The prehistory of handedness: Archaeological data and comparative ethology☆
Introduction
Human hand use patterns can be characterised as complementary role differentiation (CRD). The CRD model of bimanual action derives from the Kinematic Chain model, which was proposed by Guiard (1987) and applied by Uomini (2006a) to the prehistoric activities that offer evidence for handedness (Steele and Uomini, 2009). In this model, one hand executes high frequency tasks (involving finer temporal and spatial resolution) while the other hand performs low frequency tasks (such as supporting an object). Rather than one hand being “dominant,” the CRD model recognises that both hands have different but equally important roles (Corbetta and Thelen, 1996). “Right-handers” are thus defined as people who prefer to adopt the high frequency role with the right hand and the low frequency role with the left hand, as shown by experiments on humans for a complementary bimanual task requiring precision versus support (Hinckley, 1996, Hinckley et al., 1997). A hand role dichotomy appears to emerge between seven and thirteen months of age (Bresson et al., 1977, Ramsay et al., 1979, Michel et al., 1985, Kimmerle et al., 1995, Michel, 1998) and is well established by age three (Ingram, 1975, Gaillard, 1996), yet its genetic determinants are still unknown (Crow, 1998, Van Agtmael et al., 2001).
Beyond the individual, Homo sapiens sapiens displays lateralised hand preference at the species level. This means that a bias to the right-handed CRD pattern is found in all human populations around the world (reviewed in Llaurens et al., 2009), with the frequency of right handed persons in any given population varying between 74% and 96% (Hardyck and Petrinovich, 1977, Porac and Coren, 1981, McManus, 1991, Connolly and Bishop, 1992, Perelle and Ehrman, 1994, Annett, 2002, Raymond and Pontier, 2004, Faurie et al., 2005). There has never been any report of a human population in which left handed individuals predominate (Llaurens et al., 2009). In contrast to the human bias, it is clear from observations of experimental and spontaneous hand actions in captive and wild subjects that the non human primates do not show a species wide consistency in hand use patterns (Colell et al., 1995, Papademetriou et al., 2005). While group level biases can occur in some populations of chimpanzees (e.g., a rightward bias at Yerkes [Hopkins et al., 2007]) and for certain manual actions in some great apes (e.g., gorillas feeding on plants [Corp and Byrne, 2004]), there is no consistent pattern across populations at the species level, since some populations are claimed to show a leftward bias (e.g., for termite fishing in Gombe's wild chimpanzees [Lonsdorf and Hopkins, 2005]).
Furthermore, these biases do not extend to all actions, nor do they reach the extreme degree of consistency seen in humans across all tasks including unimanual actions (McManus, 1985, Hopkins, 2006). As discussed below in more detail, the compilation of research led by various authors suggests that a species-wide group level manual preference across all tasks is not the norm in primates. Therefore, human handedness is unique in both its direction (rightward CRD pattern) and its strength (species wide preference), and remains to be explained in evolutionary terms.
The timing and context for the emergence of species handedness in hominins is therefore of much interest to palaeoanthropologists. Combining palaeoanthropology with primatology can help us decide whether human and nonhuman hand preference patterns are part of the same continuum, or are qualitatively different. A selection of archaeological and primatological findings for hand preference is discussed in a comparative framework, with a special focus on the population- versus species-level distinction.
Section snippets
Lateralised ancestors
Markers of hand preference in prehistory are found in material culture from the actions of lateralised tool manufacture and use that leave traces on objects, and in fossil skeletal asymmetries resulting from asymmetric use of the upper limb muscles over an individual's lifetime. These data have been extensively reviewed by Steele, 2000, Weaver et al., 2001, Steele and Uomini, 2005, Steele and Uomini, 2009, Auerbach and Ruff, 2006, Cashmore et al., 2008, and Uomini, 2008, Uomini, in press.
Comparing apes
In an ideal comparative perspective, the archaeological handedness data available for hominins would be directly compared to excavated material culture from extinct members of the extant great apes Pan, Gorilla, and Pongo. However, archaeological investigations of chimpanzee sites (Mercader et al., 2002, McGrew et al., 2006, Haslam, 2009) are just beginning, and there are virtually no post cranial pongid fossils that would be diagnostic for laterality. The most useful category of evidence
Discussion
As reviewed above, the most robust evidence for population level handedness in prehistory begins with the Neanderthals, although data from East African sites as early as 1 Ma indicate right handedness was also the norm. As in the living human hand preference surveys, there are no reports of predominant left handedness in any prehistoric data sets. These findings contrast sharply with the situation in non human primates, especially great apes, in which we find large geographical variation in
Acknowledgements
I would like to thank Bill McGrew and LCHES for inviting me to participate in the fascinating conference, Palaeoanthropology Meets Primatology 1, that led to this paper. This research was presented jointly with Linda Marchant, to whom I am grateful for valuable advice on previous versions of the manuscript. Susan Antón, John Gowlett, Chris McManus, and anonymous reviewers also offered helpful critiques on a previous draft. The experiments were funded by a grant Land of Legends Lejre (formerly
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2020, Journal of Archaeological Science: ReportsCitation Excerpt :The control of fine manual actions (called “complex” motor actions) is attributed to the left hemisphere which would explain the preferential use of the right hand for this type of action (e.g. Greenfield, 1991; Johnson-Frey et al., 2005; Lewis, 2006; Schluter et al., 2001). Some authors have thus hypothesized the development of a strong manual preference in relation to the manufacture and complexity of tools (Corballis, 1987; Stock et al., 2013; Uomini and Gowlett, 2013; Uomini, 2009). Other studies show a greater effect of the nature and demands of the task as well as the type of object involved and the speed of the action (Forrester et al., 2013, 2012, 2011; Pouydebat et al., 2014; Quaresmini et al., 2014; Rogers, 2009).
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This article is part of the ‘Palaeoanthropology Meets Primatology’ Special Issue.