The Evolution of Complementary Cognition: Cooperatively Adapt and Evolve through a System of Collective Cognitive Search

Helen Taylor , Brice Fernandes & Sarah Wraight

We propose a new theory of cognitive evolution, which we term Complementary Cognition. We build on evidence for individual neurocognitive specialization regarding search abilities in the modern population, and propose that our cooperatively searches and adapts through a system of group-level cognition. This paper sets out a coherent theory to explain why Complementary Cognition evolved and the conditions responsible for its emergence. Using the framework of search, we show that Complementary Cognition can be contextualized as part of a hierarchy of systems including genetic search and cognitive search. We propose that, just as genetic search drives phenotypic and evolution, complementary cognitive search is central to understanding how our species adapts and evolves through culture. Complementary Cognition has far-reaching implications since it may help to explain the emergence of behavioural modernity and provides a new explanatory framework for why and many aspects of cooperation evolved. We believe that Complementary Cognition underpins our species’ success and has important implications for how modern-day systems are designed.

Introduction we begin to see that they belong to a greater complex adaptive system (Taylor & Lockett In this article we propose a new theory of human forthcoming). cognitive evolution, which we argue lies at the core To reveal the significance of Complementary of explaining the exceptional adaptiveness of our Cognition, we begin by defining search and show species. In particular, we propose that members of that Complementary Cognition can be contextua- our species are individually specialized in different lized as part of a hierarchy of systems of search, at but complementary neurocognitive search strategies the level of the genome; cognition; and collective cog- and that consequently we regulate search for adap- nition. These systems of search are significant: results tive information at the group level, adapting of adaptive search can be inherited and so evolve cooperatively. The theory is grounded in Complex over time. Systems theory (e.g. Mitchell 2011) and the frame- Neurocognitive specialization in search predicts work of Search (e.g. (Hills et al. 2015). We call this that our species evolved in a highly variable environ- emergent system of collective cognitive search ment, since uncertainty is a key driver in the selection Complementary Cognition. of search capability, and variability makes search Cross-cultural patterns that occur in human cog- optimization at the individual level difficult. We out- nition suggest specialization in cognitive search, with line how such conditions prevailed during the evolu- large portions of the population having noticeable tionary history of our ancestors (Potts 1998) and must cognitive search biases. Looking at these comple- have created strong selection pressures for efficiency mentary patterns of specialization comprehensively, and capability in search.

Cambridge Archaeological Journal Page 1 of 17 © The Author(s), 2021. Published by Cambridge University Press on behalf of the McDonald Institute for Archaeological Research. This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (http:// creativecommons.org/licenses/by/4.0/), which permits unrestricted re-use, distribution, and reproduction in any medium, provided the original work is properly cited.

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We propose that such selection pressures, coupled with fundamental constraints in individual Definition of Complementary Cognition cognition, induced division and specialization in neurocognitive capabilities; that is, selection pres- Complementary Cognition is the theory that our species sures acting at the individual level resulted in the cooperatively adapts and evolves through a system of col- lective cognitive search. evolution of individual hominins with different but It is proposed that Complementary Cognition emerged as a complementary neurocognitive search strategies, consequence of individual neurocognitive specialization in resulting in the emergence of a new system of search search and co-evolution with language and aspects of at the collective level. cooperation. Specialization in search would only be possible Cooperative search between specialized individuals enables the co-creation of of higher fitness value. in the context of appropriate means of collaboration. It does not imply that individuals carry out exploratory or We propose that Complementary Cognition co-evolved exploitative activities exclusively, but rather are specialized so in a positively reinforcing feedback loop with aspects that they differ with regard to the neurocognitive capabilities of communication and cooperation. This has implica- that support search, and how information search is balanced. tions for an evolutionary theory of language, provid- Complementary Cognition can be contextualized as part of a fi hierarchy of systems through which our species adapts and ing two key reasons for its evolution: rst, as a evolves which includes genetic evolution. mechanism to facilitate collaborative search between Complementary Cognition evolved due to high environ- cognitively specialized individuals; secondly, as a mental variability during our species’ evolution, which cre- new inheritance channel to share the results of cogni- ated strong selection pressures for cognitive search capacity tive search. Particular features of human language and efficiency. Complementary Cognition contributes to our understand- conform with this theory. For example, the integra- ing of behavioural modernity and the emergence of cumula- tion of information from different search strategies tive cultural evolution. creates information of unbounded complexity, and so communicating the results of complementary cog- nitive search requires an open system, a key feature of human language. systems of genetic search and cognitive search differ We suggest that the evolution of Complementary in that the results can be stored, inherited and Cognition can be characterized as a Major Transition updated through further search. This leads to evolu- (Maynard Smith & Szathmáry 1995) and represents tion of adaptations over time. Complementary cogni- 1 asignificant transition in evolvability, enabling sub- tive search also has this property. stantially greater capacity, speed and flexibility to adapt than adaptation at evolutionary or cognitive What do we mean by search? scales. Living systems need to acquire and update adaptive We propose that the emergence of Comple- information: information that is of most adaptive mentary Cognition plays a central role in explaining value will vary over time and space, for example as our species’ remarkable adaptiveness. Humans predators or prey move, or as resources are depleted have come to thrive in nearly every terrestrial envir- or change with seasons and environmental variabil- onment on the planet. From tropical rainforests, ity, creating uncertainty that necessitates search savannah to tundra, our phenotypic adaptations (Hills et al. 2015). Search in some form is thus funda- are almost identical. Adaptation to such a range of mental to adaptation across species. has primarily been achieved through extra- The optimal search strategy will involve a mix- somatic or cultural adaptations—including behav- ture of exploration and exploitation. This can be ioural and technological adaptations (Binford 1962; viewed as a continuum. At extremes, all resources Richerson & Boyd 2005). The evolution of these cul- are fully allocated to either exploration or exploit- tural adaptations is open-ended and cumulative. ation of existing information. Consider a simple We propose that the reason for this adaptive cap- search for food in physical space. An organism can ability is Complementary Cognition and that it lies exploit the known area where it stands, exploiting a at the heart of explaining our cumulative cultural known patch of resources, or it could search more evolution. globally, exploring unknown areas for new patches of resources, or mix these behaviours. Search, adaptation and evolution Search is not restricted to physical domains. Abstract search can involve information domains, Although many systems of adaptation exist within such as the social domain or abstract design ideas, an organism (McGlade & Allen 1986), the adaptive or searching for a relevant piece of

