Human Behavioural Traits and the Polycrisis: A Systematic Review

Abstract

Polycrisis has recently emerged as a term to capture the human predicament, and analyses of the drivers of this have identified causes such as economic growth. However, identification of the fundamental, underpinning causes is needed to effectively mitigate the polycrisis, and human behavioural traits are likely to comprise such fundamental drivers. Specifically, traits that have become maladaptive in the context of modernity are particularly important. We identify three particular maladaptations that are likely the most direct contributors to the polycrisis: warfare; resource overexploitation; and human cognitive biases. A systematic literature review using these maladaptations identified key studies from which behavioural traits underpinning the maladaptations were highlighted. These identified traits form the basis of suggested leverage points in the global system to reduce the likelihood of a polycrisis.

1. Introduction

The global high-energy, high-complexity technological civilisation that humans have developed over approximately the last 100 centuries finds itself in an increasingly serious situation with respect to the functioning of Earth systems. This is because collective human civilisation acting at global scale is perturbing the overall Earth system to an increasingly severe degree. This growing predicament (which is taking place in a period of time broadly described as the ‘Anthropocene’ [1]) increasingly threatens the sustainability of human societies in their different forms across the planet, and their capacity to persist at contemporary scales and forms into the future [2]. The impacts of human civilisation on the Earth system are manifold but amongst the most significant are destabilisation of the climatic system due to emission of greenhouse gases (GHG), destruction of habitats and ecosystems leading to a sixth mass extinction event [3], depletion of the resources, and toxification of the global environment as a result of growing industrial emissions [4].

The large and complex human societies that developed underwent a series of energy transitions and technological developments which culminated in the Industrial Revolution [5] (which may trace its origins in Great Britain to timeframes as early as the 17th century [6]) and Green Revolution (characterised by the global spread of changes in the use of land, machinery, and synthetic chemicals that started around 1960 [7]), which have together ‘supercharged’ human numbers and the collective impact of human activities at a global scale. A key aspect of recent timeframes (covering approx. the last 200 years of industrialisation, and the eight decades following World War II) is the ongoing acceleration and intensification of aggregate human activity, which is resulting in compounding impacts on the Earth system, and feedbacks from the Earth system on human systems due to this destabilisation [8,9,10].

A term has recently emerged in the scientific and general discourse that captures the predicament of human civilisation in contemporary timeframes, and which may increasingly develop in coming years and decades: ‘polycrisis’. Polycrisis remains a relatively loosely defined term but broadly speaking it captures the potential for the occurrence and accumulation of parallel and interconnected crises and shocks (which may occur at different scales, ranging from global to regional to local) in the highly complex Earth–human ‘supra-system’. Crises emerging in this situation are prone to causally entangle, compound, synchronise, and interact, and thereby create crises greater than the sum of their parts [11,12].

Although it is evident that global civilisation has been accelerating towards polycrisis conditions for many decades, this has not been due to a lack of foresight in different forms. Reasoned arguments have been put forward by many in the scientific community that a reduction in the magnitude of human activity will be necessary to avoid such a polycrisis. Amongst the most insightful of these early warnings is the seminal study in The Limits to Growth (LTG) [13], published in the early 1970s (with updates being produced in the 1990s and 2000s) [14]. The key findings of the LTG study were that under the scenarios seeking to pursue ongoing growth both global population and industrial capacity would likely experience abrupt and unmanageable future declines (i.e., ‘collapse’) due to different causes such as resource depletion and accumulating pollution. Only the scenarios in which an equilibrium state (i.e., no net growth) is sought and achieved, is collapse shown to be averted [13,14].

Although it was undoubtedly influential and has been the subject of a resurgence in interest from approximately the late 2000s onwards [15], the key messages of LTG were not heeded in any serious or systemic way in the years and decades following its original publication. This is mostly attributed to its call to action having challenged the incumbent economic system and status quo. Its key message was variously regarded as untrue, alarmist, undesirable, and too fundamentally difficult to implement [16,17].

There may, however, be another perspective on the failure of global society to heed the findings of LTG. This may be that there are deeper root causes of the human predicament than those highlighted by LTG. Put another way, it may be that the societal goal of seeking growth is not the distal cause of the human predicament, but may in fact be a proximate cause of yet more fundamental drivers.

In this context, proximate refers to the immediate or closest apparent causative factor(s) for the occurrence of a given phenomenon, and distal refers to the ultimate, fundamental, or underpinning diver. Therefore, proximate causes may be considered to be emergent, downstream, or first (and nth) order effects of distal drivers. Effective mitigations should therefore target distal drivers, as these are most likely to dampen the downstream effects. Identification and comprehension of distal causes is therefore imperative for mitigation of the current human predicament, and for future societies facing similar challenges.

Humans have, of course, been the fundamental and consistent agents in the evolution of the human system through time (noting that the extent of direct human agency may have been reduced in recent timeframes due to the autopoietic effects associated with the global ‘superorganism’ [18]), so logically human evolution, and the behaviours associated with/emergent from the various aspects of this process, are likely be fundamentally important in the global predicament. The key factors to consider in the context are traits (or phenotypes), which in any given organism refer to the suite of physical (e.g., morphological, physiological, developmental, etc.) and behavioural (e.g., instinctual, cognitive, etc.) characteristics that it employs to achieve survival and reproduction [19].

Human traits (and more specifically, behavioural traits) may be considered as fundamental or distal drivers of the human/Earth system predicament. It is important to note that distal refers to the underlying traits and behaviours, and proximate the emergent phenomena (e.g., the cultural, economic, governance, and technological constructs that humans have progressively developed) arising from the distal traits [20]. The phenomena that emerge from humans interacting in accordance with these traits are quite direct in their emergence from collective human behaviours, namely the tendency toward economic growth (i.e., from maximisation of resource usage with behavioural modernity and technology), the power dynamics inherent within human constructs such as capitalism or global institutions, and approaches to governance such as democracy or autocracy. These may therefore be considered as ‘sub-distal’, i.e., not the ultimate underlying drivers, but closely associated with them.

However, some of these patterns of behaviour, actions, and features that result from traits, which can therefore be considered as ‘subdistal’ emergent phenomena, have developed into maladaptations. Maladaptations are those behavioural patterns that have a negative or disbeneficial effect on the rate of survival and reproduction for individual organisms, groups, or species. They are generally not selected for and so arise from changes in the context of adaptations such that their previous mechanism of effect changes from net positive to negative in terms of survival benefit. Due to the external factors involved, the shift from adaptive to maladaptive may occur at different rates (gradual or sudden) and be subject to delay and ‘cliff-edge’ effects.

In the context of humans, maladaptations are the behavioural patterns that primarily underpin Earth system destabilisation, and, as such, contribute to the potential future development of polycrisis conditions. Therefore, to effectively understand and address these maladaptations, this paper attempts to identify the traits which comprise their distal drivers. By doing this, the ultimate drivers of the human predicament may be identified.

1.1. Research Question and Objectives

The research question this systematic literature review aims to address is whether the current global predicament of accelerating Earth system destabilisation and a developing polycrisis may be underpinned fully or in part by a key distal driver (or drivers) that are more fundamental than those identified, assessed, or explored in the available literature on the Earth system and the human predicament.

