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The size of a bird's nest can play a key role in ensuring reproductive success and is determined by a variety of factors. The primary function of the nest is to protect offspring from the environment and predators. Field studies in a number of passerine species have indicated that higher-latitude populations in colder habitats build larger nests with thicker walls compared to lower-latitude populations, but that these larger nests are more vulnerable to predation. Increases in nest size can also be driven by sexual selection, as nest size can act as a signal of parental quality and prompt differential investment in other aspects of care. It is unknown, however, how these microevolutionary patterns translate to a macroevolutionary scale. Here, we investigate potential drivers of variation in the outer and inner volume of open cup nests using a large dataset of nest measurements from 1117 species of passerines breeding in a diverse range of environments. Our dataset is sourced primarily from the nest specimens at the Natural History Museum (UK), complemented with information from ornithological handbooks and online databases. We use phylogenetic comparative methods to test long-standing hypotheses about potential macroevolutionary correlates of nest size, namely nest location, clutch size and variables relating to parental care, together with environmental and geographical factors such as temperature, rainfall, latitude and insularity. After controlling for phylogeny and parental body size, we demonstrate that the outer volume of the nest is greater in colder climates, in island-dwelling species and in species that nest on cliffs or rocks. By contrast, the inner cup volume is associated solely with average clutch size, increasing with the number of chicks raised in the nest. We do not find evidence that nest size is related to the length of parental care for nestlings. Our study reveals that the average temperature in the breeding range, along with several key life-history traits and proxies of predation threat, shapes the global interspecific variation in passerine cup nest size. We also showcase the utility of museum nest collections-a historically underused resource-for large-scale studies of trait evolution., (© 2022 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Callitrichidae is a unique primate family not only in terms of the large number of food transfers to infants but also for the prevalence of transfers that are initiated by the adults. It has been hypothesized that, as well as provisioning infants, callitrichid food transfers might function to teach the receiver what food types to eat. If food provisioning has a teaching function, we would expect successful food transfers to be more likely with food types that are novel to the juveniles. We would also expect juveniles to learn about foods from those transfers. We introduced different types of food (some familiar, some novel) to wild groups of golden lion tamarins (Leontopithecus rosalia). While novel foods were not more successfully transferred than familiar food in the experiment, transfers were more successful (i.e., the receiver obtained food) when the donor had previous experience with that food. Moreover, we found evidence suggesting that food transfers influenced the future foraging choices of juveniles. Our findings are consistent with the first and third criteria of the functional definition of teaching, which requires that tutors (the adults) modify their behavior in the presence of a naïve individual (a juvenile), and that the naïve individual learns from the modified behavior of the demonstrator. Our findings are also consistent with the provisioning function of food transfer. Social learning seems to play an important role in the development of young tamarins' foraging preferences.

Observational learning is a form of social learning in which a demonstrator performs a target task in the company of an observer, who may as a consequence learn something about it. In this study, we approach social learning in terms of the dynamics of coordination rather than the more common perspective of transmission of information. We hypothesised that observers must continuously adjust their visual attention relative to the demonstrator's time-evolving behaviour to benefit from it. We eye-tracked observers repeatedly watching videos showing a demonstrator solving one of three manipulative puzzles before attempting at the task. The presence of the demonstrator's face and the availability of his verbal instruction in the videos were manipulated. We then used recurrence quantification analysis to measure the dynamics of coordination between the overt attention of the observers and the demonstrator's manipulative actions. Bayesian hierarchical logistic regression was applied to examine (1) whether the observers' performance was predicted by such indexes of coordination, (2) how performance changed as they accumulated experience, and (3) if the availability of speech and intentional gaze of the demonstrator mediated it. Results showed that learners better able to coordinate their eye movements with the manipulative actions of the demonstrator had an increasingly higher probability of success in solving the task. The availability of speech was beneficial to learning, whereas the presence of the demonstrator's face was not. We argue that focusing on the dynamics of coordination between individuals may greatly improve understanding of the cognitive processes underlying social learning., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier B.V. All rights reserved.)

Recognition plays an important role in the formation and organization of animal groups. Many animals are capable of class-level recognition, discriminating, for example, on the basis of species, kinship or familiarity. Individual recognition requires that animals recognize distinct cues, and learn to associate these with the specific individual from which they are derived. In this study, we asked whether sticklebacks ( Gasterosteus aculeatus and Pungitius pungitius ) were capable of learning to recognize individual conspecifics. We have used these fish as model organisms for studying selective social learning, and demonstrating a capacity for individual recognition in these species would provide an exciting opportunity for studying how biases for copying specific individuals shape the dynamics of information transmission. To test for individual recognition, we trained subjects to associate green illumination with the provision of a food reward close to one of two conspecifics, and, for comparison, one of two physical landmarks. Both species were capable of recognizing the rewarded landmark, but neither showed a preference for associating with the rewarded conspecific. Our study provides no evidence for individual recognition in either species. We speculate that the fission-fusion structure of their social groups may not favour a capacity for individual recognition., Competing Interests: The authors declare no competing interests., (© 2020 The Authors.)