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information. Abstract searches occur over informa- Inheritance enables the results of previously tion landscapes instead of physical ones. successful searches to be exploited. Further search Navigating this ubiquitous ‘exploration- or variation allows the possibility of increased exploitation’ trade-off is a search optimization prob- adaptation or adaptation to a changing environ-

lem (Crepinšek et al. 2013). Organisms continuously ment. The degree to which search is explorative face the dilemma of whether to pursue actions that may depend upon the processes that produced the exploit existing but possibly suboptimal information, variation. For example, asexual reproduction may or explore uncertain but potentially more profitable be less explorative than sexual reproduction as the solutions. While exploration may lead to better variation stems only from mutation and not recom- resources, exploring to the exclusion of exploitation bination as well (Page 2011). Sexual reproduction might result in too many undeveloped new ideas appears to be very effective at balancing the trade-off and a lack of refined skills and expertise. By contrast, between exploitation of alleles that were fitonaver- focusing too much on exploitation risks being age in the past, and sampling alleles in new combina- trapped in a local optimum or failure to adapt to tions (Chastain et al. 2014; see Watson & Szathmáry environmental change (March 1991). 2016). The most adaptive strategy depends on context The resulting combination of inherited and new and factors such as environmental familiarity, information will play a major role in determining resource richness and level of variability. The relative an organism’s fitness. in effect value of exploiting known resources versus the cost of searches for organisms that are adapted to their reducing uncertainty through exploration will also environment, each the result of a balance between guide the appropriate strategy. The optimal strategy exploitation and exploration. Thus genetic search may vary in time and space, as well as across drives phenotypic evolution. domains of search relevant to survival. Finding a generally optimal solution to this optimization prob- Variability, uncertainty and the evolution of cognitive lem is extremely challenging, and probably impos- search sible (Cohen et al. 2007). Genetic search has obvious limitations. It does not Regulating the balance between exploration and enable an organism to adapt to environmental exploitation in search is fundamental to adaptive changes during its lifetime. If changes occur faster success (Cohen et al. 2007). This trade-off arises in than can be adapted for genetically, adaptation many fields, often under a different name. In organ- must occur through other mechanisms for the izationalresearch, explorationencompassesthings such organism to survive and reproduce. Many species as search, variation, risk-taking, flexibility, experimen- have evolved the capacity for cognitive search, enab- tation, discovery and innovation. Exploitation ling behavioural adaptation during an organism’s includes aspects such as refinement, choice, produc- lifespan. In effect, the rate of change in the environ- tion, efficiency, selection and implementation (March ment relative to the frequency of genetic evolution 1991). In , the contrast is between extensive contributes to the need for other means of adapting versus intensive search (Benhamou 2007); in artificial to change. The notion that cognitive search is intelligence, breadth versus depth-first search (Korf selected for by high rates of variability is supported 1985); in time, long- versus short-term; in visual atten- by models in the field of cultural evolution which tion, diffuse versus focused (Rivière et al. 2017); in show that capacities for learning are selected for , global versus local (Todd et al. 2012). This over hard-wired behaviours in the context of envir- disparity in terminology has made it harder for spe- onmental variability (see Henrich & McElreath cialists in different areas to recognize these varied 2003). facets as part of the same underlying pattern. Cognitive search Genetic search Cognition is not generally viewed from the perspec- Darwin’s theory of evolution through natural selec- tive of search in the field of cultural evolution. tion can be interpreted as a search process (e.g. see Emphasis has been on how cultural adaptations are Watson & Szathmáry 2016 ) by which successful inherited and maintained between generations, adaptations are inherited and updated over time. with importance placed on mechanisms such as Every organism’s genotype can be regarded as a bal- social learning (Henrich 2017; Laland 2017; ance between inherited information that existed in Richerson & Boyd 2005) as well as other aspects the previous generation and variation arising for which support high fidelity and bandwidth of inher- example from mutation or recombination. itance such as niche construction (Sterelny 2011). In

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other words, how past adaptive knowledge is gradually co-opted for regulating goal-directed cog- exploited. nition (Hills 2006). So to complement physical search Notable exceptions include Jablonka and Lamb for tangible resources, organisms evolved cognitive (2005), who argue that focusing on replication and abilities to search for information related to those selection rather than on generation of variants and resources in internal and external information spaces reconstruction is counterproductive to understanding (Hills 2006). cultural evolution. Using the framework of cognitive Minimizing uncertainty is one of the key com- search to understand how behavioural adaptations putational principles that drives the search process. are inherited, and adaptive knowledge updated, Such uncertainty may relate to all aspects of an helps to address these concerns. organism’s world, from how relevant features are Hills et al. (2015) propose that cognition can be represented to the nature of its interaction with the envisioned as a search process, characterized by the world. The degree of uncertainty plays a key role exploration–exploitation trade-off. Search can be in both the initiation and termination of cognitive used as a common framework for understanding search (Winstanley et al. 2012). An organism familiar cognitive behaviour and the function of cognitive with an environment can use its knowledge and control across domains (Hills et al. 2015). internal models to make predictions about the conse- Regulating search is so central to adaptive success quences of cues, events and actions. If observations that optimally balancing exploration and exploitation differ from expectations to a degree that is not is believed to be one of the most important selective expected, i.e. unexpected uncertainty, this is likely to forces operating in the evolution of cognition (Cohen motivate search to update the understanding of the et al. 2007; Hills et al. 2015). environment’s properties (Winstanley et al. 2012). Search for food, water, mates or any other Even more fundamental is estimation uncertainty— resource is not a straightforward matter of simply referring to the degree of uncertainty in the estima- searching in physical space for a particular goal: it tions an organism makes based on previous experi- is a complex and multi-layered process. One way of ences (Winstanley et al. 2012). High levels of breaking this down is to consider search at the per- environmental variability will cause both unexpected ceptual level, search at the level of causal structure, uncertainty and estimation uncertainty to be high, the level of goal selection and the level of action potentially making search necessary at multiple selection (Winstanley et al. 2012, 128). In combin- levels of inference (Winstanley et al. 2012). ation, these different aspects of search enable an By being driven by environmental uncertainty, organism to construct models of the world so that cognitive search enables an organism to identify it can more successfully navigate its environment and characterize discrepancies between existing and locate resources necessary for survival and knowledge and observations, enabling knowledge reproduction. to be efficiently updated or improved to enable bet- Search at any level, will involve navigating the ter adaptation. trade-off between exploration and exploitation. With regard to external search, an individual’s sen- Variability and uncertainty in hominin evolution sory organs can usually only capture a small propor- It is well established that hominin evolution occurred tion of that information which is relevant to in the context of extremely high levels of variability adaptation, and even this exceeds the brain’srateof and thus uncertainty. Environmental evidence, such information processing (Hills & Dukas 2012). Thus as oxygen isotope measurements, reveals that the limited attentional resources must be allocated to period of human evolutionary history over the past that portion of relevant information that has the six million years corresponds with one of the most greatest effect on fitness (Dukas & Ellner 1993; Hills dramatic periods of climate oscillation of the past & Dukas 2012) in a similar way to how resources 65 million years (Potts 1998). Hominid evolution must be allocated during search in physical space. coincides with longer and even more extreme cli- Beyond its external perceptual landscape, an matic oscillations (Potts 1998). The rate of change, organism may also search an internal information degree of variability or predictability as well as the space to retrieve relevant information from memory. level of resource richness or scarcity will all have Internal search involves navigating the exploration- been subject to significant variation, with variability exploitation trade-off similarly to search in physical occurring at all time scales from daily to millennial or external space (Hills & Dukas 2012). It is thought over which longer-term evolution occurs (Potts & that the same molecular machinery that first evolved Faith 2015). Adapting to these conditions would for goal-directed search in physical space was have required exceptional versatility.