The objectives are to elucidate and explore themes that emerge from evolutionary psychology, anthropology, and human behavioural science disciplines, among others. The aim of this is to compose a coherent narrative that may allow the identification of any key behavioural traits and maladaptive tendencies in humans that may contribute towards the development of an integrated view of the drivers of the global predicament. This then forms the basis for discussion of what responses and actions may be implemented to mitigate the impacts of these traits and maladaptations.

2. Materials and Methods

In order to identify the key human behavioural traits underpinning maladaptations (and therefore the global predicament), a systematic literature review was carried out [21,22]. The method applied in this study applied a three-stage approach, which is summarised in the following bullets, and expanded on in the following sections:

• Identification of key maladaptations, which underpin phenomena impacting the Earth system, through high level review of available studies.
• Identification and application of keywords to generate a ‘long list’ of studies from Web of Science (WoS), and then keyword/thematic review to produce a rationalised list of studies to undergo a systematic literature review using the PRISMA guidelines [23].
• Thematic review of the identified studies to interpret and extract key findings and concepts to allow the identification of key human behavioural traits associated with the maladaptations.

2.1. Identification of Themes Relating to Maladaptation

Searches were undertaken using the Google Scholar search engine to identify key peer-reviewed journal studies and other sources, which provided high level, summary and overview viewpoints on the human predicament, impacts on the Earth system, and key aspects of human nature and behaviour underpinning this.

This search was undertaken as an initial scoping (or ‘first pass’) search to identify key themes and discipline areas in which maladaptations were described and explored, in order to inform the search subsequently undertaken using Web of Science. Keywords and search terms used (in isolation and in combination) for the Google Scholar search included maladaptations; evolutionary psychology; anthropology; human behaviour; human evolution; human nature; human cognition; sustainability; predicament; and polycrisis.

Google Scholar was used for the purpose of an initial search for studies describing maladaptations, and Web of Science for the subsequent detailed literature search using identified keywords, due to the relative merits of these resources. Google Scholar was deemed suitable for provision of highly interdisciplinary studies (including a wide range of journal articles, preprints, and books), whilst Web of Science was well suited to a more targeted search of up-to-date and high-impact sources (journal articles and conference proceedings).

The use of other research databases in lieu of or in addition to these (e.g., Scopus and JSTOR) was considered, but it was decided that these would likely not offer any advantages over and above Google Scholar and Web of Science (such as larger databases and/or more effective search functions), and would add more complexity in terms of tracking the search procedures and outputs. The studies identified via Google Scholar are summarised in

Table 1. Studies used to identify maladaptations.

Study Authors	Year of Publication	Title
Marean	2015	An evolutionary anthropological perspective on modern human origins [24]
Waring et al.	2023	Characteristic processes of human evolution caused the Anthropocene and may obstruct its global solutions [25]
Hagens	2020	Economics for the future–Beyond the superorganism [18]
Li et al.	2017	The evolutionary mismatch hypothesis: Implications for psychological science [26]
Rees	2010	What’s blocking sustainability? Human nature, cognition, and denial [27]
Merz et al.	2023	World scientists’ warning: The behavioural crisis driving ecological overshoot [28]
Varki	2019	Did human reality denial breach the evolutionary psychological barrier of mortality salience? A theory that can explain unusual features of the origin and fate of our species [29]
Ehrlich & Ehrlich	2022	Returning to “normal”? Evolutionary roots of the human prospect [30]
Constantino et al.	2021	Cognition and behavior in context: a framework and theories to explain natural resource use decisions in social-ecological systems [31].
Bentley et al.	2015	Collective behaviour, uncertainty and environmental change [32]
Whitehouse	2024	Inheritance–The evolutionary origins of the modern world [33]

.

Thematic analysis of the studies listed in Table 1 identified a number of maladaptations in human behaviour and actions.

A consistent overarching theme through these studies was cooperation—in the general sense of the tendency for humans to form cooperative, highly socialised groups that provide far greater problem-solving and survival capabilities than humans have on an individual basis. This behavioural trait is novel amongst the hominids in terms of the extent and efficacy of its operation and has been the fundamental factor underpinning the establishment of humans as apex predators, allowing the overall human population to grow to eight billion individuals, and underpinning the building of socio-political complexity and technological civilisations that have come to dominate the planet.

From a purely biological perspective, cooperation can be said to have been highly adaptive, but in recent timeframes the phenomena emergent from cooperation (particularly the recent building of technical civilisations) have been fundamental drivers of Earth system destabilisation. Therefore, cooperation may be becoming increasingly maladaptive. Given that the central aim of this study is to link Earth system impacts and maladaptation, cooperation may seem a logical focus; however, it is an ‘umbrella’ term that captures a grouping of related behaviours (so may be considered a distal maladaptation). To analyse maladaptations, more discrete and readily definable human behaviours, actions, and phenomena, which by themselves have clear tendencies to drive Earth system impacts, should be the focus.

Therefore, from these papers and their thematic analysis, three key, interrelated maladaptations under the umbrella of cooperation have been identified, as follows:

• Warfare—A proclivity for warfare is described as a key driver for early human migration and destruction of other hominids, and an endpoint of escalating between-group competition and perpetual growth is conflict with global-scale impacts [11].
• Resource Overexploitation—The tendency towards resource exploitation is described as having been selected for by evolution [26], which has been enhanced by technology [27], and availability of resource surpluses has been a key factor in the emergence of dominance hierarchies [11].
• Cognitive Biases—These are described as beliefs, behaviours, and predispositions [28] that are biologically fundamental to K-strategists [27], which helped maintain social groups [18] and shaped cultural tendencies [33]. The combination of theory and mind and death denial may be unique human biases [29], and biases are a contributor to complex and context-dependent human decision-making [31].

2.2. Identification of Keywords

Keywords associated with the different maladaptations were identified through thematics analysis of the applicable studies identified above (Table 1); these are listed in

Table 2. Keywords used against each maladaptation.

Maladaptation	Keywords
Warfare	Adaptation; Human Evolution; Psychology; Human Traits; Anthropocene (total: 5)
Resource Overexploitation	Human Traits; Evolution; Anthropocene (total: 3)
Cognitive Biases	Adaptation; Human Evolution; Psychology; Human Traits; Heuristics; Anthropocene (total: 6)

. The procedure used for undertaking the searches in Web of Science was to enter into the search bar the maladaptation in question for the search, i.e., (warfare; resource overexploitation; or cognitive bias), along with the term (human behaviour) (i.e., this was entered into every search combination, as trial and error identified this to be effective for narrowing search results to relevant topic areas) and the (keyword) in question (refer to Table 2). This procedure was repeated for each maladaptation and keyword, and is summarised as follows: (maladaptation) and human behaviour AND (keywords identified for each maladaptation).