Human technology is characterized by cumulative cultural knowledge gain, yet researchers have limited knowledge of the mix of copying and innovation that maximizes progress. Here, we analyze a unique large-scale dataset originating from collaborative online programming competitions to investigate, in a setting of real-world complexity, how individual differences in innovation, social-information use, and performance generate technological progress. We find that cumulative knowledge gain is primarily driven by pragmatists, willing to copy, innovate, explore, and take risks flexibly, rather than by pure innovators or habitual copiers. Our study also reveals a key role for prestige in information transfer., (Copyright © 2020 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works. Distributed under a Creative Commons Attribution License 4.0 (CC BY).)

As a form of adaptive plasticity that allows organisms to shift their phenotype toward the optimum, learning is inherently a source of developmental bias. Learning may be of particular significance to the evolutionary biology community because it allows animals to generate adaptively biased novel behavior tuned to the environment and, through social learning, to propagate behavioral traits to other individuals, also in an adaptively biased manner. We describe several types of developmental bias manifest in learning, including an adaptive bias, historical bias, origination bias, and transmission bias, stressing that these can influence evolutionary dynamics through generating nonrandom phenotypic variation and/or nonrandom environmental states. Theoretical models and empirical data have established that learning can impose direction on adaptive evolution, affect evolutionary rates (both speeding up and slowing down responses to selection under different conditions) and outcomes, influence the probability of populations reaching global optimum, and affect evolvability. Learning is characterized by highly specific, path-dependent interactions with the (social and physical) environment, often resulting in new phenotypic outcomes. Consequently, learning regularly introduces novelty into phenotype space. These considerations imply that learning may commonly generate plasticity first evolution., (© 2019 Wiley Periodicals, Inc.)

This article consists of commentaries on a selected group of papers of Marc Feldman published in Theoretical Population Biology from 1970 to the present. The papers describe a diverse set of population-genetic models, covering topics such as cultural evolution, social evolution, and the evolution of recombination. The commentaries highlight Marc Feldman's role in providing mathematically rigorous formulations to explore qualitative hypotheses, in many cases generating surprising conclusions., (Copyright © 2018 Elsevier Inc. All rights reserved.)

Culture (behaviour based on socially transmitted information) is present in diverse animal species, yet how it interacts with genetic evolution remains largely unexplored. Here, we review the evidence for gene-culture coevolution in animals, especially birds, cetaceans and primates. We describe how culture can relax or intensify selection under different circumstances, create new selection pressures by changing ecology or behaviour, and favour adaptations, including in other species. Finally, we illustrate how, through culturally mediated migration and assortative mating, culture can shape population genetic structure and diversity. This evidence suggests strongly that animal culture plays an important evolutionary role, and we encourage explicit analyses of gene-culture coevolution in nature.

Whether learning primarily reflects general processes or species-specific challenges is a long-standing matter of dispute. Here, we present a comprehensive analysis of public information use (PI-use) in sticklebacks (Gasterosteidae). PI-use is a form of social learning by which animals are able to assess the relative quality of resources, here prey patches, by observing the behaviour of others. PI-use was highly specific with only Pungitius and their closest relative Culaea inconstans showing evidence of PI-use. We saw no effects of ontogenetic experience upon PI-use in Pungitius pungitius . Experiments with live demonstrators and animated fish revealed that heightened activity and feeding strikes by foraging conspecifics are important cues in the transmission of PI. Finally, PI-use was the only form of learning in which P. pungitius and another stickleback, Gasterosteus aculeatus differed. PI-use in sticklebacks is species-specific and may represent an 'ecological specialization' for social foraging. Whether this reflects selection on perception, attentional or cognitive processes remains to be determined., Competing Interests: We have no competing interests to declare.

Why groups of individuals sometimes exhibit collective 'wisdom' and other times maladaptive 'herding' is an enduring conundrum. Here we show that this apparent conflict is regulated by the social learning strategies deployed. We examined the patterns of human social learning through an interactive online experiment with 699 participants, varying both task uncertainty and group size, then used hierarchical Bayesian model fitting to identify the individual learning strategies exhibited by participants. Challenging tasks elicit greater conformity among individuals, with rates of copying increasing with group size, leading to high probabilities of herding among large groups confronted with uncertainty. Conversely, the reduced social learning of small groups, and the greater probability that social information would be accurate for less-challenging tasks, generated 'wisdom of the crowd' effects in other circumstances. Our model-based approach provides evidence that the likelihood of collective intelligence versus herding can be predicted, resolving a long-standing puzzle in the literature.