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Local climatic conditions for early hominins also Resource-scarce environments are therefore likely to show exceptional variability. African climate is espe- exert a particularly strong selection pressure for high- cially influenced by variation in solar radiation fidelity exploitation and local or depth-first search. (Pearson 2013)—an important factor in explaining Concurrently, adaptation to a variable environment the dynamics of the earth’s climatic systems, particu- relies on allocating resources towards exploration. larly air circulation and rainfall. Hydroclimate The strength of the selection pressure for exploratory extremes and variability are thought to be cognitive search correlates directly to the degree of key environmental drivers of African hominin evolu- uncertainty and variability in domains relevant to tion (Potts et al. 2018). East African climate shows survival. periods of significant instability with drastic shifts Selection for behavioural flexibility and adapt- between arid and moist conditions amongst periods ability to complex changing and unpredictable con- of greater stability (Potts & Faith 2015). In some per- ditions has long been argued to be one of the key iods, major shifts occurred rapidly due to factors drivers in hominid encephalization (Potts 1998; such as volcanic or tectonic activity, while in other Trauth et al. 2010). However, capacity in cognitive periods change was slower, with vegetation, water search cannot increase indefinitely. Eventually, and other resources changing over spans extending improvements meet impassable functional and effi- beyond individual lifetimes (Potts 1998). Thus the ciency constraints. Due to strong selection pressures rate as well as the range of change differed in time for greater effectiveness and efficiency in cognitive and space. search, as well as difficulties in optimizing this pro- Potts and Faith (2015) identify 32 periods of cess, we propose that environmental selection pres- high climate variability over the past five million sures induced a division and specialization in years. They found that nearly all key technological hominin cognition. and biogeographic milestones in eastern Africa, as well as the first appearances of new hominin species, The evolution of complementary cognitive search coincided with periods of high variability. Of these 32 periods, they identify six, which represent the Division and specialization in cognitive search most prolonged and intense periods of variability. While differences in human search behaviours have Our own species, Homo sapiens, emerged during the been observed (e.g. Hutchinson et al. 2012), the sixth period which lasted from c. 358,000 to 50,000 notion that the individual members of our species BP (Potts & Faith 2015, table 1) and included a hyper- are neurocognitively specialized in complementary arid phase during the period 186,000–127,000 BP cognitive search strategies has not been previously (Pearson 2013). proposed. Division and specialization are common Optimizing search is difficult to achieve at the throughout nature. In the context of an already individual level, particularly when there is variabil- cooperating group, within-species division and spe- ity in the rate of change. Further, variability may cialization are favoured when features that confer fit- favour selection of those organisms that have a var- ness benefits are functionally incompatible (Rueffler ied diet, with this and other adaptive behaviours et al. 2012) or when efficiency benefits to reduction such as tool use necessitating search over a broader of task-switching costs or specialization reach a cer- range of domains. Corresponding cognitive search tain threshold (Cooper & West 2018; West et al. capabilities might need to co-evolve to support this. 2015). These criteria correspond well to the condi- The need to search across a broader range of tions outlined above and characteristics of human domains, related to food acquisition, social search, cognition. technology creation and so on, also creates further difficulties in search optimization as different Selection pressures to reduce task-switching costs domains may require different search strategies. Given the level of variability, efficiency gains from Increased search capacity is also likely to have avoiding task-switching costs are likely to have been under strong selection. Survival during hyper- been an important factor. The role of task- arid periods of resource scarcity would have switching costs in division of labour is well recog- depended on the accurate replication of existing sur- nized. In human economies, Adam Smith (1776) vival strategies, as well as further refinement to opti- identified that avoidance of task-switching costs mize adaptive knowledge. This concurs with patch through division of labour led to greater efficiency exploitation theory, which predicts that organisms and productive gains ‘saving of the time which is will stay longer and exploit the same patch when commonly lost in passing from one species of resources are scarce (Stephens et al. 2012). work to another’.

