.     

Development of microsatellite markers in the grey‐crowned babbler (Pomatostomus temporalis)

The grey‐crowned babbler (Pomatostomus temporalis) is a cooperative breeding bird species in which nonbreeding helpers of both sexes care for the young of breeding individuals. To measure the genetic relatedness between breeders and their offspring and helpers, we developed nine microsatellite markers. Most of the loci were highly polymorphic. These loci will be useful in understanding the evolution and maintenance of cooperative breeding and helping behaviour in this species.     

On the Theory of Evolution Versus the Concept of Evolution: Three Observations

Here we address three misconceptions stated by Rice et al. in their observations of our article Paz-y-Mi?o and Espinosa (Evo Edu Outreach 2:655–675, 2009), published in this journal. The five authors titled their note “The Theory of Evolution is Not an Explanation for the Origin of Life.” First, we argue that it is fallacious to believe that because the formulation of the theory of evolution, as conceived in the 1800s, did not include an explanation for the origin of life, nor of the universe, the concept of evolution would not allow us to hypothesize the possible beginnings of life and its connections to the cosmos. Not only Stanley Miller’s experiments of 1953 led scientists to envision a continuum from the inorganic world to the origin and diversification of life, but also Darwin’s own writings of 1871. Second, to dismiss the notion of Rice et al. that evolution does not provide explanations concerning the universe or the cosmos, we identify compelling scientific discussions on the topics: Zaikowski et al. (Evo Edu Outreach 1:65–73, 2008), Krauss (Evo Edu Outreach 3:193–197, 2010), Peretó et al. (Orig Life Evol Biosph 39:395–406, 2009) and Follmann and Brownson (Naturwissenschaften 96:1265–1292, 2009). Third, although we acknowledge that the term Darwinism may not be inclusive of all new discoveries in evolution, and also that creationists and Intelligent Designers hijack the term to portray evolution as ideology, we demonstrate that there is no statistical evidence suggesting that the word Darwinism interferes with public acceptance of evolution, nor does the inclusion of the origin of life or the universe within the concept of evolution. We examine the epistemological and empirical distinction between the theory of evolution and the concept of evolution and conclude that, although the distinction is important, it should not compromise scientific logic.     

Sex,long life and the evolutionary transition to cooperative breeding in birds

Long life is a typical feature of individuals living in cooperative societies. One explanation is that group living lowers mortality, which selects for longer life. Alternatively, long life may make the evolution of cooperation more likely by ensuring a long breeding tenure, making helping behaviour and queuing for breeding positions worthwhile. The benefit of queuing will, however, depend on whether individuals gain indirect fitness benefits while helping, which is determined by female promiscuity. Where promiscuity is high and therefore the indirect fitness benefits of helping are low, cooperation can still be favoured by an even longer life span. We present the results of comparative analyses designed to test the likelihood of a causal relationship between longevity and cooperative breeding by reconstructing ancestral states of cooperative breeding across birds, and by examining the effect of female promiscuity on the relationship between these two traits. We found that long life makes the evolution of cooperation more likely and that promiscuous cooperative species are exceptionally long lived. These results make sense of promiscuity in cooperative breeders and clarify the importance of life-history traits in the evolution of cooperative breeding, illustrating that cooperation can evolve via the combination of indirect and direct fitness benefits.     

Why Are Some Evolutionary Trees in Natural History Museums Prone to Being Misinterpreted?

Today, the picture of an evolutionary tree is a very well-known visual image. It is almost impossible to think of the ancestry and relationships of living beings without it. As natural history museums play a major role in the public understanding of evolution, they often present a wide variety of evolutionary trees. However, many studies have shown (Baum and Offner 2008; Baum et al. 2005; Catley and Novick 2008; Evans 2009; Gregory 2008; Matuk 2007; Meir et al. 2007b; Padian 2008) that even though evolutionary trees have the potential to engage visitors of natural history museums with the phenomena of evolution, many of them unwittingly might lead to misunderstandings about the process. As valuable research and educational institutions, one of the museum’s important missions should be the careful design of their exhibits on evolution considering, for example, common preconceptions visitors often bring, such as the notion that evolution is oriented from simple toward complex organisms (incarnating the idea of a single ladder of life amidst the extraordinary diversity of organisms) and that humans are at the pinnacle of the evolutionary story, as well as na?ve interpretations of phylogenies. Our aim in this article is to show from history where many of these misunderstandings come from and to determine whether five important Western natural history museums inadvertently present “problematic” evolutionary trees (which might lead to non-scientific notions).     

Development of three allele-specific codominant rice <Emphasis Type="Italic">Waxy</Emphasis> gene PCR markers suitable for marker-assisted selection of amylose content and paste viscosity

Four Waxy haplotypes, previously identified as each having a different combination of three single nucleotide polymorphisms (SNPs) in the Waxy gene, were highly correlated with apparent amylose content and pasting properties, which are important grain quality traits for predicting cooked rice (Oryza sativa L.) texture and processing properties (Chen et al. in J Cereal Sci 47:536–545, 2008a; Chen et al. in J Cereal Sci 48:781–788, 2008b). Three allele-specific PCR markers were developed to genotype the three aforementioned functional SNPs in a single PCR amplification. Each marker contained two allele-specific primers and one common primer. For each marker, the two allele-specific primers differed by one base at the 3′-end to provide discrimination of SNP alleles, and were labeled with unique fluorescence probes. An additional mismatched base, the third base from the 3′-end, was inserted in some allele-specific primers to increase selectivity. The amplification step of the PCR thermal cycling program was initially set for 20× touch-down cycles with the annealing temperature of the first cycle approximately 6°C above the thermal melting temperature of all three primers at a touch-down rate of −0.3°C per cycle, and followed by 25× regular thermal cycles with the annealing temperature at their thermal melting temperature. The allelic genotypes for each SNP were distinguished from each other by both their differential primer-allele fluorescences and their amplification product lengths. The simplicity of these assays makes it easy to utilize these markers as part of a marker-assisted selection strategy in rice breeding programs selecting for these important grain quality traits.     