Non-human primate neuroimaging is a rapidly growing area of research that promises to transform and scale translational and cross-species comparative neuroscience. Unfortunately, the technological and methodological advances of the past two decades have outpaced the accrual of data, which is particularly challenging given the relatively few centers that have the necessary facilities and capabilities. The PRIMatE Data Exchange (PRIME-DE) addresses this challenge by aggregating independently acquired non-human primate magnetic resonance imaging (MRI) datasets and openly sharing them via the International Neuroimaging Data-sharing Initiative (INDI). Here, we present the rationale, design, and procedures for the PRIME-DE consortium, as well as the initial release, consisting of 25 independent data collections aggregated across 22 sites (total = 217 non-human primates). We also outline the unique pitfalls and challenges that should be considered in the analysis of non-human primate MRI datasets, including providing automated quality assessment of the contributed datasets., (Copyright © 2018 The Authors. Published by Elsevier Inc. All rights reserved.)

Human children, in contrast to other species, are frequently cast as prolific "over-imitators". However, previous studies of "over-imitation" have overlooked many important real-world social dynamics, and may thus provide an inaccurate account of this seemingly puzzling and potentially maladaptive phenomenon. Here we investigate this topic using a cultural evolutionary approach, focusing particularly on the key adaptive learning strategy of majority-biased copying. Most "over-imitation" research has been conducted using consistent demonstrations to the observer, but we systematically varied the frequency of demonstrators that 4- to 6-year-old children observed performing a causally irrelevant action. Children who "over-imitate" inflexibly should copy the majority regardless of whether the majority solution omits or includes a causally irrelevant action. However, we found that children calibrated their tendency to acquire the majority behavior, such that copying did not extend to majorities that performed irrelevant actions. These results are consistent with a highly functional, adaptive integration of social and causal information, rather than explanations implying unselective copying or causal misunderstanding. This suggests that our species might be better characterized as broadly "optimal-" rather than "over-" imitators., (© 2017 John Wiley & Sons Ltd.)

Phenotypic variation is generated by the processes of development, with some variants arising more readily than others-a phenomenon known as "developmental bias." Developmental bias and natural selection have often been portrayed as alternative explanations, but this is a false dichotomy: developmental bias can evolve through natural selection, and bias and selection jointly influence phenotypic evolution. Here, we briefly review the evidence for developmental bias and illustrate how it is studied empirically. We describe recent theory on regulatory networks that explains why the influence of genetic and environmental perturbation on phenotypes is typically not uniform, and may even be biased toward adaptive phenotypic variation. We show how bias produced by developmental processes constitutes an evolving property able to impose direction on adaptive evolution and influence patterns of taxonomic and phenotypic diversity. Taking these considerations together, we argue that it is not sufficient to accommodate developmental bias into evolutionary theory merely as a constraint on evolutionary adaptation. The influence of natural selection in shaping developmental bias, and conversely, the influence of developmental bias in shaping subsequent opportunities for adaptation, requires mechanistic models of development to be expanded and incorporated into evolutionary theory. A regulatory network perspective on phenotypic evolution thus helps to integrate the generation of phenotypic variation with natural selection, leaving evolutionary biology better placed to explain how organisms adapt and diversify., (Copyright © 2018 by the Genetics Society of America.)

While social learning is widespread, indiscriminate copying of others is rarely beneficial. Theory suggests that individuals should be selective in what, when, and whom they copy, by following 'social learning strategies' (SLSs). The SLS concept has stimulated extensive experimental work, integrated theory, and empirical findings, and created impetus to the social learning and cultural evolution fields. However, the SLS concept needs updating to accommodate recent findings that individuals switch between strategies flexibly, that multiple strategies are deployed simultaneously, and that there is no one-to-one correspondence between psychological heuristics deployed and resulting population-level patterns. The field would also benefit from the simultaneous study of mechanism and function. SLSs provide a useful vehicle for bridge-building between cognitive psychology, neuroscience, and evolutionary biology., (Copyright © 2018. Published by Elsevier Ltd.)

The ability to build progressively on the achievements of earlier generations is central to human uniqueness, but experimental investigations of this cumulative cultural evolution lack real-world complexity. Here, we studied the dynamics of cumulative culture using a large-scale data set from online collaborative programming competitions run over 14 years. We show that, within each contest population, performance increases over time through frequent 'tweaks' of the current best entry and rare innovative 'leaps' (successful tweak:leap ratio = 16:1), the latter associated with substantially greater variance in performance. Cumulative cultural evolution reduces technological diversity over time, as populations focus on refining high-performance solutions. While individual entries borrow from few sources, iterative copying allows populations to integrate ideas from many sources, demonstrating a new form of collective intelligence. Our results imply that maximising technological progress requires accepting high levels of failure.

Women appear to copy other women's preferences for men's faces. This 'mate-choice copying' is often taken as evidence of psychological adaptations for processing social information related to mate choice, for which facial information is assumed to be particularly salient. No experiment, however, has directly investigated whether women preferentially copy each other's face preferences more than other preferences. Further, because prior experimental studies used artificial social information, the effect of real social information on attractiveness preferences is unknown. We collected attractiveness ratings of pictures of men's faces, men's hands, and abstract art given by heterosexual women, before and after they saw genuine social information gathered in real time from their peers. Ratings of faces were influenced by social information, but no more or less than were images of hands and abstract art. Our results suggest that evidence for domain-specific social learning mechanisms in humans is weaker than previously suggested.