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Similarly, organisms may need to change activ- For example, procedural memory enables skills to ities or move locations. Switching between any tasks be automatized, supporting efficient exploitation of will involve costs in time and energy, but higher information (Lum et al. 2013; Nicolson & Fawcett costs are incurred when switching between tasks 2000). Conversely, if an individual has difficulty that are very different or incompatible, such as acquiring automaticity of a skill, they may still have exploration and exploitation. Beyond avoiding declarative (conscious) awareness of the process. extrinsic costs which occur due to the context in This way of processing information is less efficient which the task is carried out, the avoidance of with regard to exploitation, but the trade-off is that internal metabolic costs associated with cognitive it enables exploratory search to continue, so that task-switching may also be a factor in division of the process might be improved or integrated with labour (Chittka et al. 1997; Goldsby et al. 2012). other declarative information (Nicolson 2014). Specifically, different large-scale networks Another fundamental example of physical con- within the brain are involved in different aspects of straints relates to the structure of minicolumns in cognitive search. For example, the default network the brain. Minicolumns are an elementary unit in is involved in the exploration of new behavioural the of all mammalian brains and are essen- patterns, whereas the dorsal attention network is tial in cortical information processing (Buxhoeveden broadly associated with high-order processing, such & Casanova 2002). Differences in connectivity within as evaluation, revision and exploitation (Beaty et al. and between modular cortical circuits result in differ- 2018; Mittner et al. 2016). Such networks are inhibi- ences in how information is processed (Casanova & tory of one another and switching between them Tillquist 2008; Williams & Casanova 2010). The incurs a measurable energy cost. Increasing uncer- width, density and connectivity of minicolumns all tainty has been found to be associated with less affect their function, and at a certain threshold cannot efficient (slower and less accurate) cognitive task- be further optimized for global and local information switching performance (Cooper et al. 2015). In the processing simultaneously (Williams & Casanova context of strong selection pressures for different 2010). Optimization in one direction must come at search behaviours, the avoidance of cognitive switch- the cost of a deficiency in the complementary area ing costs alone might therefore account for some as a result of physical trade-offs (Williams & level of division and specialization in cognitive Casanova 2010). search between different individuals. It is worth noting more generally that studies of brain evolution indicate that any significant enhance- Functional constraints ment of brain power would require a simultaneous Another contributing factor to division and special- improvement in neural organization, signal process- ization is functional constraints of the brain itself. ing and thermodynamic efficiency, with such a scen- At a certain threshold, a generalized brain reaches ario being unrealistic due to trade-offs that exist limits on its search capacity and efficiency. Beyond between these factors (Hofman 2001). In other words, those limits, the only way to increase capacity is due to physical constraints, and despite their plasti- through specialization. city (La Rosa et al. 2020) there is little scope for In the brain, there are many examples where individual human brains fundamentally to increase increased capability in global versus local search in capability (Hofman 2001). must be traded off (see Taylor & Lockett). Such con- We postulate that selection to reduce task- straints are observable in evidence from very differ- switching costs may initially have induced division ent areas of cognitive psychology and neuroscience. of labour in hominin cognition. It is possible that With regard to perceptual search, for example, some level of division of labour in cognitive search various studies indicate that increased capability in may have evolved fairly early on in hominin evolu- global and local visual search are mutually antagon- tion. Once some degree of specialization in an area istic (Geiger & Lettvin 1987; Schneps et al. 2012). This of search had evolved, an individual may have trade-off appears to be due to physical constraints become predisposed to further specialization in a that at least partly relate to the location and density of similar search strategy in other domains due to certain (magnocellular) cells in the brain (Schneps shared mechanisms. For example, specialization in et al. 2012). more exploratory search in physical space might pre- An example of functional incompatibility also dispose an individual to specialization in aspects of relates to the way information is stored in memory exploratory internal search. —this will affect the availability of information and We propose that due to functional constraints how it can be used for internal cognitive search. the only way that search capability could increase

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beyond a certain threshold was through the evolu- Mechanisms evolved in our own lineage to help tion of neurocognitive specialization: that is, indivi- with coordination, such as: gaze following (Brooks duals were specialized to navigate the search & Meltzoff 2005), iconic gesturing (Tomasello & process in complementary ways, with neurocogni- Call 2019), second-order (Gowlett tive specializations that supported search at different et al. 2012) and, of course, language. scales from more global to local or exploitation Division of labour and specialization must have (Taylor & Lockett forthcoming). Some differences in occurred in the context of an initially cooperative neural structures might also have evolved so that fur- group (West et al. 2015). Specialization creates a sec- ther specialization exists regarding capability in dif- ondary selection pressure for further lowering coord- ferent domains of search, but this remains ination costs. We propose that many features of speculative. cooperation, particularly language, may have We do not suggest specialization would pre- emerged as part of a coevolutionary process with clude an individual specialized more in explorative specialization in cognitive search. cognitive search from exploiting information, or vice Specialization typically comes at a trade-off to versa; however, since specialization comes at a trade- reduced complementary abilities. When high effect- off in complementary realms, they will be less effi- iveness is required in multiple domains, specializa- cient and have less aptitude than an individual tion is only possible if the loss of functionality in with a more depth-first search or exploitative complementary domains are compensated through strategy. other means (also called ‘compensated trait loss’). Such specialization in cognitive search has sig- Notably, compensated trait loss can involve essential nificant implications for our understanding of why traits and can occur at various gradations, ranging many aspects of cooperation and communication from complete loss of a trait to vestigialization also evolved. (Ellers et al. 2012). Compensated trait loss transforms facultative relationships into obligatory ones: that is, Co-evolution with lowered coordination costs it makes individuals inter-dependent for survival, Our species’ unique ability to collaborate in complex tightening the ecological relationship (Ellers et al. ways at scale (Melis & Semmann 2010) has been dif- 2012). In the case of Complementary Cognition, ficult to explain from an evolutionary perspective. trait loss must be mitigated through interactions For example, there remains significant debate as to with other individuals that have complementary cog- why language evolved (see Szathmáry 2010, fig. 1, nitive search abilities. We propose that specialization for an overview of theories). Complementary in cognitive search created a secondary selection Cognition provides a new explanatory framework pressure for aspects of cooperation necessary for for understanding why such sophisticated levels of combining these different search capabilities. cooperation and means of communication such as The interaction between specialization and low- language evolved. ered coordination costs can be viewed as a positively As established in economic theory, division of reinforcing feedback loop. Environmental selection labour and specialization can only be beneficial in the pressures induce a division of labour and specializa- context of lowered coordination costs. Coordination tion in cognitive search, which in turn creates sec- costs are incurred in coordinating work or combining ondary selection pressures for lowered coordination the output from agents performing complementary costs as individuals become increasingly inter- tasks (Becker & Murphy 1992). Time and energy dependent for survival. The evolution of traits that may be lost due to misinformation, conflict and mis- lower coordination costs then enables further special- trust and poor coordination of tasks (Becker & ization to occur; thus cognitive specialization and Murphy 1992). The difficulty of coordination in real- lowered coordination costs co-evolve in a positively istic, noisy environments limits division of labour reinforcing feedback loop. and specialization. Given the importance of coordin- The evolution of cognitive specialization is fur- ation costs in specialization, mechanisms for redu- ther complicated by the interplay between different cing them can confer strong fitness benefits. search strategies. For example, specialization in Many examples of adaptations to reduce coord- explorative global search and consequent trait loss ination costs exist in nature, for example the coordin- may result in fitness benefits to other individuals ation mechanisms evolved by eusocial insects, such who have a local, exploitative search bias. There is as pheromone deposition in ants (Wilson 1962), likely to be a complex interplay between direct envir- or the waggle dance in honeybees (Frisch 1967). onmental selection pressures and secondary selection Other factors can reduce the cost of coordination. pressures, dependent upon the complementary search

