Hominin reliance on Oldowan stone tools – which appear from 2.5mya and are believed to have been socially transmitted – has been hypothesised to have led to the evolution of teaching and language. Here we present an experiment investigating the efficacy of transmission of Oldowan tool-making skills along chains of adult human participants (N=184) using 5 different transmission mechanisms. Across six measures, transmission improves with teaching, and particularly with language, but not with imitation or emulation. Our results support the hypothesis that hominin reliance on stone tool-making generated selection for teaching and language and imply that (i) low-fidelity social transmission, such as imitation/emulation, may have contributed to the ~700,000 year stasis of the Oldowan technocomplex, and (ii) teaching or proto-language may have been pre-requisites for the appearance of Acheulean technology. This work supports a gradual evolution of language, with simple symbolic communication preceding behavioural modernity by hundreds of thousands of years.

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BackgroundThe popular theory that complex tool-making and language co-evolved in the human lineage rests on the hypothesis that both skills share underlying brain processes and systems. However, language and stone tool-making have so far only been studied separately using a range of neuroimaging techniques and diverse paradigms.Methodology/Principal FindingsWe present the first-ever study of brain activation that directly compares active Acheulean tool-making and language. Using functional transcranial Doppler ultrasonography (fTCD), we measured brain blood flow lateralization patterns (hemodynamics) in subjects who performed two tasks designed to isolate the planning component of Acheulean stone tool-making and cued word generation as a language task. We show highly correlated hemodynamics in the initial 10 seconds of task execution.Conclusions/SignificanceStone tool-making and cued word generation cause common cerebral blood flow lateralization signatures in our participants. This is consistent with a shared neural substrate for prehistoric stone tool-making and language, and is compatible with language evolution theories that posit a co-evolution of language and manual praxis. In turn, our results support the hypothesis that aspects of language might have emerged as early as 1.75 million years ago, with the start of Acheulean technology.

Traditional attempts to understand the evolution of human cognition compare humans with other primates. This research showed that relative brain size covaries with cognitive skills, while adaptations that buffer the developmental and energetic costs of large brains (e.g. allomaternal care), and ecological or social benefits of cognitive abilities, are critical for their evolution. To understand the drivers of cognitive adaptations, it is profitable to consider distant lineages with convergently evolved cognitions. Here, we examine the facilitators of cognitive evolution in corvid birds, where some species display cultural learning, with an emphasis on family life. We propose that extended parenting (protracted parent–offspring association) is pivotal in the evolution of cognition: it combines critical life-history, social and ecological conditions allowing for the development and maintenance of cognitive skillsets that confer fitness benefits to individuals. This novel hypothesis complements the extended childhood idea by considering the parents' role in juvenile development. Using phylogenetic comparative analyses, we show that corvids have larger body sizes, longer development times, extended parenting and larger relative brain sizes than other passerines. Case studies from two corvid species with different ecologies and social systems highlight the critical role of life-history features on juveniles’ cognitive development: extended parenting provides a safe haven, access to tolerant role models, reliable learning opportunities and food, resulting in higher survival. The benefits of extended juvenile learning periods, over evolutionary time, lead to selection for expanded cognitive skillsets. Similarly, in our ancestors, cooperative breeding and increased group sizes facilitated learning and teaching. Our analyses highlight the critical role of life-history, ecological and social factors that underlie both extended parenting and expanded cognitive skillsets. This article is part of the theme issue ‘Life history and learning: how childhood, caregiving and old age shape cognition and culture in humans and other animals’.

In this review and position paper we explore the neural substrates for manual specialization and their possible connection with language and speech. We focus on two contrasting hypotheses of the origins of language and manual specialization: the language-first scenario and the tool-use-first scenario. Each one makes specific predictions about hand-use in non-human primates, as well as about the necessity of an association between speech adaptations and population-level right-handedness in the archaeological and fossil records. The concept of handedness is reformulated for archaeologists in terms of manual role specialization, using Guiard's model of asymmetric bimanual coordination. This focuses our attention on skilled bimanual tasks in which both upper limbs play complementary roles. We review work eliciting non-human primate hand preferences in co-ordinated bimanual tasks, and relevant archaeological data for estimating the presence or absence of a population-level bias to the right hand as the manipulator in extinct hominin species and in the early prehistory of our own species.

Using the comparative approach, researchers draw inferences about the evolution of cognition. Psychologists have postulated several hypotheses to explain why certain species are cognitively more flexible than others, and these hypotheses assume that certain cognitive skills are linked together to create a generally “smart” species. However, empirical findings suggest that several animal species are highly specialized, showing exceptional skills in single cognitive domains while performing poorly in others. Although some cognitive skills may indeed overlap, we cannot a priori assume that they do across species. We argue that the term “cognition” has often been used by applying an anthropocentric viewpoint rather than a biocentric one. As a result, researchers tend to overrate cognitive skills that are human-like and assume that certain skills cluster together in other animals as they do in our own species. In this paper, we emphasize that specific physical and social environments create selection pressures that lead to the evolution of certain cognitive adaptations. Skills such as following the pointing gesture, tool-use, perspective-taking, or the ability to cooperate evolve independently from each other as a concrete result of specific selection pressures, and thus have appeared in distantly related species. Thus, there is not “one cognition”. Our argument is founded upon traditional Darwinian thinking, which—although always at the forefront of biology—has sometimes been neglected in animal cognition research. In accordance with the biocentric approach, we advocate a broader empirical perspective as we are convinced that to better understand animal minds, comparative researchers should focus much more on questions and experiments that are ecologically valid. We should investigate nonhuman cognition for its own sake, not only in comparison to the human model.

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The cumulative modification of tool designs over time is a crucial development for technological evolution. Cognitive-related prerequisites for this technological capability are innovative behaviour and the faithful inter-generational transmission and maintenance of tool designs by accurate social learning processes. Here, we investigated the hypothesis that a complex of morphological and behavioural adaptations specifically for tool skills is also required. In a novel analysis we compared the tool-associated adaptive patterns in Homo erectus and the New Caledonian crow. Both species provide the most convincing early Homo and nonhuman evidence, respectively, for the making of cumulatively modified tools. We identified probable shared traits in H. erectus and the New Caledonian crow that include morphological adaptations specifically for enhanced tool manipulation and a significant component of daily diet from hunting and/or processing animal food with tools. We propose that a tool-using lifestyle based on animal food that confers a reproductive advantage and evolves enhanced tool manipulation skills, together with appropriate innovative ability and social learning processes, may be essential for cumulative modification of tool designs.