2.3. Web of Science Search Outputs

The maladaptation/keyword search criteria defined above and in Table 2 yielded the following results (i.e., these are the collated lists of studies resulting from the different keyword combinations used):

• Warfare: 125 studies
• Resource Overexploitation: 322 studies
• Cognitive Biases: 216 studies

The next stage of the systematic literature review [23] was to review the studies identified via Web of Science using the following stages and criteria:

• The studies returned from these searches varied in terms of applicability and relevance to the research objectives of this review, so the first stage was to undertake a manual review of the titles for keywords and thematic applicability (the titles of several of the studies indicated the themes were from wholly or partially unrelated disciplines, e.g., computer science, management, medical science, and philosophy). This ‘coarse screening’ stage eliminated a significant proportion of the studies for each maladaptation from further consideration.
• Following the initial ‘coarse screening’ against study titles, manual assessment of the abstracts and body text for each of the studies was undertaken, again applying the keywords and thematic applicability. This was to ascertain applicability to the specific requirements of the systematic literature review (in some cases the study title indicated applicability, but further consideration of the objectives and findings of the study revealed the subject matter was of no or limited relevance to the objectives of this review), and this ‘deeper dive’ allowed another significant proportion to be eliminated from further consideration.
• Time limits (i.e., time since publication) were not applied apart from the literature had to be included in the Web of Science; this is because some more dated publications (i.e., published prior to 2000) contain key concepts that are still current.

This process allowed the number of studies per maladaptation to be significantly reduced, with the final study lists numbering 25 for warfare, 20 for resource overexploitation, and 18 for cognitive biases (full lists of studies are presented in Appendix A,

Table A1. List of studies for warfare.

Study Authors	Year of Publication	Title
Dawson	1999	Evolutionary theory and group selection: the question of warfare [34]
Sell et al.	2012	The importance of physical strength to human males [35]
Durham	1976	Resource competition and human aggression [42]
Sell et al.	2017	Physically strong men are more militant: A test across four countries [36]
Durrant	2011	Collective violence: An evolutionary perspective [45]
Zefferman & Mathew	2015	An evolutionary theory of large-scale human warfare: Group-structured cultural selection [46]
Lopez	2015	The evolution of war: theory and controversy [47]
Sugiyama	2014	Fitness costs of warfare for women [54]
Rusch	2014	The two sides of warfare—an extended model of altruistic behavior in ancestral human intergroup conflict [53]
Cashdan & Downes	2012	Evolutionary perspectives on human aggression [48]
Mathew & Boyd	2014	The cost of cowardice: punitive sentiments towards free riders in Turkana raids [50]
Boyd & Richerson	2021	Large-scale cooperation in small-scale foraging societies [41]
Bellamy et al.	2018	A computational framework for modelling inter-group behaviour using psychological theory [51]
Cacault et al.	2015	Do we harm others even if we don’t need to? [52]
Rusch	2013	Asymmetries in altruistic behavior during violent intergroup conflict [49]
Hammerstein	2013	What theoretical biology has to say on aggression in humans and animals [57]
Pitman	2011	The evolution of human warfare [58]
Hames	2019	Pacifying hunter–gatherers [56]
Carter & Kuchnick	2018	Male aggressiveness as intrasexual contest competition in a cross-cultural sample [43]
Taylor	2019	The puzzle of altruism: Why do ‘selfish genes’ behave so unselfishly? [40]
Gómez et al.	2016	The phylogenetic roots of human lethal violence [39]
Glowacki & McDermott	2022	Key individuals catalyse intergroup violence [37]
Henriques et al.	2019	Acculturation drives the evolution of intergroup conflict [55]
Fuentes	2017	Human niche, human behaviour, human nature [38]
Gayo et al.	2023	Towards understanding human–environment feedback loops: the Atacama Desert case [44]

,

Table A2. List of studies for resource overexploitation.

Study Authors	Year of Publication	Title
Brandt et al.	2012	Human adaptive behavior in common pool resource systems [72]
Janssen & Scheffer	2004	Overexploitation of renewable resources by ancient societies and the role of sunk-cost effects [75]
Anderies	2000	On modeling human behavior and institutions in simple ecological economic systems [74]
Dacko	2015	The issue of environmental resources management in the light of the model of Tragedy of the Commons—systemic approach [69]
Joshi et al.	2020	Emergence of social inequality in the spatial harvesting of renewable public goods [73]
Vuorinen et al.	2021	Why don’t all species overexploit? [60]
Abrams	2019	How roes the evolution of universal ecological traits affect population size? Lessons from simple models [61]
Bayham et al.	2019	Social boundaries, resource depression, and conflict: A bioeconomic model of the intertribal buffer zone [62]
Bagawade et al.	2023	Multi-scale effects of habitat loss and the role of trait evolution [63]
ten Broeke et al.	2019	Cooperation can improve the resilience of common-pool resource systems against over-harvesting [70]
Tu et al.	2021	The emergence of cooperation from shared goals in the Systemic Sustainability Game of common pool resources [71]
Vermeij	2012	The limits of adaptation: Humans and the predator-prey race [104]
Gowdy	1995	Trade and environmental sustainability: An evolutionary perspective [76]
Silliman et al.	2013	Consumer fronts, global change, and runaway collapse in ecosystems [105]
Aoki & Wakano	2022	Hominin forager technology, food sharing, and diet breadth [66]
Foley & Lahr	2015	Lithic landscapes: Early human impact from stone tool production on the Central Saharan environment [64]
Porcasi et al.	2000	Trans-Holocene marine mammal exploitation on San Clemente Island, California: A tragedy of the commons revisited [106]
Young et al.	2016	Patterns, causes, and consequences of Anthropocene defaunation [65]
Barnosky et al.	2016	Avoiding collapse: Grand challenges for science and society to solve by 2050 [67]
Rammel et al.	2007	Managing complex adaptive systems—A co-evolutionary perspective on natural resource management [68]

and

Table A3. List of studies for cognitive biases.

Study Authors	Year of Publication	Title
Haselton et al.	2009	Adaptive rationality: An evolutionary perspective on cognitive bias [85]
Tobena et al.	1999	Advantages of bias and prejudice: an exploration of their neurocognitive templates [86]
Jefferson	2017	Born to be biased? Unrealistic optimism and error management theory [94]
Griffiths et al.	2008	Theoretical and empirical evidence for the impact of inductive biases on cultural evolution [107]
Santos & Rosati	2015	The evolutionary roots of human decision making [83]
Boudry et al.	2015	Can evolution get us off the hook? Evaluating the ecological defence of human rationality [84]
Efferson et al.	2020	The evolution of distorted beliefs vs. mistaken choices under asymmetric error costs [108]
Butterworth et al.	2022	The better to fool you with: Deception and self-deception [92]
Azzollini et al.	2023	A study on five cognitive biases [80]
Meneganzin et al.	2020	Anthropogenic climate change as a monumental niche construction process: background and philosophical aspects [82]
Ortman & Spiliopoulos	2023	Ecological rationality and economics: where the Twain shall meet [91]
Beckage et al.	2022	Incorporating human behaviour into Earth system modelling [109]
Varella	2018	The biology and evolution of the three psychological tendencies to anthropomorphize biology and evolution [88]
Nicholson	1997	Evolutionary psychology: Toward a new view of human nature and organizational society [87]
Catalano et al.	2017	Black swans, cognition, and the power of learning from failure [89]
Brosnan & Jones	2023	Using an evolutionary approach to improve predictive ability in the social sciences: Property, the endowment effect, and law [95]
Arce & Winkelman	2021	Psychedelics, sociality, and human evolution [110]
Shults	2015	How to Survive the Anthropocene: Adaptive Atheism and the Evolution of Homo deiparensis [90]

).