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strategies of other members of the social group. of cognitively specialized individuals as envisaged Different selection pressures acting at the individual by Complementary Cognition. The results of comple- level lead to the emergence of collectives with com- mentary search strategies mean that information plementary cognitive search abilities. from different search strategies must be combined. The evolution of aspects of cooperation and We view language as evolving firstly as a means communication in humans due to interdependence of facilitating co-operative search, particularly of has previously been theorized by Tomasello et al. more abstract domains. Through language, indivi- (2005). Tomasello and colleagues propose that duals with complementary search strategies can humans exhibit ‘shared intentionality’, that is, collab- search a common information landscape and so oration with others towards shared goals, requiring a co-create models of the world and behavioural adap- unique motivation to share psychological states tations that are beyond the capability of any one indi- (Tomasello & Rakoczy 2003; Tomasello et al. 2005). vidual, or even cognitively similar individuals. This aligns with Complementary Cognition and the Environmental variability and antagonistic selection proposal that our species searches cooperatively pressures for search capability (exploratory versus towards shared goals. exploitative, different domains, etc.) made coopera- Complementary Cognition also may contribute tive search critical to successful adaptation. to our understanding of personality traits, since we Secondly, we follow Maynard Smith and can expect specialization to influence how indivi- Szathmáry (1995) in their proposal that language duals best collaborate. Different kinds of social can be viewed as a new channel of inheritance for aptitudes for information sharing and collaboration information as part of a Major Transition. However, will align with different cognitive specializations. Complementary Cognition provides a more compre- We speculate that there is some measurable correl- hensive explanation of why such a system was ation between personality traits and cognitive search needed: namely, as well as facilitating co-operative bias. search, creating composite information from results of different scales of search and different domains Specialization in cognitive search and co-evolution with of search would lead to a marked increase in the language complexity of the resulting information. Passing on Specialization will affect which cooperative traits are this complex information requires a mechanism of selected for. For example, in physical or perceptual inheritance capable of transferring information with space, forms of gestural communication such as greater bandwidth and unbounded complexity, a pointing, or shared intentionality, may have been key feature of human language (Hauser et al. 2002). adequate for cooperative search towards shared We therefore view language as an integral part goals. As capability and specialization evolved in of the system of Complementary Cognition, and sug- other search domains, other forms of information gest that the evolution of language and probably sharing were probably required. It becomes almost other cooperative abilities may be better understood impossible to search certain domains cooperatively, when contextualized as part of the evolution of especially abstract domains, without means of shar- Complementary Cognition. ing complex abstract information. Language enables such sharing. Human language differs qualitatively Emergent benefits of Complementary Cognition from that of other communication systems in the use of recursion, which endows human lan- Navigating search at the group level confers a number guage with a uniquely open-ended capacity to com- of important benefits such as: signifi cantefficiency municate abstract concepts (Hauser et al. 2002) savings; globally increased capacity in search; risk Complementary Cognition provides an explanatory mitigation at the individual and group level; and framework for language by outlining the selection recombination of different search strategies. Overall, pressures that could have led to its evolution, and the combination of these benefits significantly why a new channel of information transfer and increases the robustness of the group to environmen- inheritance with these properties was required. tal variability. It can be regarded as a meta-adaptation Two justifications stand out in support of the idea which represents a substantial qualitative improve- that language emerged as part of a co-evolutionary ment in evolvability. process with division and specialization in cognitive search. Firstly, the requirements on the channel of Efficiency and capacity information sharing differ between cooperation Specialization confers efficiency gains through self- of cognitively similar individuals and cooperation organization as a result of reduced time and energy

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cost in negotiation of task allocation.2 Specialized indi- functional requirement. Computer simulations of viduals also have greater efficiency and effectiveness biological evolution show that unchanging environ- in their specialist domain compared to a generalist at ments usually lead to non-modular networks that the cost of complementary capabilities. However, a are slower to adapt to new environments (Clune group of specialists will have significantly improved et al. 2013; Kashtan & Alon 2005). Modular networks effectiveness and efficiency overall compared to a evolve when a rapidly changing environment has group of generalists, as their individual deficiencies different overall problems made up of common sub- are compensated for by other specialists. This leads problems (Kashtan & Alon 2005). When the overall to a considerable group capacity advantage. goal changes, the connections between modules can rapidly rearrange in order to adapt. Simulations Risk mitigation show that modular separation is logarithmically pro- Collaborative search amongst specialist individuals portional to the rate of variation (Lipson et al. 2002); mitigates risks better than collaborative search studies in nature also find that modularity correlates amongst generalists, particularly in a variable envir- with frequency of environmental change (Parter et al. onment. Simple cooperation through risk pooling is 2007). quite common. A group of hunters using the same If search capabilities are modularized, retargeting strategy may mitigate risks by pooling the the search to a different goal, or shifting the strategy to resources acquired and sharing the rewards of their adapt to new environmental conditions, becomes a hunt. This preserves them from going hungry when the matter of changing how individuals communicate, likelihood of catching prey is less than 100 per cent. rather than requiring any fundamental change in However, simply pooling risks still exposes col- cognitive capability. This changes the structure of the laborative groups to the risk of their common hunt- network of collaborating individuals, without funda- ing methods becoming inadequate due to changes mentally changing the cognitive task each individual in their environment. In this case, all hunters carries out, leading to adaptation at the group level, would go hungry. They are in essence pooling the while greatly reducing the cost to adapt. risks of a single strategy amongst themselves. If each hunter used a different hunting strategy, or per- Synergistic effects haps mixed this with foraging, fishing, agriculture, In addition to an increase in total capacity, modular- etc., then the risk is pooled across all strategies, mak- ity/specialization and coordination grant an increase ing the entire group more robust to a strategic failure. in capability—the ability to perform qualitatively dif- This effect extends to cognitive search and ferent tasks. This has been demonstrated in digital search in abstract domains. Distributing risks among organisms, able to complete tasks that could not be different strategies allows a failure in one to be computed by any single individual by sharing partial absorbed by the payoffs from the others. Notably, task results with each other (Goldsby et al. 2012). This this requires pre-existing collaboration. If individuals qualitative difference in tasks that can be performed do not already pool risks, division of labour in differ- confers a fitness advantage to individuals who are ent strategies will yield no benefits, and a strategy part of collaborating groups. For Complementary that can potentially have large payoffs with a low Cognition, this would manifest in an ability to probability may be detrimental to individual survival. co-create adaptations qualitatively superior to those Combining information from multiple search possible for a single individual or a cooperating strategies also increases the search space and the group of cognitively similar individuals. ‘quality’ of search, avoiding the risk of being trapped Robustness can be defined as ‘the ability of a in a local optimum or missing the global optimum system to maintain functionality in the face of some through too diffuse a search. change or disturbance, which could be internal or external to the system’ (Jen 2005). All the factors dis- Modularity cussed here, risk mitigation, modularity, overall Modularity in search strategies enables reconfigur- increased capacity, provide the flexibility to respond ation to adapt search activities to different circum- and adapt to shifting selection pressures at the group stances in time, space and across domains. Modular level, leading to an overall significant increase in systems (i.e. those composed of specialized, cooper- robustness. ating subunits) tend to be more adaptive in general. Specialized subunits are more easily re-composed Interplay between search strategies and rearranged than tightly coupled systems, enab- Modularity in search enables search strategies to be ling the reconfiguration of a system to match a new recombined in a variety of ways to increase search

