Tinbergen’s classic “On Aims and Methods of Ethology” (_Zeitschrift für Tierpsychologie, 20_, 1963) proposed four levels of explanation of behavior, which he thought would soon apply to humans. This paper discusses the need for multilevel explanation; Huxley and Mayr’s prior models, and others that followed; Tinbergen’s differences with Lorenz on “the innate”; and Mayr’s ultimate/proximate distinction. It synthesizes these approaches with nine levels of explanation in three categories: phylogeny, natural selection, and genomics (ultimate causes); maturation, sensitive period effects, and routine (...) environmental effects (intermediate causes); and hormonal/metabolic processes, neural circuitry, and eliciting stimuli (proximate causes), as a respectful extension of Tinbergen’s levels. The proposed classification supports and builds on Tinbergen’s multilevel model and Mayr’s ultimate/proximate continuum, adding intermediate causes in accord with Tinbergen’s emphasis on ontogeny. It requires no modification of Standard Evolutionary Theory or The Modern Synthesis, but shows that much that critics claim was missing was in fact part of Neo-Darwinian theory (so named by J. Mark Baldwin in _The American Naturalist_ in 1896) all along, notably reciprocal causation in ontogeny, niche construction, cultural evolution, and multilevel selection. Updates of classical examples in ethology are offered at each of the nine levels, including the neuroethological and genomic findings Tinbergen foresaw. Finally, human examples are supplied at each level, fulfilling his hope of human applications as part of the biology of behavior. This broad ethological framework empowers us to explain human behavior—eventually completely—and vindicates the idea of human nature, and of humans as a part of nature. (shrink)

This paper presents and criticizes. Alexander's evolutionary theory of morality (1987). Earlier research, on which Alexander's theory is based, is also reviewed. The propensity to create moral systems evolved because it allowed ancestral humans to limit conflict within cooperating groups and thus form larger groups, which were advantageous because of intense between-group competition. Alexander sees moral codes as contractual, and the primary criticism of his theory is that moral codes are not completely contractual but also coercive. Ways of evaluating Alexander's (...) theory as well as modified versions of it are discussed. (shrink)

Alloparenting, wherein people provide care to children who are not their biological offspring, is a key aspect of human child-rearing. In the Pacific, many children are adopted or fostered by custodial alloparents even when both biological parents are still alive. From a behavioral ecology perspective, such behaviors are puzzling: why parent someone else’s child at your expense? Furthermore, little is known about how these arrangements are made in Pacific Islander societies today, who provides care, and what kinds of outcomes fostered (...) children experience. A better understanding of these proximate factors may help reveal the ultimate drivers behind custodial alloparenting. Here, we report findings from a survey carried out with the caregivers of 282 children in rural areas of Vanuatu, an island nation in Melanesia. Most fostered and adopted children lived with relatives such as aunts, uncles, and grandparents (87.5%) rather than unrelated caregivers, with a strong preference for maternal kin. The most common reasons for these arrangements were that the parents had separated (16.7%), were engaging in labor migration (27.1%), or a combination of both (27.1%). Results for investment in children’s education and their educational outcomes were mixed, although children removed from crisis situations did more poorly than children removed for aspirational reasons. Our findings suggest that custodial alloparenting helps families adapt to socioeconomic transitions and changing marriage practices. Outcomes may depend on a range of factors, such as the reason children were transferred out of the natal home to begin with. (shrink)

Proponents of the standard evolutionary biology paradigm explain human “altruism” in terms of either nepotism or strict reciprocity. On that basis our underlying nature is reduced to a function of inclusive fitness: human nature has to be totally selfish or nepotistic. Proposed here are three possible paths to giving costly aid to nonrelatives, paths that are controversial because they involve assumed pleiotropic effects or group selection. One path is pleiotropic subsidies that help to extend nepotistic helping behavior from close family (...) to nonrelatives. Another is “warfare”—if and only if warfare recurred in the Paleolithic. The third and most plausible hypothesis is based on the morally based egalitarian syndrome of prehistoric hunter-gatherers, which reduced phenotypic variation at the within-group level, increased it at the between-group level, and drastically curtailed the advantages of free riders. In an analysis consistent with the fundamental tenets of evolutionary biology, these three paths are evaluated as explanations for the evolutionary development of a rather complicated human social nature. (shrink)

Chagnon’s analysis of a well-known axe fight in the Yanomamö village of Mishimishiböwei-teri (Chagnon and Bugos 1979) is among the earliest empirical tests of kin selection theory for explaining cooperation in humans. Kin selection theory describes how cooperation can be organized around genetic kinship and is a fundamental tool for understanding cooperation within family groups. Previous analysis on groups of cooperative Lamaleran whale hunters suggests that the role of genetic kinship as a principle for organizing cooperative human groups could be (...) less important in certain cases than previously thought (Alvard Human Nature 14:129–163, 2003b). Evidence that supports a strong role for genetic kinship—groups are found to be more related than expected by chance—may be spurious because of the correlation between social structure and genetic kinship. Reanalysis of Chagnon’s data using matrix regression techniques, however, confirms that genetic kinship was the primary organizing principle in the axe fight; affinal relations were also important, whereas lineage identity explained nothing. (shrink)