Many animals emit calls in the presence of food, but researchers do not always know the function of these calls. Evidence suggests that adult golden lion tamarins ( Leontopithecus rosalia ) use food-offering calls to teach juveniles which substrate (i.e., microhabitat) to forage on, or in, for food. However, we do not yet know whether juveniles learn from this aspect of the adults' behavior. Here we examine whether juveniles learn to associate food-offering calls with a foraging substrate, as a step toward assessing whether these calls qualify as teaching behavior. We compared the performance of four wild juvenile golden lion tamarins that were introduced to a novel substrate while exposed to playbacks of food-offering calls (experimental condition) to the performance of three juveniles that were exposed to the novel substrate without the presence of food-offering playbacks (control condition). We varied the location of the novel substrate between trials. We found that food-offering calls had an immediate effect on juveniles' interactions with the novel substrate, whether they inserted their hands into the substrate and their eating behavior, and a long-term effect on eating behavior at the substrate. The findings imply that juvenile golden lion tamarins can learn through food-offering calls about the availability of food at a substrate, which is consistent with (but does not prove) teaching in golden lion tamarins through stimulus enhancement. Our findings support the hypothesis that teaching might be more likely to evolve in cooperatively breeding species with complex ecological niches.

Since the publication of the primate brain volumetric dataset of Stephan and colleagues in the early 1980s, no major new comparative datasets covering multiple brain regions and a large number of primate species have become available. However, technological and other advances in the last two decades, particularly magnetic resonance imaging (MRI) and the creation of institutions devoted to the collection and preservation of rare brain specimens, provide opportunities to rectify this situation. Here, we present a new dataset including brain region volumetric measurements of 39 species, including 20 species not previously available in the literature, with measurements of 16 brain areas. These volumes were extracted from MRI of 46 brains of 38 species from the Netherlands Institute of Neuroscience Primate Brain Bank, scanned at high resolution with a 9.4-T scanner, plus a further 7 donated MRI of 4 primate species. Partial measurements were made on an additional 8 brains of 5 species. We make the dataset and MRI scans available online in the hope that they will be of value to researchers conducting comparative studies of primate evolution., (© 2018 S. Karger AG, Basel.)

Lack of confidence in one's own ability can increase the likelihood of relying on social information. Sex differences in confidence have been extensively investigated in cognitive tasks, but implications for conformity have not been directly tested. Here, we tested the hypothesis that, in a task that shows sex differences in confidence, an indirect effect of sex on social information use will also be evident. Participants (N = 168) were administered a mental rotation (MR) task or a letter transformation (LT) task. After providing an answer, participants reported their confidence before seeing the responses of demonstrators and being allowed to change their initial answer. In the MR, but not the LT, task, women showed lower levels of confidence than men, and confidence mediated an indirect effect of sex on the likelihood of switching answers. These results provide novel, experimental evidence that confidence is a general explanatory mechanism underpinning susceptibility to social influences. Our results have implications for the interpretation of the wider literature on sex differences in conformity., (© 2016 The British Psychological Society.)

Explanations for primate brain expansion and the evolution of human cognition and culture remain contentious despite extensive research. While multiple comparative analyses have investigated variation in brain size across primate species, very few have addressed why primates vary in how much they use social learning. Here, we evaluate the hypothesis that the enhanced reliance on socially transmitted behavior observed in some primates has coevolved with enlarged brains, complex sociality, and extended lifespans. Using recently developed phylogenetic comparative methods we show that, across primate species, a measure of social learning proclivity increases with absolute and relative brain volume, longevity (specifically reproductive lifespan), and social group size, correcting for research effort. We also confirm relationships of absolute and relative brain volume with longevity (both juvenile period and reproductive lifespan) and social group size, although longevity is generally the stronger predictor. Relationships between social learning, brain volume, and longevity remain when controlling for maternal investment and are therefore not simply explained as a by-product of the generally slower life history expected for larger brained species. Our findings suggest that both brain expansion and high reliance on culturally transmitted behavior coevolved with sociality and extended lifespan in primates. This coevolution is consistent with the hypothesis that the evolution of large brains, sociality, and long lifespans has promoted reliance on culture, with reliance on culture in turn driving further increases in brain volume, cognitive abilities, and lifespans in some primate lineages., Competing Interests: The authors declare no conflict of interest.

Competing Interests: The authors declare no conflict of interest.

Gupta et al., in their article in this issue ('Niche construction in evolutionary theory: the construction of an academic niche?'. doi:10.1007/s12041-017-0787-6), lament 'serious problems with the way science is being done' and suggest that 'niche construction theory exemplifies this state of affairs.' However, their aggressively confrontational but superficial critique of niche construction theory (NCT) only contributes to these problems by attacking claims that NCT does not make. This is unfortunate, as their poor scholarship has done a disservice to the evolutionary biology community through propagating misinformation.We correct Gupta et al.'s misunderstandings, stressing that NCT does not suggest that the fact that organisms engage in niche construction is neglected, nor does it make strong claims on the basis of its formal theory. Moreover, the treatment of niche construction as an evolutionary process has been highly productive, and is both theoretically and empirically well-validated.We end by reflecting on the potentially deleterious implications of their publication for evolutionary science.