The number of studies identified in the initial Google Scholar search (11) and the total number used in the review (63) were relatively low compared to the total yielded by the Web of Science search (663). However, as stated, a large proportion of the search results were found to be unrelated to the subject matter, which is likely a function of the number of search terms and keywords used, and could not be used. The final ‘shortlist’ of studies taken forward for review was also of a size amenable to practical review whilst providing sufficient breadth, so was therefore an appropriate balance of representation and detail against practicality.

Figure 1 shows the flow diagram for the systematic review using the PRISMA guidelines [23] and PRISMA checklist (see Supplementary File).

The procedure and approach for undertaking the thematic analysis of the studies identified through the literature searches was to review in detail each document identified from the search (i.e., full review of aims, objectives, findings, analysis, and conclusions) to identify content that thematically linked to the research questions and objectives of this study.

Where relevant content was identified, key themes, concepts, and findings were extracted and summarised. Given that multiple points and concepts were identified from single studies in many cases, and from across all of the studies reviewed, these were integrated into coherent narratives that linked to and underpinned the research questions and objectives, and provided the basis for discussion and for conclusions to be drawn. Note that thematic content was not extracted from all of the shortlisted studies that were reviewed, although these studies had been identified as of general relevance. In some cases, these did not provide any content that robustly supported the objectives of this study.

3. Results

3.1. Warfare

The identified traits of the warfare maladaptation can be found in Box 1. Several hominid species appear to have evolved the capacity for close social interaction, and thereby attained the capacity for developing and transmitting cultural information within their groups. Humans were likely the most successful at this (evidenced to a large degree by humans being the sole surviving hominid species), and may have experienced a ‘breakthrough’ in terms of language and cultural development, which allowed them to win the ‘cognitive arms race’ with other hominids. The context for the development of these features was the thousands of generations spent living in small bands of genetically/ethnically related hunter–gatherers, and one key feature that may have emerged from this is ethnocentricity, or groupishness. This is the tendency to form exclusive, ethnically centred social groups that encourage the favouring of other members of the group and the fearing of outsiders. This likely has a genetic component (selected for by the enhanced survival of groups with internal loyalty), which has been enhanced by cultural transmission. Supporting evidence for this includes symbolically marked boundaries demarcating ethnic groups (e.g., cave paintings), which proliferated starting approximately 35 thousand years ago [34].

Box 1. Identified traits of the warfare maladaptation.

Groupishness; ethnocentrism; territoriality; parochial altruism; male violence proclivity; male pacification; collective/coalitional violence; war norm psychology; fighting ability cues; overconfidence; group selection; risk-taking; dominance; enculturation; cultural drive; bellicosity.

A tendency to fear strangers and act aggressively towards out-groups in accordance with ethnocentricity may be one of the key precursors to warfare. In this context, males have likely developed a proclivity for violence. This is due at least in part to the dual pressures of males of having a high potential upper limit to reproductive success (i.e., there is a driver to fulfil this), along with risk of violent death without leaving descendants, creating a selective pressure for males to use violence for conflict resolution. This has resulted in features such as aggressive attitudes and upper body strength (a proxy for fighting ability) acting as cues for enhanced leadership ability, which may have been a driver for warfare. Aggressive men took leadership positions and applied this aggression at group scale, and gained a selective advantage from coordinating coalitional aggression (or collective violence, which is defined as that undertaken by people who identify as part of a group). This is underpinned by the potentially large influence individuals can have on group behaviour, with traits such as boldness and exploratory behaviour being correlated with leaders, which are more likely to initiate intergroup conflict (though in humans this is highly social-context dependent) [35,36,37].

Ethnocentrism appears to be a feature of human nature, with negative responses to out-group members occurring fairly readily (and is potentially modulated at the neural level by the peptide oxytocin), which may have required a mental shift to start to see other groups as ‘the enemy’ [38]. This may also be part of the genetic inheritance of humans as member of the class Mammalia; lethal violence is common amongst mammalian species (observed in up to 40% of all mammals) and is more prevalent in more socially and territorially oriented species in the group. Given that humans have both of these characteristics, it is likely that a propensity towards violence is phylogenetically inherited. Given this, the ability of humans to live in high density societies in the modern era may have been made possible only through the cultural pacification of these tendencies [39].

An alternative view is that for hunter–gatherer societies in Pleistocene environments, population density was so low (the population of the entire European continent may have only been 30 thousand individuals 15 thousand years ago, compared with 500 million in contemporary timeframes) that there was limited scope for resource competition. Furthermore, these bands were likely highly egalitarian in terms of the status of men and women, and therefore had no economic stratification. These social features might suggest that the use of violence is not an innate tendency in humans [40], and there is also strong evidence that ancient hunter–gatherer societies cooperated at large scale to build sizeable ‘collective goods’ (e.g., animal traps and irrigation works), which would have required a high degree of planning and cooperation [41].

This is supported by a lack of direct evidence for large-scale and coordinated violence for most of human history (prior to approx. 7500 years ago), which may imply that the lethal violence that did occur primarily comprised interpersonal events such as revenge and honour killings, disputes over reproductive partners, etc. However, from approximately 6–7 thousand years ago evidence of large-scale and coordinated violence becomes apparent, which appears to have risen in prevalence alongside shifts in human societies, including domestication, sedentism, technological changes, and the embedding of concepts such as property ownership [38].

Whether or not humans have a strong genetic predisposition towards violence, the tendency towards warfare has likely been enhanced by cultural evolution. Where assumed to be innate, the organisation and social sanction of killing (in the context of warfare but generally not in interpersonal interactions) would tend to become an embedded part of culture where aggression provides consistent net benefits (or a higher probability of this) to the aggressors (e.g., gaining or retaining fitness-enhancing resources) [42]. Where assumed to not be innate (i.e., the establishment of cooperative social behaviour in some circumstances), cultural factors may have still resulted in its evolution following the densification of populations following the widespread adoption of agricultural lifestyles [39].

The tendency towards inter-group collective violence has likely persisted, as it has conferred benefits including enhanced access to resources (e.g., food, territory, reproductive partners; societies with relatively more female-biased sex ratios are known to show higher levels of aggressiveness [43]); and safety (by eliminating threats). It is also likely an outcome of general human cognitive capacity (i.e., advanced mental models and forward planning), and another important dynamic is likely to have been environmental conditions; competition between groups (and therefore warfare) has been observed to intensify when environmental conditions became unpredictable and/or adverse (e.g., persistent droughts) [44]. Inter-group violence has therefore likely been an important selective force in human evolution that helped shape key traits such altruism and ultrasociality (as well as leading to approx. 150 million human fatalities since approx. 2300 BC) [45].