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capacity and capability. Explorative search captures Implications for cultural evolution less detail but enhances ability to identify deeper glo- Neurocognitive specialization in search has not pre- bal patterns—these can guide individuals with more viously been considered, nor has its potential effect local search strategies who then avoid being trapped on cultural evolution. With regard to inheritance of in a local maximum. As such, global search can adaptive knowledge (i.e. the results of previous improve the quality and efficiency of local search. searches), one aspect to consider is that individuals In the domain of physical space, for example, a glo- with different cognitive specializations may be bal search of an area may identify a promising area more reliable sources for different kinds of socially of food that a local search would have not reached, learned information. For example, specialists with but the local search can then harvest it more effi- an exploitative bias may be better at faithfully copy- ciently. Global search may also guide search across ing and recalling detailed or procedural information different domains. For example, a global search passed down through generations, such as the correct may identify a broader variety of cues in an ecosys- sequence and way of making a tool. Conversely, tem, enabling more accurate, longer-term predictions those with an explorative search will be better able regarding environmental change. This can in turn to identify global patterns in inherited information, guide local search in relevant domains, for example enabling generalizations and predictions about to exploit different resources not so affected by the unknown or ambiguous situations. In this way, cog- predicted changes. nitive specialization may also increase the bandwidth Conversely, local search will be more effective and fidelity of inherited adaptive information as well at identifying local patterns potentially missed by as increasing capability to update shared knowledge. global search. Local search can further refine the Cooperative search enables group knowledge to search outcomes of individuals with a global search undergo a process of ‘quality control’. A prediction bias. Considering a technological adaptation as an error between knowledge held in the social group example, an exploratory search might recombine (i.e. the results of previous searches) and observa- the use of a wooden tool and a stone hand-axe in a tions, or misalignment between different kinds of shafted axe. This would require a creative leap, but knowledge in the social group, can target further will better succeed in the context of detailed knowl- search to resolve those discrepancies specifically. edge of which tree at what time gives the appropriate Expending resources to targeted search confers sig- wood with sufficient strength. Essential in the suc- nificant efficiency benefits, continuously improving cess of the final shafted axe is the outcome of global the adaptiveness of inherited information. Different search making the leap to recombine wood and stone search strategies enable the creation of different mod- in a novel manner, but also detailed in-depth under- els of the world and will identify different kinds of standing of wood and stone as materials, and the prediction errors. Cooperative search therefore local depth-first search of these domains. enables individuals to co-create adaptations of Creating a common information landscape from much greater adaptive value. As a consequence of different search strategies using language leads to this process of inheritance, search through different information that is more adaptive than is possible strategies to update adaptive knowledge and co- through individual cognition, or with populations creation of adaptive information, we see evolution in sharing common cognitive search strategies. These those adaptations over time, that is, cultural evolution. collaborations may be possible with protolanguage, or without language, but many domains of search, Discussion: relation to existing research and areas especially abstract domains, become almost impos- for future research sible to search cooperatively without means of shar- ing complex abstract information. Alignment with variability selection Cognitive search from global/exploratory to Complementary Cognition aligns strongly with Potts’ local/exploitative may build on the results of previous ‘Variability Selection’ hypothesis, which proposes searches. These may be sourced from memory or that, rather than being adapted to a particular envir- acquired from other individuals. Through social learn- onment, humans are adaptive with respect to envir- ing mechanisms, individuals can ‘inherit’ information onmental variability, that is, inconsistency of gathered through previous searches from parents selective conditions (Potts 1998; Potts & Faith 2015; (vertical inheritance) or peers (horizontal inheritance). Potts et al. 2018). Complementary Cognition enables Information may be passed down through multi- precisely this—significantly, it increases the capacity to ple generations, potentially updated and modified adapt not to a particular environment, but to any kind of through multiple searches across generations. environmental context—any information landscape.

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Cultural evolution specialization and compensated trait loss. Trait loss Cultural innovation and transmission have long been is hidden at the functional level as long as the eco- seen as analogous to genetic mutation and transmis- logical interaction is maintained and so its preva- sion (Boyd & Richerson 1985; Cavalli-Sforza & lence tends to be grossly underestimated (Ellers Feldman 1981). The framework of search highlights et al. 2012). We suggest that its visual as well as func- the similarity between these processes. Genetic tional invisibility explains why specialization in cog- search drives phenotypic adaptation and cognitive nitive search has hitherto gone unrecognized. search drives behavioural adaptation. In a similar manner, we propose that Complementary Cognition Evolutionary theory of language and cooperation (cooperative cognitive search) lies at the heart of Several different theories have be proposed to explaining the exceptional level of cultural adaptation explain the evolution of language (Hauser et al. in our species. 2014; Szathmáry 2010, fig. 1). Complementary As highlighted by Page (2011, 160), ‘for any Cognition provides a new theory of how language type of entity, the appropriate level of variation evolved as a means of enabling cooperative search [search] will eventually emerge from the system. and as an inheritance mechanism for the more com- Moreover, that level will tend to track the rate at plex adaptive information that resulted. which the system churns’. In this manner, the evolu- Beyond language, certain cooperative traits may tion of Complementary Cognition enabled us to also have co-evolved with cognitive specialization. It adapt to the high levels of variability found in our is beyond the scope of this paper to provide an species’ evolutionary history. adequately in-depth exploration of these possibilities. When variability or scarcity diminishes, however, Some examples might include second- or third-order the increased collective search capacity represents theory of mind (Gowlett et al. 2012), which would an over-adaptation relative to the environment in enable convergence to a common understanding fas- which it evolved. Information is still created and ter than first-order theory of mind, and if so would updated, but becomes obsolete much more slowly. facilitate co-operative search towards shared goals. This would lead to an accumulation of adaptive infor- Complementary Cognition may also provide insights mation: cumulative cultural evolution. The mechan- into why humans are characterized by such extensive isms of accumulation described by Complementary non-kin cooperation. Cognition may be tested and help improve our exist- ing models of cultural evolution. Major transition We propose that the evolution of Complementary Apparent gap between behaviourally and anatomically Cognition can be characterized as a major transi- modern humans tion (Taylor & Fernandes forthcoming). These rare Complementary Cognition could contribute to evolutionary events occur when evolution favours understanding the apparent delay of around cooperation, division of labour, specialization and 100,000 years between anatomically modern humans interdependence to such a degree that there is a and typically modern human cultural behaviour loss of individuality (Maynard Smith & Szathmáry (Sterelny 2011). It seems that behavioural flexibility 1995; West et al. 2015). The transition brings about increased without any sign of concomitant change a new level of complexity that enables behaviour in physical adaptations, including increase in brain not previously possible (Suki 2012). size—a phenomenon that Renfrew dubbed the ‘sapi- The Major Transitions framework was first pro- ent paradox’ (see e.g. Renfrew 2008). Potts and Faith posed by Maynard Smith and Szathmary (1995). (2015) identified that the period from 358,000 to They regarded the transition from societies with 50,000 BP was among the most prolonged and intense protolanguage to modern societies with language periods of variability in our evolutionary history. as the last of the major evolutionary transitions in These observations concur with the theory of their framework (Maynard Smith & Szathmáry Complementary Cognition. We expect increase in 1995; Szathmáry 2010). The notion that a major tran- behavioural flexibility and adaptiveness to have sition occurred with regard to our own species has been selected for during the period of variability, remained controversial, however (see Calcott et al. but more clearly manifested once that variability 2011). We suggest that the proposed theory on the had subsided (no longer running to stand still). evolution of Complementary Cognition supports However, we would not necessarily expect con- Maynard Smith and Szathmary’s argument that lan- current encephalization, since we propose that guage evolved as part of a major transition, but pro- increased capability emerged through a process of vides a new explanation of this transition that aligns