Out of Africa and back again

If the genusHomo did indeed originate in Africa, then it must have spread by about 2 m.y. ago into Asia where it is represented at 1.8 m.y. ago byHomo erectus fossils. This latter species in turn eventually spread back into Africa, as indicated by the 1.4 m.y. old OH 9 calvaria from Olduvai, and into Europe, as indicated by the 800,000 year old Ceprano calvaria from Italy. These hominids are associated only with Oldowan style artefacts of cores, choppers and flakes and were apparently not conversant with Acheulean handaxe technology. It seems that they most probably evolved viaHomo heidelbergensis into the Neanderthals. Meanwhile, a completely separate development originating withHomo ergaster of about 1.7 m.y. ago in Africa and possessing Acheulean handaxe technology evolved via such forms as Ndutu and Steinheim intoHomo sapiens.     

Hadamard Phylogenetic Methods and the <Emphasis Type="Italic">n</Emphasis>-taxon Process

The Hadamard transform (Hendy and Penny, Syst. Zool. 38(4):297–309, 1989; Hendy, Syst. Zool. 38(4):310–321, 1989) provides a way to work with stochastic models for sequence evolution without having to deal with the complications of tree space and the graphical structure of trees. Here we demonstrate that the transform can be expressed in terms of the familiar P[τ]=e ^Q[τ] formula for Markov chains. The key idea is to study the evolution of vectors of states, one vector entry for each taxa; we call this the n-taxon process. We derive transition probabilities for the process. Significantly, the findings show that tree-based models are indeed in the family of (multi-variate) exponential distributions.     

Phenotypic variation in nestlings of a bird of prey under contrasting breeding and diet conditions

Environmental conditions often vary in space and time, and this may explain variation in the expression of phenotypic traits related to individual quality, such as ornamental coloration. Furthermore, the direction and strength of the relationship between coloured trait expression and individual quality might vary under contrasting conditions. These issues have been explored in adult birds but much less so in nestlings, which are more likely to experience different selective pressures and different physiological trade‐offs than adults. Here, we empirically investigated the effects of contrasting breeding and diet conditions on the expression of carotenoid‐based colour traits displayed by marsh harrier (Circus aeruginosus) nestlings. We studied the variation in coloration, body condition, and immune responsiveness of nestlings in four populations over a 5‐year period. We characterized spatiotemporal differences in rearing conditions experienced by C. aeruginosus nestlings in terms of breeding (laying date, clutch size, and number of nestlings hatched and fledged) and diet (percentage of mammal in diet and prey diversity) conditions. We found that breeding conditions influenced the co‐variation between coloration and immune responsiveness in female nestlings, and that diet conditions influenced the condition‐dependence of nestling coloration in later‐hatched nestlings. In addition, breeding conditions influenced nestling body condition and immune responsiveness, whereas diet conditions influenced nestling coloration and body condition. Our study highlights that nestling phenotype (levels of signalling, circulating carotenoids, and immunity) varies both spatially and temporally, and that some of this variation is related to differences in breeding and diet conditions. Moreover, under contrasting conditions, the direction of the relationships between nestling carotenoid‐based coloration and nestling quality may also vary. In order to fully understand the evolution and maintenance of colour traits in nestling birds, studies and experiments should ideally be replicated under contrasting rearing conditions. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Traveller or tourist? Jack Kerouac and the commodification of culture

Jack Kerouac, the author of On The Road, was a central figure of the Beat Generation, a generation which rebelled against middle-class conformity in post–World War II America. Kerouac described himself as “a religious wanderer” (Kerouac 2006: 2), but an examination of his texts and life suggest his travels may also be understood as tourism. Viewed through the prism of tourism, this study will argue, for example, that MacCannell’s notion of the tourist’s quest for reality and authenticity (MacCannell 1989: 3) provides some insight into why Kerouac wrote that just south of Macon, Georgia, he and his travelling companion Neal Cassady stopped and got out of the car, “and suddenly both of us were stoned with joy to realize that in the darkness all around us was fragrant green grass and the smell of fresh manure and warm waters” (Kerouac 1957: 115). As Kerouac rebelled against being, as one of his protagonists in The Dharma Bums put it, “imprisoned in a system of work, produce, consume, work, produce, consume” (Kerouac 2006: 73) he travelled across America on a rapidly improving network of highways, turning “mobility into a retreat” (Holladay and Holton 2009: 42). Kerouac alternately identified himself as a hobo (Kerouac 1973: 181) and “not a real hobo” (Kerouac 1973: 173), but this article asks whether Kerouac’s travels were those of the last in a line of wanderers rebelling against conformity and modernization or a precursor of mobile mass tourism in America.