Many factors, including the demonstrator's sex, status, and familiarity, shape the nature and magnitude of social learning. Given the important role of pair bonds in socially-monogamous animals, we predicted that these intimate relationships would promote the use of social information, and tested this hypothesis in zebra finches (Taeniopygia guttata). Observer birds witnessed either their mate or another familiar, opposite-sex bird eat from one, but not a second novel food source, before being allowed to feed from both food sources themselves. Birds used social information to make foraging decisions, but not all individuals used this information in the same way. While most individuals copied the foraging choice of the demonstrator as predicted, paired males did not, instead avoiding the feeder demonstrated by their mate. Our findings reveal that sex and pairing status interact to influence the use of social information and suggest that paired males might use social information to avoid competing with their mate., (Copyright © 2016 The Authors. Published by Elsevier B.V. All rights reserved.)

November 7-9, 2016 witnessed a joint discussion meeting of the Royal Society and the British Academy (the UK national academies for the sciences and social sciences, respectively) entitled 'New Trends in Evolutionary Biology: Biological, Philosophical and Social Science Perspectives'. The meeting, anticipated with a mix of feverish enthusiasm and dread, sold out months in advance, the eager audience perhaps expecting radical and traditional evolutionists to go toe to toe, rather than the constructive dialogue among biologists, social scientists, and researchers in the humanities that the academies advertised. One issue under discussion was whether or not the explanatory core of evolutionary biology requires updating in the light on recent advances in evo-devo, epigenetics, ecosystem ecology, and elsewhere., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

There is a tension between the conception of cognition as a central nervous system (CNS) process and a view of cognition as extending towards the body or the contiguous environment. The centralised conception requires large or complex nervous systems to cope with complex environments. Conversely, the extended conception involves the outsourcing of information processing to the body or environment, thus making fewer demands on the processing power of the CNS. The evolution of extended cognition should be particularly favoured among small, generalist predators such as spiders, and here, we review the literature to evaluate the fit of empirical data with these contrasting models of cognition. Spiders do not seem to be cognitively limited, displaying a large diversity of learning processes, from habituation to contextual learning, including a sense of numerosity. To tease apart the central from the extended cognition, we apply the mutual manipulability criterion, testing the existence of reciprocal causal links between the putative elements of the system. We conclude that the web threads and configurations are integral parts of the cognitive systems. The extension of cognition to the web helps to explain some puzzling features of spider behaviour and seems to promote evolvability within the group, enhancing innovation through cognitive connectivity to variable habitat features. Graded changes in relative brain size could also be explained by outsourcing information processing to environmental features. More generally, niche-constructed structures emerge as prime candidates for extending animal cognition, generating the selective pressures that help to shape the evolving cognitive system.

Access to information is a key advantage of grouping. Although experienced animals can lead others to solve problems, less is known about whether partially informed individuals can pool experiences to overcome challenges collectively. Here we provide evidence of such 'experience-pooling'. We presented shoals of sticklebacks (Gasterosteus aculeatus) with a two-stage foraging task requiring them to find and access hidden food. Individual fish were either inexperienced or had knowledge of just one of the stages. Shoals containing individuals trained in each of the stages pooled their expertise, allowing more fish to access the food, and to do so more rapidly, compared with other shoal compositions. Strong social effects were identified: the presence of experienced individuals increased the likelihood of untrained fish completing each stage. These findings demonstrate that animal groups can integrate individual experience to solve multi-stage problems, and have implications for our understanding of social foraging, migration and social systems.

Background: Advanced cognitive abilities are widely thought to underpin cultural traditions and cumulative cultural change. In contrast, recent simulation models have found that basic social influences on learning suffice to support both cultural phenomena. In the present study we test the predictions of these models in the context of skill learning, in a model with stochastic demographics, variable group sizes, and evolved parameter values, exploring the cultural ramifications of three different social learning mechanisms., Results: Our results show that that simple forms of social learning such as local enhancement, can generate traditional differences in the context of skill learning. In contrast, we find cumulative cultural change is supported by observational learning, but not local or stimulus enhancement, which supports the idea that advanced cognitive abilities are important for generating this cultural phenomenon in the context of skill learning., Conclusions: Our results help to explain the observation that animal cultures are widespread, but cumulative cultural change might be rare.