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much more closely with their own theoretical frame- The shift in the topology and nature of information work than previously posited descriptions (Taylor & sharing is sufficient that we propose Complementary Fernandes forthcoming). Cognition can be characterized as a Major Transition. We suggest that the evolution of this system reached Implications and significance its greatest extent during the last of the intense peri- ods of variability highlighted by Potts and Faith Summary of Complementary Cognition (2015)(c. 358,000 to 50,000 BP) with the transition in We have proposed that humans primarily adapt adaptive capability it afforded being manifested in and evolve via a hitherto unrecognized system of the origins of behavioural modernity. complementary cognitive search, which we call Finally, we propose that Complementary Complementary Cognition. Cognition lies at the heart of explaining the exceptional High levels of variability prevailed throughout levels of cumulative cultural evolution that character- our species’ evolutionary history. These created ize our species. We suggest that accumulation of adap- strong selection pressures for greater effectiveness tive information occurred at times when our evolved and efficiency in cognitive search, as well as difficulties ability to search—which evolved due to high levels in optimizing this process. These selection pressures of variability and scarcity—became hyper-adaptive induced a division and specialization in human cogni- during periods of relative stability and abundance. tive search in different but complementary cognitive search strategies. Complementary Cognition within a hierarchy of This division and specialization created second- evolutionary systems ary selection pressures, co-evolving with language Complementary Cognition can be contextualized as and other aspects of cooperation, enabling coopera- part of a hierarchy of self-similar systems, each of tive search and more effective adaptation across mul- which contributes to the adaptation and evolution tiple domains. Significantly, a consequence of this of our species. This hierarchy includes phenotypic adaptation is that our species navigates cognitive adaptation and evolution, which is driven by genetic search at the group level. search and natural selection and behavioural adapta- We propose a new evolutionary theory of tion, driven by cognitive search (Fig. 1). The levels language, namely that as well as enabling greater of search, adaptation and evolution identified here cooperative search, it also evolved as a new inherit- correspond to those proposed by Eva Jablonka and ance channel to enable sharing of the more complex Marion Lamb in their book Evolution in Four adaptive information that resulted from recombin- Dimensions (2005). While we do not discuss epigen- ing information from multiple search strategies. etic evolution, this can also be viewed from the Language can thus be contextualized as an integral perspective of search (Stolfi & Alba 2017). part of this system of Complementary Cognition. Although evolution and cognition are clearly Through specialization in complementary search established areas of scholarship, they are not always strategies, as well as new mechanisms of inheritance, considered from the perspective of search in archae- this system enabled higher-fidelity inheritance of ology. The framework of search enables us to see adaptive information. It also provided a system of how adaptation and evolution at the genetic, cogni- quality control for adaptive information—search tive and complementary cognitive scales are analo- across different scales from more global to local as gous. All three identified systems have mechanisms well as across domains (e.g. time, social space, aspects by which previous search results (adaptive informa- of the natural world) to identify errors with pre- tion) can be selected, stored, updated and inherited. existing adaptations that could be updated to improve Because new searches can build on the results of their fitness value. Through searching cooperatively past searches, we see evolution over time, be that and combining the results of different strategies, it manifested in phenotypic or extra-somatic (including also enabled the co-creation of cultural adaptations behavioural and cultural) adaptations. At each scale, of much higher adaptive value. variability and uncertainty create the selection pres- Given the meta-adaptive nature of Complemen- sure for improved search capabilities, leading to the tary Cognition, its emergence represents a significant emergence of a new level of search to adapt. transition in evolvability, enabling substantially greater capability, speed and flexibility to adapt, Limitations than search at the level of genetic evolution, individ- ual cognition, or collaboration of neurocognitively It is important to not over-simplify our discus- non-specialized individuals alone. sion regarding Complementary Cognition. Whilst

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Figure 1. Hierarchy of search.

grounded in the theory of complex adaptive systems, evolution, rather than an end. There remains a great this article should be seen as an overview to contextual- deal of research to be carried out to understand the ize further development and publication of specific implications of Complementary Cognition for the forthcoming research. We have provided a broad over- modern population and systems, an area of particular view of the evolution of cooperative cognitive search, practical interest. We hope Complementary Cognition but it is by no means a comprehensive overview of cog- can be used as a lens to explore these areas fruitfully. nitive evolution in humans. For the sake of brevity, for example, many details were omitted regarding the cog- Closing remarks nitive evidence which are presented elsewhere (Taylor &Lockettforthcoming) and more in-depth treatments The fact is that no species has ever had such wholesale of the complex interactions between search biases and control over everything on earth, living or dead, as we domains of search were left undiscussed. We also specu- now have. That lays upon us, whether we like it or late that domains of search play an important role in spe- not, an awesome responsibility. In our hands now lies not only our own future, but that of all other living crea- cialization and affect the evolutionary trajectory of our tures with whom we share the earth. species, but these have not been categorized and (Attenborough 1979, 308) explored. It is likely there exists some level of sub- specialization regarding preferred domains of search Our species’ evolutionary history was shaped by in addition to general search tendencies. climate change and substantial environmental vari- Complementary Cognition should be seen as a ability. We are now facing a period of dramatic starting point in exploring a rich area of human climate change, in which our environment will vary

