Chapter 18 ................................ ................................ Body Size and Intelligence in Hominoid ................................ ................................ chapter Evolution ................................ ................................ Carol Ward ................................ Department of Anthropology – Department of Pathology and Anatomical Sciences ................................ 107 Swallow Hall, University of Missouri ................................ Columbia, MO 65211 USA ................................ [email protected] ................................ Mark Flinn ................................ Department of Anthropology, 107 Swallow Hall, University of Missouri ................................ Columbia, MO 65211 USA ................................ [email protected] ................................ David Begun ................................ Department of Anthropology, University of Toronto ................................ Toronto, ON M5S 3G3, Canada ................................ [email protected] ................................ ................................ Word count: text (6526) XML Typescript c Cambridge University Press – Generated by TechBooks. Body Size and Intelligence in Hominoid Evolution Page 1218 of 1365 A-Head 1 Introduction ................................ ................................ 2 Great apes and humans are the largest brained primates. Aside from a few ................................ 3 extinct subfossil lemurs, they are also the largest in body mass. Body size ................................ 4 is a key aspect of a species’ biology, a large organism having different ................................ 5 energetic, ecological and physical constraints than a small one. Brain size, ................................ 6 insofar as it determines abilities to acquire, process, and act on ................................ 7 information, is also a key aspect of a species’ biology and is linked to body ................................ 8 size. Large animals have different informational problems to solve than do ................................ 9 small ones, hence their respective sensory organs and nervous systems are ................................ 10 sized and organized differently. ................................ 11 Mammalian body and brain size scale consistently with each other ................................c18fig001 12 (Figure 18.1). This relation is generally described by allometric exponents c18rfa014 ................................c18rfa015 13 that vary between 2/3 and 3/4 (e.g., Bauchot & Stephan 1966, 1969; c18rfa041 ................................c18rfa043 14 Hofman 1982; Jerison 1973; Lande 1979; Martin 1981; Martin & Harvey c18rfa053 ................................c18rfa057 15 1985; Stephan 1972). From a paleontological perspective, the body-brain ................................ 16 size relation offers an appealing way to evaluate the cognitive abilities of c18rfa061 ................................c18rfa095 17 fossil taxa, a problem of particular importance for understanding hominid ................................ 18 evolution. ................................ Inset Figure 18.1 about here ................................ XML Typescript c Cambridge University Press – Generated by TechBooks. The Evolution of Thought Page 1219 of 1365 1 A convincing theoretical basis for this general allometric statistical ................................ c18rfa040 2 pattern, however, remains elusive (Deacon 1997; Harvey & Krebs 1990). ................................ 3 Body mass is not a strict determinant of brain size, as species of similar ................................ 4 size can have different brain sizes and cognitive abilities (Pagel & Harvey ................................ c18rfa068 5 1989). In addition, comparative analyses indicate considerable variation ................................ c18rfa068 6 among taxa from general mammalian patterns (Pagel & Harvey 1989). ................................ 7 Hominid brains, for example, are double or more their expected size as ................................ 8 mammals. There are also phylogenetic differences in typical brain-body ................................ 9 size relations within primates that reflect grade shifts in encephalization ................................ c18rfa007 c18rfa008 10 across taxa (Armstrong 1985a, b; Martin & Harvey 1983, Pagel & Harvey ................................ c18rfa059 11 1989). ................................ c18rfa068 12 One reason for the lack of a universal brain-body size correlation ................................ 13 among mammalian species is that factors other than body mass or ................................ 14 metabolism, such as locomotion, diet, predation risk, social structure, and ................................ 15 life history, affect relations between body and brain size (see recent ................................ c18rfa021 16 reviews in de Waal & Tyack 2003; other chapters in this volume). All of ................................ 17 these factors and others may contribute to selective pressures for cognitive ................................ 18 abilities. As such, allometric scaling models developed from analyses of ................................ 19 relations between physical variables such as metabolic rate and body mass ................................ 20 may not be appropriate models for relations between body and brain size. ................................ XML Typescript c Cambridge University Press – Generated by TechBooks. Body Size and Intelligence in Hominoid Evolution Page 1220 of 1365 1 Evolving a large brain depends upon a complex balance of costs and ................................ 2 benefits, which vary from species to species. There are not likely to be ................................ 3 simple explanations based upon simple physical principles. Observed ................................ 4 correlations between brain size and body size, the variability in these ................................ 5 relations, and the reasons underlying phylogenetic differences, require ................................ 6 consideration of both direct and indirect influences. Direct influences ................................ 7 include structural and metabolic constraints on encephalization and ................................ 8 size-related needs: Large-bodied animals are better able to support large ................................ 9 crania and energetically expensive neural tissue than are small-bodied ................................ 10 animals, and larger bodies may require more neurons to control. If this ................................ 11 were all there was to the relation between body size and brain size, then we ................................ 12 would expect simple and consistent statistical associations. However ................................ 13 additional indirect influences can independently affect both body size and ................................ 14 intelligence, including the effects of selective pressures shaping other ................................ 15 aspects of a species’ biology, such as locomotion, diet, predation risk, ................................ 16 social interactions, and life history. If these indirect influences are ................................ 17 important for determining body size-brain size relations, then we expect ................................ 18 more complex and variable statistical patterns, as indicated by analyses of ................................ c18rfa061 19 inter-taxa differences (Martin & Harvey 1985). ................................ 20 Because of the apparent complexities of brain-body relations and ................................ XML Typescript c Cambridge University Press – Generated by TechBooks. The Evolution of Thought Page 1221 of 1365 1 neurobiological differences among extant mammals, simple consideration ................................ 2 of relative brain size provides only an incomplete picture of the cognitive ................................ 3 abilities of fossil species. To understand scaling relations among body size, ................................ 4 brain size, cognition and other aspects of a species’ adaptation, we need to ................................ 5 first understand the underlying selective pressures shaping cognitive ................................ 6 function and related variables. Complex models involving ecological and ................................ 7 social factors are required. Such models may provide new insights into the ................................ 8 causal relations underlying statistical associations between body and brain ................................ 9 size. This chapter first examines the interrelations among multiple relevant ................................ 10 variables and their relations with cognitive capacities and brain size that ................................ 11 apply generally in primates, especially those linked with body size. This ................................ 12 puts us in a stronger position to interpret the cognitive capabilities of ................................ 13 extinct taxa, and therefore to understand the evolution of intelligence in the ................................ 14 hominids. ................................ ................................ A-Head 15 Previous hypotheses relating body mass to intelligence ................................ 16 Logarithmic scaling between brain size and body size in mammals is