Innovative behaviour in animals, ranging from invertebrates to humans, is increasingly recognized as an important topic for investigation by behavioural researchers. However, what constitutes an innovation remains controversial, and difficult to quantify. Drawing on a broad definition whereby any behaviour with a new component to it is an innovation, we propose a quantitative measure, which we call the magnitude of innovation , to describe the extent to which an innovative behaviour is novel. This allows us to distinguish between innovations that are a slight change to existing behaviours (low magnitude), and innovations that are substantially different (high magnitude). Using mathematical modelling and evolutionary computer simulations, we explored how aspects of social interaction, cognition and natural selection affect the frequency and magnitude of innovation. We show that high-magnitude innovations are likely to arise regularly even if the frequency of innovation is low, as long as this frequency is relatively constant, and that the selectivity of social learning and the existence of social rewards, such as prestige and royalties, are crucial for innovative behaviour to evolve. We suggest that consideration of the magnitude of innovation may prove a useful tool in the study of the evolution of cognition and of culture., (© 2017 The Author(s).)

I introduce seven criteria for determining the validity of competing theories for the original function of language. I go on to present a novel explanation that meets all the criteria: language originally evolved to teach kin. I suggest that the use of symbols subsequently generated evolutionary feedback at two levels, in the form of self-modified selection pressures that favored structures in the mind that functioned to manipulate and use symbols with efficiency, and cultural selection on languages for learnability.

Humans exhibit extensive large-scale cooperation, of a form unprecedented in the natural world. Here I suggest that this cooperation arises in our species alone because of our uniquely potent capacities for social learning, imitation and teaching, combined with the co-evolutionary feedbacks that these capabilities have generated on the human mind. Culture took human populations down evolutionary pathways not available to non-cultural species, either by creating conditions that promoted established cooperative mechanisms, such as indirect reciprocity and mutualism, or by generating novel cooperative mechanisms not seen in other taxa, such as cultural group selection. In the process, gene-culture co-evolution seemingly generated an evolved psychology, comprising an enhanced ability and motivation to learn, teach, communicate through language, imitate and emulate, as well as predispositions to docility, social tolerance, and the sharing of goals, intentions and attention. This evolved psychology is entirely different from that observed in any other animal, or that could have evolved through conventional selection on genes alone.

Background: Social learning is potentially advantageous, but evolutionary theory predicts that (i) its benefits may be self-limiting because social learning can lead to information parasitism, and (ii) these limitations can be mitigated via forms of selective copying. However, these findings arise from a functional approach in which learning mechanisms are not specified, and which assumes that social learning avoids the costs of asocial learning but does not produce information about the environment. Whether these findings generalize to all kinds of social learning remains to be established. Using a detailed multi-scale evolutionary model, we investigate the payoffs and information production processes of specific social learning mechanisms (including local enhancement, stimulus enhancement and observational learning) and their evolutionary consequences in the context of skill learning in foraging groups., Results: We find that local enhancement does not benefit foraging success, but could evolve as a side-effect of grouping. In contrast, stimulus enhancement and observational learning can be beneficial across a wide range of environmental conditions because they generate opportunities for new learning outcomes., Conclusions: In contrast to much existing theory, we find that the functional outcomes of social learning are mechanism specific. Social learning nearly always produces information about the environment, and does not always avoid the costs of asocial learning or support information parasitism. Our study supports work emphasizing the value of incorporating mechanistic detail in functional analyses.

In birds and primates, the frequency of behavioural innovation has been shown to covary with absolute and relative brain size, leading to the suggestion that large brains allow animals to innovate, and/or that selection for innovativeness, together with social learning, may have driven brain enlargement. We examined the relationship between primate brain size and both technical (i.e. tool using) and non-technical innovation, deploying a combination of phylogenetically informed regression and exploratory causal graph analyses. Regression analyses revealed that absolute and relative brain size correlated positively with technical innovation, and exhibited consistently weaker, but still positive, relationships with non-technical innovation. These findings mirror similar results in birds. Our exploratory causal graph analyses suggested that technical innovation shares strong direct relationships with brain size, body size, social learning rate and social group size, whereas non-technical innovation did not exhibit a direct relationship with brain size. Nonetheless, non-technical innovation was linked to brain size indirectly via diet and life-history variables. Our findings support 'technical intelligence' hypotheses in linking technical innovation to encephalization in the restricted set of primate lineages where technical innovation has been reported. Our findings also provide support for a broad co-evolving complex of brain, behaviour, life-history, social and dietary variables, providing secondary support for social and ecological intelligence hypotheses. The ability to gain access to difficult-to-extract, but potentially nutrient-rich, resources through tool use may have conferred on some primates adaptive advantages, leading to selection for brain circuitry that underlies technical proficiency., (© 2016 The Author(s).)