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at a rate hitherto unforeseen in human history. believe that only through such understanding, and However, this time, human activity is the cause of an understanding of the evolution of cognitive variability. We have in a sense come full circle. This search, can we harness our full potential and design may be more connected than we realize. systems, institutions and policies that enable us to This paper has proposed that our species adapts create an adaptive and sustainable society. and evolves via a system of group level cognition, that is analogous to evolution at the genetic level. Notes Hierarchical systems evolve from the bottom up—the reason the upper level evolves is to serve the purposes 1. Non-genetic (epigenetic) gene regulatory mechanisms of the lower levels (Meadows & Wright 2015). That is, may facilitate adaptation in response to novel environ- if adaption at the level of Complementary Cognition mental conditions, potentially contributing to evolu- fails, adaptation will eventually fail more generally. tion through transgenerational epigenetic inheritance, Successful adaptation relies on a careful balance and by altering gene expression states and thus phenotypes (Bošković& Rando 2018). Its role in of cognitive specializations and effective collabor- is, however, still poorly understood ation. Not balancing this well can lead directly to (Horsthemke 2018) and beyond our expertise: so we maladaptive behaviours and culture. Given the would like to acknowledge its potential relevance scale at which we now operate, such maladaptive here, but do not discuss it for these reasons. behaviour can have a rapid and significant impact. 2. Similarly, ant or bee foragers do not have to negotiate Complementary Cognition has enabled us to adapt their role; rather they undertake tasks depending on to different environments, and may be at the heart biological factors such as evolved stimuli response. of our species’ success, enabling us to adapt much Such behavioural differences may evolve when selec- faster and more effectively than any other highly tion is strongest for the amount of work performed, complex organism. However, our specialization and and task-switching is costly (Duarte et al. 2012), or to reliance on the appropriate balance of capabilities minimize neural or other costs associated with the task allocation process itself, all of which may create may also be our species’ greatest vulnerability. fitness benefits (Dornhaus 2008). The impact of human activity on the environ- ment is the most pressing and stark example of Acknowledgements this. Ultimately, climate change is caused by a failure to update our behaviour and adapt. Thank you to Mona Khatibshahidi and Dr Adrian We are facing severe negative consequences as a Wallbank for advice and help with writing and editing result of not acting on information we have known and Dr Brett Calcott for reading drafts and very helpful about for decades, with the scope and scale of suggestions. The research was supported by Professor those consequences increasing month on month. Nigel Lockett and Dr Cameron Petrie. How did the most adaptive complex organism on earth reach this point? Helen Taylor The challenge of collaborating and co-adapting Complementary Cognition, Entrepreneurship & Societal at scale creates many difficulties. We believe that Adaptation we have also unwittingly put in place a number of Hunter Centre for Entrepreneurship cultural systems and practices that may be under- University of Strathclyde mining our ability to adapt (see Taylor & Lockett 199 Cathedral Street forthcoming). These self-imposed limitations disrupt Glasgow G4 0QU our complementary cognitive search capability and UK may restrict our capacity to find innovative and cre- & ative solutions to the problems that we face. McDonald Institute Through evolutionary chance, we have been University of Cambridge placed in a privileged position to shape not only Downing Street the destiny of our own species, but that of the rest Cambridge CB2 3ER of the planet. We face a period of unprecedented UK environmental and cultural changes brought about Email: [email protected] by human activity, and the rate of change itself is continually increasing. Brice Fernandes It is critical that we both understand and har- Independent Scholar ness this system of Complementary Cognition. We Email: [email protected]

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T.T. Hills & T.W. Robbins. Cambridge (MA): MIT T.T. Hills & T.W. Robbins. Cambridge (MA): MIT Press, 1–7. Press, 125–56. Tomasello, M. & J. Call, 2019. Thirty years of great ape ges- tures. Animal Cognition 22, 461–9. Author biographies Tomasello, M., M. Carpenter, J. Call, T. Behne & H. Moll, 2005. Understanding and sharing intentions: the ori- Helen Taylor completed her PhD on the emergence of gins of cultural cognition. Behavioral and Brain social complexity in the fifth millennium at the Sciences 28, 675–91. University of Cambridge, and has since been affiliated Tomasello, M. & H. Rakoczy, 2003. What makes human with the McDonald Institute. She is currently Research cognition unique? From individual to shared to col- Associate and project lead on Complementary Cognition, lective intentionality. Mind & Language 18, 121–47. Entrepreneurship & Societal Adaptation at the Hunter Trauth, M.H., M.A. Maslin, A.L. Deino, et al., 2010. Human Centre for Entrepreneurship, University of Strathclyde. evolution in a variable environment: the amplifier Her research spans cognitive psychology, archaeology, lakes of Eastern Africa. Quaternary Science Reviews evolutionary theory, entrepreneurship and organizational 29, 2981–8. research—areas which are all connected by complex adap- Watson, R.A. & E. Szathmáry, 2016. How can evolution tive systems theory. learn? Trends in Ecology & Evolution 31, 147–57. West,S.A.,R.M.Fisher,A.Gardner&E.T.Kiers,2015.Major Brice Fernandes is a consultant, systems engineer and evolutionary transitions in individuality. Proceedings of entrepreneur in the field of cloud native software. He is the National Academy of Sciences 112, 10112–19. particularly interested in declarative software architecture, Williams, E.L. & M.F. Casanova, 2010. and dys- functional , flow-based programming, functional- lexia: a spectrum of cognitive styles as defined by reactive programming, polyglot frameworks and data-centric minicolumnar morphometry. Medical Hypotheses 74, software design. His current work is on GitOps, a declarative 59–62. approach to infrastructure operations based on reconciliation Wilson, E.O., 1962. Chemical communication among instead of events. Brice is a maintainer of the GitOps working workers of the fire ant Solenopsis saevissima (Fr. group, an industry initiative to define the GitOps principles Smith) 3. The experimental induction of social clearly and provide a common framework for modern soft- responses. Animal Behaviour 10, 159–64. ware and infrastructure operations. Winstanley, C.A., T.W. Robbins, B.W. Balleine, et al., 2012. Search, goals, and the brain, in Cognitive Search: Sarah Wraight is a passionate human rights advocate and Evolution, algorithms, and the brain, eds P.M. Todd, international development specialist.

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