The psychological mechanisms that drove small-scale warfare may have continued to operate in later, complex societies (where great personal risks may be taken for the benefit of remote leaders and non-kin); this war norm psychology may be the driver for ongoing human warfare at scales and intensities far greater than analogues in the animal kingdom (e.g., chimpanzee ‘raiding’ behaviour, which is usually limited to small numbers of individuals), even if evolutionarily novel [46,47]. Furthermore, cultural factors may amplify the benefits to fighting beyond the cost/benefit risk assessment that past societies engaging in violence undertook. More specifically, overconfidence may have emerged as a signalling and feedback mechanism (potentially in conjunction with scarification type signals, e.g., tribal tattooing) to enhance morale and intimidate rivals, but may have also served to amplify the perceived benefits (‘prestige’) of warriors and fighting (which is likely mostly lost in the complexity of modern warfare) [43,48].

Warfare may also have conversely been a driver for the evolution of altruistic behaviours, specifically in the context of defence of in-groups (which may be described as a parochial altruism heuristic). This is because risk-taking in the collective defence of the in-group will have promoted survival and resource protection for the whole in-group, so would have been strongly selected (whereas altruism in attack would have been less likely to directly confer fitness benefits) along with punishments and sanctioning (e.g., ostracism from groups) for failures to cooperate for the group’s benefit. Parochial altruism may also have been used by leaders, authorities, etc., in many contexts as a mechanism to promote and stabilise in-group co-operation by invoking external (out-group) threats. This may therefore have been a root cause, but potentially not the only cause, for many genocides, war crimes, and atrocities through history, i.e., to motivate the defence of in-group interests at moments of critical danger (whether real or perceived) [49,50,51,52,53].

The fitness benefits of warfare have not always been evenly distributed; the capacity for inter-group war was primarily selected in males (as warriors have been predominantly male in virtually all societies throughout history [43]), and costs were likely disproportionately experienced by females. Women were likely frequently the target of collective violence actions (i.e., capture for reproductive purposes), which may have driven a tendency to cooperate with attackers (i.e., for survival, through conforming with and manipulating captors), an ability to assess attacker’s formability (through cues such as upper body strength), and encouragement of ferocity and/or disparaging timidity within groups (i.e., to reduce overall vulnerability to out-groups). Thus, collective female psychology may have also been significantly shaped by lethal violence [54].

Group selection (i.e., natural selection acting at group scale) would have also acted in these circumstances. Well-led, cohesive groups would have out-competed any uncoordinated groups that emerged from large group fissions. Warrior-rich and engagement-prone groups likely gained selective advantage and were able to spread their cultural variants (i.e., high aggressiveness). In this interpretation, groups act as the hosts or bearers of memes and institutions, and cultural traits of aggressiveness (which may be labelled as cultural drive) may spread by promoting its own transmission at the expense of its hosts’ fitness (such as overproduction of warriors, many of whom die before reproducing). This would also provide an explanation as to why conflict has continued to spread even when it has become maladaptive to groups and societies [55].

The rates of interpersonal and group violence experienced in modern, complex state-based societies has decreased by a large degree relative to historical societies (by up to a factor of 40 in state vs. non-state societies [56]), but this cannot be primarily attributed to genetic changes. Therefore, social learning, and, in particular, enculturation (alternatively acculturation; a bias that promoted the formation and spread of organisational, legal, educational, etc., institutions) has been the primary driver of this change. Aggression may be genetically inherited, but ‘male pacification’ is a culturally derived trait that emerges in the right conditions. It is likely the driver of institutions such as trade and marriage, may also be a key marker of human cognitive capacities (i.e., for traits such as reconciliation and empathy to come to the fore) and a driver of cultural evolution. However, even with embedding of acculturation, aggressive traits and group selection are likely to have been fundamental underpinning factors in the evolution of the hierarchal social systems that dominate modern societies [56,57,58].

3.2. Resource Overexploitation

The identified traits of the resource overexploitation maladaptation can be found in Box 2. Humans are defined as a K-selected species (or K-strategists), which are typically large-bodied, long-lived, and late-maturing species that produce small numbers of offspring. Populations of K-selected species tend to generate populations commensurate with the carrying capacities (K) of their local environments/ecosystems, and the survival of individuals, along with the overall evolutionary success of the species, depends on competitive superiority at high population densities under conditions of resource scarcity. Due to the competitive drive for habitat and resources that this mode of living and reproduction produces, natural selection has tended to favour those most adept at satisfying short-term selfish needs (whether this occurs via individual competition, or through in-group cooperative means). Future discounting and innate competitive drive (which do not have a natural ‘limiting function’) therefore likely have a heritable aspect, and over time may habituate to any level of available consumption (i.e., satisfaction quickly diminishes at new levels of consumption), meaning the tendency to consume and accumulate tends to ratchet up [27].

Box 2. Identified traits of the resource overexploitation maladaptation.

K-strategy/selection; individual competitive drive; in-group cooperation; instant gratification (future discounting); preferential harvesting; prioritisation of individual needs; joint resource use; anonymous free-riding; transgressor sanctioning; future productivity investment; prioritisation of losses/gains, imperception of depletion; sunk cost behaviour, elite co-opting.

In the ecological context, these drives were generally ‘held in check’ (i.e., preventing them attaining destructive levels of overexploitation) via a number of complex feedback mechanisms. These mechanisms are emergent features of highly networked ecosystems, and arise from the fact that exploitation–maximisation traits are not unique to humans. These traits are, in fact, evolutionary imperatives, given that natural selection will tend to propagate genes that promote prioritisation of survival/reproduction over sustainable resource use, and these competitive drives to maximise resource exploitation are fundamental to all living organisms [59].

The ‘baseline’ for all organisms will therefore be destructive resource use to expand populations and maximise survival potential. Given that the seeking of individual benefits is ‘prioritised’ by genes, the drive towards evolutionary suicide should be frequently observed; this is, however, not the case. This is because mechanisms that reduce the intensity of exploitation are common in biological systems, and operate as part of multispecies food webs, creating interactions between ecological and evolutionary processes. These exploitation limiters operate as networked, density-dependent eco-evolutionary balancing feedbacks that stabilise exploiter–resource interactions and counteract tendencies to overexploit.

These limiters operate via five principal mechanisms: top-down limitation (efficient exploiters expand their populations and become attractive resource for exploiters higher in the food chain, so become exploited); interference limitation (interference traits such as territoriality or predator alertness can appear within exploiter ‘guilds’, which increases individual resource shares, but reduces the overall intensity of exploitation, e.g., through creation of under-exploited buffer zones); cost-efficiency limitation (increased exploitation capacity is subject to diminishing returns, and also requires trade-offs with capacity for other biological functions such as reproduction); resource-trait limitation (increasing exploitation can drive counter-adaptations in the resources that reduce or annul the attack rate of the exploiter); and spatial heterogeneity limitation (exploiters will aggregate in resource ‘patches’ with the highest resource density, and the creation of localised metapopulations may reduce the scope for landscape-level overexploitation) [60,61,62,63].