Scientific activities take place within the structured sets of ideas and assumptions that define a field and its practices. The conceptual framework of evolutionary biology emerged with the Modern Synthesis in the early twentieth century and has since expanded into a highly successful research program to explore the processes of diversification and adaptation. Nonetheless, the ability of that framework satisfactorily to accommodate the rapid advances in developmental biology, genomics and ecology has been questioned. We review some of these arguments, focusing on literatures (evo-devo, developmental plasticity, inclusive inheritance and niche construction) whose implications for evolution can be interpreted in two ways—one that preserves the internal structure of contemporary evolutionary theory and one that points towards an alternative conceptual framework. The latter, which we label the 'extended evolutionary synthesis' (EES), retains the fundaments of evolutionary theory, but differs in its emphasis on the role of constructive processes in development and evolution, and reciprocal portrayals of causation. In the EES, developmental processes, operating through developmental bias, inclusive inheritance and niche construction, share responsibility for the direction and rate of evolution, the origin of character variation and organism-environment complementarity. We spell out the structure, core assumptions and novel predictions of the EES, and show how it can be deployed to stimulate and advance research in those fields that study or use evolutionary biology., (© 2015 The Author(s).)

Network-based diffusion analysis (NBDA) is a statistical method that allows the researcher to identify and quantify a social influence on the spread of behaviour through a population. Hitherto, NBDA analyses have not directly modelled spatial population structure. Here we present a spatial extension of NBDA, applicable to diffusion data where the spatial locations of individuals in the population, or of their home bases or nest sites, are available. The method is based on the estimation of inter-individual associations (for association matrix construction) from the mean inter-point distances as represented on a spatial point pattern of individuals, nests or home bases. We illustrate the method using a simulated dataset, and show how environmental covariates (such as that obtained from a satellite image, or from direct observations in the study area) can also be included in the analysis. The analysis is conducted in a Bayesian framework, which has the advantage that prior knowledge of the rate at which the individuals acquire a given task can be incorporated into the analysis. This method is especially valuable for studies for which detailed spatially structured data, but no other association data, is available. Technological advances are making the collection of such data in the wild more feasible: for example, bio-logging facilitates the collection of a wide range of variables from animal populations in the wild. We provide an R package, spatialnbda, which is hosted on the Comprehensive R Archive Network (CRAN). This package facilitates the construction of association matrices with the spatial x and y coordinates as the input arguments, and spatial NBDA analyses.

Human culture relies on extensive use of social transmission, which must be integrated with independently acquired (i.e. asocial) information for effective decision-making. Formal evolutionary theory predicts that natural selection should favor adaptive learning strategies, including a bias to copy when uncertain, and a bias to disproportionately copy the majority (known as 'conformist transmission'). Although the function and causation of these evolved strategies has been comparatively well studied, little is known of their development. We experimentally investigated the development of the bias to copy-when-uncertain and conformist transmission in children from the ages of 3 to 7, testing predictions derived from theoretical models. Children first attempted to solve a binary-choice quantity discrimination task themselves using asocial information, but were then given the decisions of informants, and an opportunity to revise their answer. We investigated whether children's revised judgments were adaptively contingent on (i) the difficulty of the trial and (ii) the degree of consensus amongst informants. As predicted, older but not younger children copied others more on more difficult trials than on easier trials, even though older children also showed a tendency to stick with their initial, asocial decision. We also found that older children, like adults, were disproportionately receptive to non-total majorities (i.e. were conformist) whereas younger children were receptive only to total (i.e. unanimous) majorities. We conclude that, whilst the mechanism for incorporating social information into decision-making is initially very blunt, across the course of early childhood it converges on the adaptive learning mechanisms observed in adults and predicted by cultural evolutionary theory. A video abstract of this article can be viewed at http://youtu.be/Qb6JINGYqVk., (© 2014 John Wiley & Sons Ltd.)

Hominin reliance on Oldowan stone tools-which appear from 2.5 mya and are believed to have been socially transmitted-has been hypothesized 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 five 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.

The directed social learning hypothesis suggests that information does not spread evenly through animal groups, but rather individual characteristics and patterns of physical proximity guide the social transmission of information along specific pathways. Network-based diffusion analysis (NBDA) allows researchers to test whether information spreads following a social network. However, the explanatory power of different social networks has rarely been compared, and current models do not easily accommodate random effects (e.g. allowing for individuals within groups to correlate in their asocial solving rates). We tested whether the spread of two novel foraging skills through captive starling groups was affected by individual- and group-level random and fixed effects (i.e. sex, age, body condition, dominance rank and demonstrator status) and perching or foraging networks. We extended NBDA to include random effects and conducted model discrimination in a Bayesian context. We found that social learning increased the rate at which birds acquired the novel foraging task solutions by 6.67 times, and acquiring one of the two novel foraging task solutions facilitated the asocial acquisition of the other. Surprisingly, the spread of task solutions followed the perching rather than the foraging social network. Upon acquiring a task solution, foraging performance was facilitated by the presence of group mates. Our results highlight the importance of considering more than one social network when predicting the spread of information through animal groups. This article is part of a Special Issue entitled: Cognition in the wild., (Copyright © 2014 Elsevier B.V. All rights reserved.)

