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Non-Spurious Correlations Between Genetic and Linguistic Diversities in the Context of Human Evolution

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Non-Spurious Correlations Between Genetic and Linguistic Diversities in the Context of Human Evolution Non-Spurious Correlations between Genetic and Linguistic Diversities in the Context of Human Evolution Dan Dediu Bsc. Math. Comp. Sci., Msc. Neurobiol. Behav. A thesis submitted in fulfillment of requirements for the degree of Doctor of Philosophy to Linguistics and English Language School of Philosophy, Psychology and Language Sciences University of Edinburgh February 2007 © Copyright 2006 by Dan Dediu Declaration I hereby declare that this thesis is of my own composition, and that it contains no material previously submitted for the award of any other degree. The work reported in this thesis has been executed by myself, except where due acknowledgment is made in the text. Dan Dediu iii iv Abstract This thesis concerns human diversity, arguing that it represents not just some form of noise, which must be filtered out in order to reach a deeper explanatory level, but the engine of human and language evolution, metaphorically put, the best gift Nature has made to us. This diversity must be understood in the context of (and must shape) human evolution, of which the Recent Out-of-Africa with Replacement model (ROA) is currently regarded, especially outside palaeoanthropology, as a true theory. It is argued, using data from palaeoanthropology, human population genetics, ancient DNA studies and primatology, that this model must be, at least, amended, and most probably, rejected, and its alternatives must be based on the concept of reticulation. The relationships between the genetic and linguistic diversities is complex, including inter- individual genetic and behavioural differences (behaviour genetics) and inter-population differences due to common demographic, geographic and historic factors (spurious correlations), used to study (pre)historical processes. It is proposed that there also exist non- spurious correlations between genetic and linguistic diversities, due to genetic variants which can bias the process of language change, so that the probabilities of alternative linguistic states are altered. The particular hypothesis (formulated with Prof. D. R. Ladd) of a causal relationship between two human genes and one linguistic typological feature is supported by the statistical analysis of a vast database of 983 genetic variants and 26 linguistic features in 49 Old World populations, controlling for geography and known linguistic history. The general theory of non-spurious correlations between genetic and linguistic diversities is developed and its consequences and predictions analyzed. It will very probably profoundly impact our understanding of human diversity and will offer a firm footing for theories of language evolution and change. More specifically, through such a mechanism, gradual, accretionary models of language evolution are a natural consequence of post-ROA human evolutionary models. The unravellings of causal effects of inter-population genetic differences on linguistic states, mediated by complex processes of cultural evolution (biased iterated learning), will represent a major advance in our understanding of the relationship between cultural and genetic diversities, and will allow a better appreciation of this most fundamental and supremely valuable characteristic of humanity – its intrinsic diversity. v vi Acknowledgments Many thanks to my supervisors, Jim Hurford and Simon Kirby, for encouragement and many critical and insightful discussions, even when I was attacking controversial issues (which I almost always did). To D. Robert Ladd for sharing his vast knowledge of linguistics and willingness to work hard on the “crazy” idea of non-spurious correlations, and for his continuous support. To Mónica Tamariz, Stefan Höfler and Dave Hawkey for discussions on linguistic diversity, its meanings and origins, to Anna Parker for help with getting this thesis right and all the other LEC members for sharing ideas and thought-provoking discussions. To Carmen Strungaru and Wulf Schiefenhövel for making me discover language evolution. Special thanks to my wife, Alexandra Dima, for her loving and funny way of being, and for her invaluable help with statistics, and her decisive role in my actually learning it. I would not have been here without her... To my parents, who made me like so many aspects of Nature, from biology to mathematics, teaching me how to think freely (especially in a time and place when this was forbidden), and helping me pursue my dreams, despite many hardships. The Overseas Research Students Awards Scheme (ORS Award 2003014001) and the College of Humanities and Social Science, University of Edinburgh's Studentship made this financially possible. I thank B. Connell, C. Kutsch Lojenga, H. Eaton, J. A. Edmondson, J. Hurford, K. Bostoen, L. Ziwo, M. Blackings, N. Fabb, O. Stegen, R. Asher, R. Ridouane, M. Endl and J. Roberts for help with language data and R. McMahon for comments on an early draft of the tone-genes correlation idea. vii viii Table of Contents Declaration............................................................................................................................. iii Abstract................................................................................................................................... v Acknowledgments................................................................................................................. vii 1. Introduction and overview................................................................................................... 1 2. Human evolution................................................................................................................. 3 2.1. The Recent Out-of-Africa Model and the Evolutionary History of Homo sapiens...... 4 2.1.1. The historical development of ROA.................................................................... 4 2.1.1.1. Mitochondrial DNA.................................................................................... 7 2.1.1.2. The Molecular Clock................................................................................. 10 2.1.1.3. How neutral really is the human mtDNA?................................................. 12 2.1.1.4. Review of critiques of Cann, Stoneking & Wilson (1987)........................ 13 2.1.1.5. Reaching the current view on modern human mtDNA.............................. 14 2.1.1.6. ROA in its current form............................................................................. 15 2.1.2. Before the moderns: the palaeoanthropological context.................................... 16 2.1.2.1. What are species?...................................................................................... 22 2.1.2.2. Homo erectus and their feats..................................................................... 26 2.1.2.3. Homo neanderthalensis............................................................................. 30 2.1.3. The evolution of modern humans: the competing models................................. 33 2.2. Problems and issues for ROA .................................................................................. 43 2.2.1. The transition to Homo sapiens was not a “revolution”..................................... 43 2.2.2. A structured population for the origins of Homo sapiens.................................. 47 2.2.3. Genes with deep, non-African branches............................................................ 48 2.2.4. Primate models and the speciosity of Homo...................................................... 52 2.2.5. Regional morphological continuity................................................................... 56 2.2.5.1. The Abrigo do Lagar Velho child.............................................................. 60 2.2.6. Global trends..................................................................................................... 61 2.2.7. Ancient DNA..................................................................................................... 63 2.2.8. The genetic structure of living populations........................................................ 68 2.2.8.1. The apportionment of genetic diversity in living humans and its interpretations......................................................................................................... 70 2.2.8.2. The evolutionary interpretations of modern human genetic diversity........ 74 2.2.9. The unexpected diversity of the genus Homo: the Flores man.......................... 78 2.3. Putting all together: what is the most plausible class of human evolutionary models? ......................................................................................................................................... 80 2.3.1. John Relethford's “Mostly Out Of Africa”........................................................ 81 2.3.2. Alan Templeton's “Out of Africa again and again”........................................... 83 2.4. Conclusions ............................................................................................................. 86 3. Language-genes correlations............................................................................................. 89 3.1. The correlations between the capacity for language and the genetic makeup............ 90 3.1.1. Methods............................................................................................................. 90 3.1.2. Measuring the effect of genes and environment: the heritability....................... 94 3.1.3. Heritability estimates for speech and language...............................................
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