Throughout evolutionary history, the counteracting forces enhancing or suppressing exploitation have tended to generate equilibria, but where suppressive feedbacks do not develop sufficiently rapidly, destructive exploiter–resource interactions can emerge. However, embedded networks of balancing feedbacks usually do eventually suppress these run-away tendencies, and for most of human history this has been in the form of pathogens and resource constraints. However, technological, purposeful human societies (from approximately the Palaeolithic onwards) using exosomatic energy [5] along with tools, weapons, and versatility of diets and environments, have been able to ‘break away’ from being strongly constrained from these feedbacks.

This resulted in a crucial adaptive shift, with the emergence of organised and directed exploitation (and associated environmental modification) of dense and predictable resources from approximately early (Middle Pleistocene) stages of hominid evolution (starting with stone tools [64], and later game animals, agricultural products, and eventually mineral ores and fuels). The effectiveness of human resource extraction even in remote timeframes is evidenced through the reduction in faunal diversity, which progressed from effective hunting at low levels, then scaling of hunting, through to the taking of ecological ‘space’, and finally perturbation of underlying ecological systems (e.g., via climatic change) [65]. These behaviours have laid the foundations for growth (and eventually exponential growth) of resource exploitation activities for human-specific purposes (the spatial spread of which is akin to that of general invasive species) [60,66,67].

Socio-political mechanisms could in principle act in an equivalent manner to the natural balancing feedbacks where overexploitation is concerned (i.e., analogous to a complex adaptive system [68]). The social–ecological systems (SESs) that human societies have inhabited through deep time have frequently depended on common pool resources (CPRs), which can experience overexploitation due to the conflict between individuals and the collective (i.e., tragedy of the commons). The impulse to prioritise individual gain is observed to be frequently overcome (particularly when the CPR is constrained) through investment in future productivity; however, due to the high intrinsic uncertainty and variability of environmental/ecological systems, future payoffs are discounted. The financialisation of human societies has also generated dishonesty, distrust, and unwillingness to communicate, which challenges the aversion of this outcome.

Olson’s theory suggests that members of a group will not work for a common good (unless in the form of pure altruism) except where they personally benefit from such actions, and in large groups users may succumb to antisocial effects such as passivity and becoming free-riders due to anonymity. Privatisation, which is often promoted as a solution to this, may be flawed or ineffective as owners can still deplete resources; not all resources can be readily divided and/or are inherently difficult to divide. Interventionism (i.e., state regulation and supervision) is prone to grow in scope and become corrupt, and can also be slow to implement [69,70,71].

Current opportunities and perceptions of future returns are highly correlated and interlinked [72]. Most SES are spatially structured such that they promote cooperation by exposing ‘defectors’ to the negative local consequences of their selfish acts and allows clusters of cooperators to form (i.e., ‘positive assortment’). However, space also allows mobility, which can allow defectors to escape the negative local consequences of their selfish acts. Cooperation is favoured when dispersal costs are high, whereas consumers all tend to be mobile and overexploitative when dispersal costs are low [73].

The key points from past experiences may be that interventions in overexploitation may need to be highly site-specific to be truly effective, and full understanding of the ‘geometries’ of problems may be crucial, along with awareness of the relationships between timescales on which resource governance systems must be developed and on which resource exploitation can intensify [74]. In the multiple large-scale societies through human history that have experienced collapse, a common contributory factor may be resource depletion driven by the sunk-cost effect. This describes decision-making that is influenced by prior investments (e.g., capital, experience), which tends to drive societies to continue with ‘business as usual’ practices (with overexploitation being a common effect) during periods of stress or disruption.

One key manifestation of this is successive increases in socio-political complexity (which has been one of the main ‘outputs’ of resource exploitation in multiple societies through time), the increasing control of economic surpluses by socio-political elites/prioritisation of elite requirements, and the degradation of peripheral zones in order to prioritise imperial centres. Conversely, sunk costs may provide benefits as they are one way in which the tragedy of the commons may be avoided. There is likely an optimal level of sunk costs where relative stability may be achieved, but too much has likely driven overharvesting in many contexts [75,76].

3.3. Cognitive Biases

The identified traits of the cognitive bias maladaptation can be found in Box 3. Cognitive biases are defined as any systematic (but unconscious) dispositions or inclinations that occur in human cognition, judgement, reasoning, processing, interpretation, decision-making, and behaviour that deviates or distorts in an observable or measurable way from objective reality, logic, probability reasoning, and plausibility.

Box 3. Identified traits of the cognitive bias maladaptation.

Error management; niche construction; patternicity; theory of mind; general purpose optimising; anthropocentrism; human supremacy; teleological reasoning; religiosity; denial; mortality salience; death anxiety tolerance; deception; self-deception.

These biases are linked to the evolution of human cognition in response to ambiguous situations and stimulus in variable and complex ancestral environments, where factors such as limited time, information, and cognitive processing capacity, along with urgent survival pressures, were constant influences. In these situations, heuristics (i.e., cognitive shortcuts or intuition used to generate rapid and adequate but not optimised decisions based on experience-informed judgement) emerged as a common form of bias. More than 200 discrete types and manifestations of biases have been described (including anchoring, clustering, confirmation, recency, and salience biases), and they are distinct from random error (i.e., which would arise from, e.g., sampling variability) [77,78,79].

Cognitive biases differ from errors in that although biases are common to most areas of human mental function and can lead to errors, they did evolve to increase fitness, whereas errors result from, e.g., fatigue and stress. The prevalence of biases also appears to generally be independent of cognitive skills and rational intelligence [80]. Humans have successfully survived and reproduced in a wide range of hostile and changeable environments over evolutionary time, which would indicate that human cognition is not as flawed or ‘muddled’ as some interpretations may suggest [81]. When tested with respect to ‘real’ environments, heuristics perform much better through their exploitation of the information structure of these environments. Nonetheless, clear biases can be seen in human cognition, e.g., giving attention to irrelevant information, being prone to be influenced by contextual and situational variables, and undertaking and rationalising bad decisions (which may lead to inadvertent large scale effects such as niche construction [82]), and these take place automatically and subconsciously [83,84].

The natural limit to human working memory capacity means that some information must always be disregarded, which can certainly lead to errors. Human heuristics appear to accommodate this by having a bias towards certain types of errors over others, which reflects asymmetries in the ‘costs’ of these errors. Bias in the direction of least costly error will favour over-reactiveness (e.g., reacting to what appears to be a snake vs. not reacting to an actual one), as this has lower costs than under-reactiveness. Error management biases minimise the costliest errors to the benefit of the decision-maker, and demonstrate that that superior decision-making processes are not always those that maximise accuracy but are systematically biased to commit false-positive errors with the lowest costs and harm [84,85,86].

Cognitive biases also reflect categorical thinking taking precedence over processual analysis (or described another way, labelling over analysis), which originate in different parts of the brain and provide energetic and time efficiencies, but may also be the source of, e.g., discriminatory social judgements and prejudices. Another perspective relates to failures in estimating probabilities and likelihoods, e.g., underestimating personal risks. Frequencies of events are observable in nature, whereas probabilities are mathematical abstractions that cannot be directly derived from sensory inputs. Additionally, when computing probabilities, information about base rates is lost; therefore, frequencies convey superior information [80,83,85,86,87].