Numerous factors affect the fine-scale social structure of animal groups, but it is unclear how important such factors are in determining how individuals encounter resources. Familiarity affects shoal choice and structure in many social fishes. Here, we show that familiarity between shoal members of sticklebacks (Gasterosteus aculeatus) affects both fine-scale social organization and the discovery of resources. Social network analysis revealed that sticklebacks remained closer to familiar than to unfamiliar individuals within the same shoal. Network-based diffusion analysis revealed that there was a strong untransmitted social effect on patch discovery, with individuals tending to discover a task sooner if a familiar individual from their group had previously done so than if an unfamiliar fish had done so. However, in contrast to the effect of familiarity, the frequency with which individuals had previously associated with one another had no effect upon the likelihood of prey patch discovery. This may have been due to the influence of fish on one another's movements; the effect of familiarity on discovery of an empty 'control' patch was as strong as for discovery of an actual prey patch. Our results demonstrate that factors affecting fine-scale social interactions can also influence how individuals encounter and exploit resources.

Many animals exhibit social learning and behavioural traditions, but human culture exhibits unparalleled complexity and diversity, and is unambiguously cumulative in character. These similarities and differences have spawned a debate over whether animal traditions and human culture are reliant on homologous or analogous psychological processes. Human cumulative culture combines high-fidelity transmission of cultural knowledge with beneficial modifications to generate a 'ratcheting' in technological complexity, leading to the development of traits far more complex than one individual could invent alone. Claims have been made for cumulative culture in several species of animals, including chimpanzees, orangutans and New Caledonian crows, but these remain contentious. Whilst initial work on the topic of cumulative culture was largely theoretical, employing mathematical methods developed by population biologists, in recent years researchers from a wide range of disciplines, including psychology, biology, economics, biological anthropology, linguistics and archaeology, have turned their attention to the experimental investigation of cumulative culture. We review this literature, highlighting advances made in understanding the underlying processes of cumulative culture and emphasising areas of agreement and disagreement amongst investigators in separate fields., (© 2013 The Authors. Biological Reviews © 2013 Cambridge Philosophical Society.)

Niche construction refers to the activities of organisms that bring about changes in their environments, many of which are evolutionarily and ecologically consequential. Advocates of niche construction theory (NCT) believe that standard evolutionary theory fails to recognize the full importance of niche construction, and consequently propose a novel view of evolution, in which niche construction and its legacy over time (ecological inheritance) are described as evolutionary processes, equivalent in importance to natural selection. Here, we subject NCT to critical evaluation, in the form of a collaboration between one prominent advocate of NCT, and a team of skeptics. We discuss whether niche construction is an evolutionary process, whether NCT obscures or clarifies how natural selection leads to organismal adaptation, and whether niche construction and natural selection are of equivalent explanatory importance. We also consider whether the literature that promotes NCT overstates the significance of niche construction, whether it is internally coherent, and whether it accurately portrays standard evolutionary theory. Our disagreements reflect a wider dispute within evolutionary theory over whether the neo-Darwinian synthesis is in need of reformulation, as well as different usages of some key terms (e.g., evolutionary process)., (© 2013 The Author(s). Evolution published by Wiley Periodicals, Inc. on behalf of The Society for the Study of Evolution.)

Social learning mechanisms are widely thought to vary in their degree of complexity as well as in their prevalence in the natural world. While learning the properties of a stimulus that generalize to similar stimuli at other locations (stimulus enhancement) prima facie appears more useful to an animal than learning about a specific stimulus at a specific location (local enhancement), empirical evidence suggests that the latter is much more widespread in nature. Simulating populations engaged in a producer-scrounger game, we sought to deploy mathematical models to identify the adaptive benefits of reliance on local enhancement and/or stimulus enhancement, and the alternative conditions favoring their evolution. Surprisingly, we found that while stimulus enhancement readily evolves, local enhancement is advantageous only under highly restricted conditions: when generalization of information was made unreliable or when error in social learning was high. Our results generate a conundrum over how seemingly conflicting empirical and theoretical findings can be reconciled. Perhaps the prevalence of local enhancement in nature is due to stimulus enhancement costs independent of the learning task itself (e.g. predation risk), perhaps natural habitats are often characterized by unreliable yet highly rewarding payoffs, or perhaps local enhancement occurs less frequently, and stimulus enhancement more frequently, than widely believed., (Copyright © 2013 Elsevier Inc. All rights reserved.)

This year is the 50th anniversary of Tinbergen's (1963) article 'On aims and methods of ethology', where he first outlined the four 'major problems of biology'. The classification of the four problems, or questions, is one of Tinbergen's most enduring legacies, and it remains as valuable today as 50 years ago in highlighting the value of a comprehensive, multifaceted understanding of a characteristic, with answers to each question providing complementary insights. Nonetheless, much has changed in the intervening years, and new data call for a more nuanced application of Tinbergen's framework. The anniversary would seem a suitable opportunity to reflect on the four questions and evaluate the scientific work that they encourage., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

Humans have a form of externalised memory. They are able to transmit information across generations in the form of learned cultural traditions and preserve this knowledge in artefacts. How this capability evolved from the simpler traditions of other animals is an active area of research., (Copyright © 2013 Elsevier Ltd. All rights reserved.)