Historically, population-level handedness has been considered a hallmark of human evolution. Whether nonhuman primates exhibit population-level handedness remains a topic of considerable debate. This paper summarizes published data on handedness in great apes. Comparative analysis indicated that chimpanzees and bonobos show population-level right handedness, whereas gorillas and orangutans do not. All ape species showed evidence of population-level handedness when considering specific tasks. Familial analyses in chimpanzees indicated that offspring and maternal (but not paternal) handedness was significantly positively correlated, but this finding was contingent upon the classification criteria used to evaluate hand preference. Overall, the proportion of right handedness is lower in great apes compared with humans, and various methodological and theoretical explanations for this discrepancy are discussed.
Keywords handedness; great apes; laterality; behavior geneticsApproximately 85% to 90% of humans report themselves as being right handed (Annett, 1985(Annett, ,2002Perelle & Ehrman, 1994;Porac & Coren, 1981;Raymond & Pontier, 2004). Right handedness presumably reflects a left-hemisphere specialization for motor skills and is thought to be an indirect marker of lateralization for language functions. For example, 96% of righthanded individuals are left-hemisphere dominant for language in contrast to about 70% in lefthanded individuals (Knecht et al., 2000;Rasmussen & Milner, 1977). The association between hand preference and language dominance has led to a number of evolutionary and biological theories proposing that hemispheric specialization and language (or perhaps other higher cognitive functions) coevolved and are unique characteristics of the human brain (see Annett, 1985;Corballis, 1992Corballis, ,2002Crow, 1998;Yeo, Thoma, & Gangestad, 2002, for reviews). The historical lack of evidence for functional asymmetries in nonhuman species, with the exception of birds, was taken as evidence in support of the uniqueness of hemispheric specialization to humans (Bradshaw & Rogers, 1993). However, based on research carried out in the past 10 to 15 years, good evidence exists of population-level hemispheric specialization in nonhuman species for various cognitive, perceptual, and to a lesser extent, motor functions (Rogers & Andrew, 2002;Vallortigara & Rogers, 2005). For example, toads prefer to the use their right paw when removing materials from their bodies and orienting themselves when floating upside-down (Bisazza, Cantalupo, Robins, Rogers, & Vallortigara, 1996). Chicks and pigeons show a host of behavioral asymmetries including left-hemisphere biases in local discrimination of food items (Andrew, 1991;Güntürkün, 1997). Chicks also show and right-hemisphere biases in predator detection (Vallortigara, 1992), as has also been reported in certain species of fish (Bisazza, Cantalupo, Capocchiano, & Vallortigara, 2000). These data clearly challenge the long held belief that hemispheric specialization is a un...