It is also significant that in the natural environments in which humans evolved, pure randomness is rare. Therefore, fitness advantages would have been gained from a propensity to focus on outcomes rather than processes, and in spotting patterns, but a minor, modern consequence of this is vulnerability to (fair) dice games and roulette wheels; with their smooth and radially symmetric design, their abstracted behaviour is expressly designed to foil and exploit the pattern-matching heuristic that was previously effective (and still is) in natural environments. For example, natural phenomena such as weather and resources (e.g., food sources) can have ‘overdueness’ (after prolonged rainy periods it can be expected for this pattern to break and be replaced by different conditions, due to the statistical behaviour of atmospheric phenomena) and clumping (fruit-bearing trees may be found in abundance in certain locations and micro-climates). Therefore, the biases to ‘gamble’ on the kinds of environments with a specific hazard function and to employ positive temporal autocorrelation would have become beneficial [80,84].

The preferential detection of false positives may also be described as patternicity. This is the tendency to perceive patterns (in sensory input such as visual and auditory, but also purely cognitive) in meaningless noise, and causes the overattribution of agency, volition, and motive where there is only vague or no suggestive similarity. It differs from analogical reasoning (where schema transfer from a familiar domain is intentionally used to clarify a problem in another domain) and manifests as phenomena such as anthropomorphism, pareidolia, and teleology. In the latter case, this may be a unique human advancement over and above the theory of mind (ToM) and may be a complexly organised adaptive system descended from predator avoidance instincts of apprehension and vigilance. Basic goal and belief stances are the cognitive mechanisms that can animate fictional agents such as religious, mythological, folkloric, and extraterrestrial, but also more rational ones such as belief in evolution seeking ‘goals’. It is this latter case that may motivate phenomena such anthropocentrism, human supremacy, and the seeking of socio-technological progress; the persuasive rationale of teleological modes of thinking may have compelling and addictive aspects [88].

To early humans, disembodied intentional forces (i.e., supernatural agents and gods) were the ‘best guesses’ available because this provided further motivation to keep trying to detect hidden agents (i.e., potential predators) in a way that was cognitively cheap and inferentially rich, and over time these agents feasibly took on human-like qualities. This may be seen as a form of narrative fallacy in linking an unrelated series of events together into a logical story by imposing a pattern of causality on observations (i.e., ‘what you see is all there is’) [89]. With the later development of large and complex societies, religions acquired a new and additional purpose in ensuring all group members followed the norms of the group or coalition. However, these religious biases, although adaptive throughout the Neolithic, axial and up to the modern ages, are likely to be less effective in pluralistic, globalised civilisations. This may especially be the case where religious thinking detracts from efforts to address environmental degradation, i.e., resistance to ‘experts’ that challenge existing senses of identity and idealised/current forms of social ordering [90].

Future discounting is also favoured by natural selection, as organisms may not survive to reap future rewards so the costs of waiting may be higher. This contrasts with selection of delayed rewards, related in contrast to prefrontal and parietal areas of the brain associated with deliberative processes and numerical computation. A potentially unique temporal bias that humans exhibit is counterfactual or hypothetical reasoning, in which people are biased by past decisions and the emotional effect of regret. This is in essence a framing bias; options are considered not in absolute terms but rather relative to salient reference points. These factors indicate that human biases are adaptively rational and well matched to recurrent, naturalistic survival, and reproductive problems faced during evolutionary history [83,86,91].

The increase in the scope and power of human cognitive tools (i.e., language, theory of mind, episodic foresight) through evolutionary time has likely been a driver in the emergence of certain biases (and capabilities) [92]. One of these may be reality denial, which is a collective term for the human propensity for false beliefs, general optimism, and irrational risk taking. Two unique and crucial features of human cognition are ToM (the ability to consider and understand the thoughts of other individuals) and mortality salience (the ability to understand and consider mortality of the self and others). The first provides fitness advantages in social interactions, but also allows the recognition of deaths of conspecifics, and combined with the second should in theory result in (potentially severe) fear and anxiety of death, which would be a strong maladaptive psychological feature [29,93]. Therefore, it seems there is a feature of human psychology that allows death awareness whilst inhibiting the fear mechanism, which may be described as reality denial. This reality denial, even if resulting from a ‘one-time’ combination of psychological traits, could have become highly adaptive through contributing to hyper-prosociality and would be a key contributor to human optimism bias, and the generation and persistence of memes such as belief in the afterlife (which is another means to manage death anxiety) [29].

Self-deception may aid in the practice of deception by reducing the cognitive demands of holding contradictory information in mind simultaneously (which may result in cognitive dissonance, the avoidance of which is a driver for self-deception [89]), and also reduces scope for detection by dampening idiosyncratic deceptive cues (which would likely have been more readily detected in small, familiar groups). It may have also been selected for as a hedonic cognitive tool, which facilitates positivity and raised self-esteem that could have been important in the continuation of societal functioning in the face of hardships, e.g., environmental variation. Self-enhancement is a widely documented form of this bias, which manifests as overconfidence in abilities and beliefs, and intergroup behaviours where this can be witnessed include the sacrificing of rationality and information integrity to demonstrate political and other group-loyal ties [92].

In common with all other organisms, humans have three main choices in response to environmental variability: respond with greater behavioural flexibility, migrate to more stable environments, or die out. Humans have employed a mixture of the first two in order to survive and flourish, but in more recent timeframes (i.e., since the Agricultural Revolution) have expanded this tendency far beyond that of any other organisms through fast-evolving socio-cultural niche construction. This involves non-random (i.e., deliberate and purposeful) environmental modification (e.g., land conversion), which in turn imposes systematic biases and feedbacks. Biases that are relevant to a global predicament dominated by climate change include shifting baseline syndrome (i.e., slowly emerging patterns are quickly normalised); cognitive myopia (recent and local data such as prevailing weather conditions are prioritised over effects that are spatially and temporally unbounded); hyperbolic discounting (future costs and benefits are discounted at non-constant rates); and failure in statistical judgement.

Many cognitive biases may reflect adaptive behavioral predispositions that only became irrational when human environments changed in ways that rendered these predispositions as less beneficial. The speed of these changes (relative to evolutionary timescales) is a driver for the switch from rational to irrational. Optimism bias, which provided the motivation, focus, and sensitivity to threats that underpinned survival in past harsh, uncertain, and variable environments, now drives phenomena such as doubling down on sunk costs (‘escalating commitment’). Loss aversion (and the related endowment effect), which is a preference for the avoidance of losses relative to the acquisition of gains, was a key driver for retention of survival resources, now drives retention of desirable but harmful gains of modernity such as ready access to abundant high-energy foods [87,92,94,95].

4. Discussion

In terms of the research question and objectives outlined in Section 1.1, the evidence and narratives obtained from the literature reviewed in this study underpin the position that human traits, and related maladaptations, are likely to comprise major distal drivers of Earth system destabilisation. In turn, economic and population growth, alongside capitalism and the governance of global institutions, comprise downstream proximate drivers emergent from these traits and maladaptations.

