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grounds, the fact is that the large brain of anatomically modern humans predates the Upper Paleolithic cultural system by perhaps 150 kya. Perhaps chronically low population densities prevented the cumulative cultural evolution of highly complex tools and symbolic behavior that characterize the Upper Paleolithic and Later Stone Age (Powell et al., 2009). Favorable circumstances that allowed more substantial populations, particularly in western Eurasia after 40 kya and more generally in the Holocene, may have allowed anatomically modern humans to create highly elaborated cultures much along the line of Ayala’s (Chapter 16, this volume) hypothesis about morality. Our hypothesis that culture was generally the leading rather than the lagging variable in the coevolutionary system may not always (or ever) be correct, even late in hominin evolution. Genomic data promise to have a large impact by shedding light on questions that are difficult to resolve with traditional methods.

NEW GENOMIC TOOLS

Whereas paleoanthropologists will make slow progress in solving the many riddles hinted at in the preceding section, the genomics revolution, made possible by the rapidly falling cost of sequencing genomes, is providing important new tools. These methods promise two important contributions. First, they already help us to better understand paleodemography (Rogers and Harpending, 1992). Second, genomic methods can be used to estimate where and when selection has occurred in the human genome.

Mitochondria and autosomal lineage coalescence times record some evidence of past genetic bottlenecks. When population sizes are small, genetic diversity is lost by drift. If a population increases suddenly, as the hominin population did when anatomically modern humans expanded out of Africa, then a larger number of genes will have coalescence times indicating the time when the human population became large enough to sustain higher genetic diversity. Coalescence times are older for autosomes than for mitochondria or Y chromosomes in part because the effective population size for diploid autosomes is four times the size of the population of maternally transmitted haploid mitochondria or paternally transmitted Y chromosomes (Garrigan and Hammer, 2006).

Studies of the mitochondrial and autosomal genomes have given an interesting picture of the demographic expansion out of Africa. A succession of population bottlenecks caused decreasing genetic diversity farther away from our ancestral African homeland (Vigilant et al., 1991; Ramachandran et al., 2005; Liu et al., 2006; Handley et al., 2007; Wallace, Chapter 7, this volume). The populations most distant from one another, measured by the length of the most likely migration path from Africa, are most distant from one another genetically. Thus, the picture



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