In line with the objectives concerning responses and mitigations, the traits and maladaptations as identified in this review are potentially particularly significant. In the LTG study, the identified leverage points were associated with growth of the human system [96,97], but distal human traits and maladaptations are likely to be amongst the most effective leverage points. This is because they could provide the most direct route for ameliorating the aggregated human impacts that are driving large scale Earth system destabilisation, and in turn development of a global polycrisis. Mitigation efforts should be focused on weakening or dampening (‘reducing the gain’) of enhancing (positive) feedback loops in preference to the strengthening of balancing (negative) feedback loops [96].

A key common theme that is apparent from the majority of the traits identified is a general tendency towards prioritisation of spatially small and temporally near-term concerns by humans. This tendency likely emerged from evolutionary pressures and variable environmental conditions humans faced through deep evolutionary time, and served to ensure that humans and groups survived and passed on genetic and cultural inheritance over a large number of generations. Warfare, overexploitation, and cognitive biases underpinned this overall purpose but have become maladaptive in the modern context due to the emergent physical magnitude, technological, and energetic power, and lack of centralised control that has come to characterise very large-scale human societies. Therefore, these tendencies to prioritise the near (in time and space) may comprise a major leverage point for reducing human impact.

A transition to greatly improved collective cognisance and action on longer-term perspectives may be described as enhancing the collective wisdom of human societies, which is currently lacking in many of the fundamental ways [98,99]. Complex human societies have been enabled, and to a large degree defined, by cultural learning and transmission of information, but another key aspect has been the collective or shared myths and narratives that large human groups agree on, and which therefore become emergent quasi-realities for these groups [100,101]. These realities help shape the worldviews of collective cultures, and establish assumptive ‘foundations’ (i.e., are distal) that contribute to how they act; these collective myths may therefore comprise major leverage points. Changes to key aspects of these myths may therefore ‘operate’ this leverage point and ameliorate egregious effects. The following bullets are suggested focus points for enacting this:

• Reduction in territoriality—The division of the world into a multitude of sovereign nation states is a relatively recent innovation, but has been the basis and driver of the territoriality, which has underpinned much of the conflict that has occurred in recent centuries.
• Breaking cycles of violence—Violence and conflict begets further violence, which is often the product of tit-for-tat cycles that are often prolonged and hard to stop once underway, as humans and human groups are averse to ‘backing down’ or ‘losing face’.
• De-prioritisation of the individual—Modern (Western) societies have prioritised an individualistic culture that has been a driver of resource overexploitation and inequality, as well as larger-scale egregious phenomena such as the privatisation of gains and socialisation of costs.
• Reduce aversion to losses—Once certain status, comforts, material gains, and levels of abundance have been experienced by individuals and human groups (e.g., nations), there can be a stronger aversion to losing these ‘banked’ gains than loss of further gains, which has manifested as resistance to changes in ‘ways of life’, even when their unsustainability and the level of externalities imposed have been made clear.
• Changing time horizons—Endemic features of modernity such as short political cycles and ‘instant gratification’ consumerism has exacerbated natural biases towards hyperbolic discounting, leading to resource drawdown and creation of persistent and long-lived legacies and liabilities (e.g., novel entities, mass concrete waste, etc.).
• Recognition of the community of life—Anthropocentrism has placed human needs as central and downplayed the importance of the needs of other organisms and the networks linking them to humans and each other.
• Reduction in teleological reasoning—Despite the central role of science and technology in modernity, irrational thought patterns are still prevalent and being enhanced by greater access to information (e.g., conspiracy theories), which are proving destabilising to societies.
• Acceptance of limits/constraints—The distractions of modernity such as information overload, the pressures of surviving in neoliberal capitalist societies, conveniences and comforts, seeking of social status, and insulation from distant events makes (short term) denial of limits to growth and resource constraints more feasible.

The leverage points above are expansive and relate to many of the fundamental functions of modern societies, so could be too highly challenging to implement. Shifting cultural attitudes such as these would likely need to start with shifts in education and cultural institutions, along with governance structures. However, to avert the growing global polycrisis predicament, such shifts would likely need to take place across an appreciable fraction of humanity in relatively quick time (years–decades) to have a chance for making a difference. This would challenge a wide array of competing agendas, interests, legacies, and intentions at a global scale.

Intelligence in the form of essentially universal tool building and use of exosomatic energy [5] is undoubtedly unique to humans, and has provided humans with distinct survival advantage (i.e., enabling humans to become the apex predator in virtually all environments). Consciousness may not be unique to humans, but the confluence of tool building and consciousness in humans is likely a key aspect of general adaptiveness [102]. However, the long-term effectiveness and adaptiveness of intelligence and consciousness in the form humans display is far from proven or certain, and observation and interpretation of the current condition of human civilisation would suggest that intelligence is providing steadily lessening adaptive capabilities. The disparity of collectively failing to recognise the changing context of our survival strategies and our growing power for causing the destruction of our planetary life support systems is fundamental to our lack of wisdom: ‘The real problem of humanity is the following: we have Paleolithic emotions, medieval institutions, and god-like technology’ (E. O. Wilson, 2009) [103].

5. Conclusions

This paper identified key maladaptations in the human system that are having the greatest impacts on the Earth system as warfare, resource overexploitation, and cognitive biases. This study addressed the central research question by identifying that the major distal drivers of these maladaptations are likely to be collective human behavioural traits. More specifically, traits that provided humans with fitness and survival benefits during deep evolutionary time have led to behaviours that have become maladaptive in more recent timeframes. The systematic review of key studies relating to these maladaptations identified a number of the key underpinning behavioural traits (

Table 3. Behavioural traits against each maladaptation.

Maladaptation	Traits
Warfare	groupishness; ethnocentrism; territoriality; parochial altruism; male violence proclivity; male pacification; collective/coalitional violence; war norm psychology; fighting ability cues; overconfidence; group selection; risk-taking; dominance; enculturation; cultural drive; bellicosity.
Resource Overexploitation	K-strategy/selection; individual competitive drive; in-group cooperation; instant gratification (future discounting); preferential harvesting; prioritisation of individual needs; joint resource use; anonymous free-riding; transgressor sanctioning; future productivity investment; prioritisation of losses/gains, imperception of depletion; sunk cost behaviour, elite co-opting.
Cognitive Biases	error management; niche construction; patternicity; theory of mind; general purpose optimising; anthropocentrism; human supremacy; teleological reasoning; religiosity; denial; mortality salience; death anxiety tolerance; deception; self-deception.

).

In terms of the objectives relating to mitigations, the identified traits are likely to be the most effective leverage points for addressing the subdistal and proximate behaviours and consequent phenomena that are causing Earth system perturbation. A subset of these traits is likely to be particularly significant and might be the best leverage points to target. However, the wider perspective indicates human cognitive characteristics and limitations may make this challenging, and indeed the manifestation of maladaptations may be something of an inevitability.

This study identifies the need to address maladaptations as the distal, or fundamental, causes of the human predicament, along with making suggestions for the types of changes that would be required of human behavioural traits to increase collective wisdom. A much harder aspect for the future is defining how these changes might be effectively implemented and sustained at large enough scale and sufficiently rapidly to avert the coming polycrisis. Further work should be undertaken with great urgency to investigate this aspect.