Abstract
E.O. Wilson’s Genesis: The Deep Origins of Societies is one of a series of short books where the author has tried to explain human societies using ideas and concepts from biology. While Wilson is to be lauded for his recent efforts to reintroduce the notions of group selection and multilevel selection, he still sustains an emphasis on only Darwinian selection and reveals a bias toward seeing selection for groups as a result of selection on individuals (as is the case for insects), perhaps entangled with selection on groups. The effort to conceive of human societies as an example of eusocieties of social insects ignores most of the sociological works on human and societal evolution; and as a result, the book is not convincing in its argument. Despite the pleasant writing style, Wilson and other biologists writing about human societies need to engage the almost 200 years of sociological work devoted to understanding the evolution of human societies.
For five decades, Edward O. Wilson has sought to link biology to the analysis of human societies. Such was the promise in his path-breaking work, Sociobiology: The New Synthesis (1975), and efforts along these lines have continued in a series of shorter books. His recent book, Genesis: The Deep Origins of Societies (2019), is a short book dealing with big issues, from the origins of life through the emergence of multicellular organisms to the evolution of group-living organisms, culminating in the emergence of large-scale insect societies and then human societies. The main thrust of this fascinating book is the effort to view the evolution of human societies as yet another eusociety.
The evolution of eusocieties
Like so much of his work, this new book is beautifully written and poses the enduring question: What is the meaning of human existence? Professor Wilson emphasizes that no longer do we have to rely upon long-standing creation stories from religion because science has been able to explain the emergence of life and its organization; and increasingly, science provides the information necessary to understand how humans, as evolved apes, came to be organized into increasingly larger societal formations. Wilson first outlines the great evolutionary transformations of life from its origins through the transformation of bacterium-level cells to more complex cells, the exchange of DNA, and the transitions of cells to multicellular organisms to the organization of many species into group-living formations. The key breakthrough of these group-organizing life forms, Wilson argues, was the arrival of eusocial groups among termites some 200 million years ago in the early Cretaceous, followed by ants some 50 million years ago. And, as Wilson argues at the climax of the book, the potential for eusociality first appeared in the hominin (human) lineage with the arrival of Homo habilis, some 2 million years ago.
Eusocial societies are rare in the animal kingdom and involve a unique cluster of traits, in which organisms live in colonies composed of overlapping generations, communal nesting behaviors and cooperative care of offspring. Eusocial colonies also have a division of labor, with some members giving up their personal reproduction for the fitness of the larger social formation. Most eusocial organisms are species of ants, termites, some bees, and social wasps. A few others have been added to the list, including naked mole rats and snapping shrimp. In eusocial societies, members have evolved mechanisms to enhance not just the survival of individuals, per se, but also the survival of the colony as a whole. Most of the book documents the rarity of eusocieties, their benefits, particularly protection from predators and advantages over competitors in food foraging, and their costs in creating altruists that sacrifice their own reproduction to engage in behaviors that increase the fitness of the colony at large.
As is well known, Wilson (1998) has argued in recent years for a kind of group selection and even multilevel selection, but in so doing he is somewhat unclear just how far he is willing to go in viewing evolution as driven by multiple-levels of group selection and particularly non-Darwinian selection (see also Wilson and Wilson, 2007). He clearly emphasizes that much selection in eusocial societies is on individual organisms, giving them genetically driven behavioral propensities to form colonies with many members, divided labor, and altruism by some members to sacrifice personal reproduction for the communal good. Thus, much of what Wilson terms ‘group selection’ is, in fact, Darwinian selection on individual phenotypes that lead to group-oriented behaviors, with the group as a whole enhancing fitness. This is not the group selection so reviled by much of biology for the second half of the 20th century since it is not the group that is being selected upon but the individuals in the group.
Wilson has also argued for multilevel selection, but again what does he mean in this assertion? Does selection directly work on the individual phenotypes or structure of the group, or both? At one point near the end of the book, he notes that ‘group selection [may be] entangled in individual selection’ (p. 116). It is never entirely clear, however, what ‘entangled’ means, but he appears willing to go this far: selection occurs on both individuals and the group, changing biological phenotypes and perhaps organizational phenotypes of groups. Still, the more Wilson begins to address the sociocultural evolution of human societies, the more importantly he and others need to make explicit what is Darwinian selection on the individual and what is non-Darwinian selection on the structure and culture of organizational formations created by humans. Yet, while Wilson is clear that groups compete with each other, and indeed, are often at war, he does not address the question: Does the nature of selection change when it is on culture-using animals that can change their sociocultural phenotypes and, indeed, the very environments to which they must adapt? In other words, once one has left the universe of insects and other animals driven by genetically-based bio-programmers and entered a social world driven by culture as much as biology, it is essential to untangle the types of selection involved, and the level of sociocultural organization being targeted by selection.
Can this kind of fairly tepid group selectionist argument, making only entangled references to group selection, also explain the evolution of human societies? Wilson certainly thinks so. As will become evident, however, explanations for the evolution of eusocieties among tiny-brained insects without culture are not so easily transferred to explanations of not only human evolution as organisms but also to the properties and dynamics of the superorganisms – societies – that humans can construct.
Before addressing this fundamental problem in Wilson’s explanation of the genesis and origins of human societies, we want to illuminate two major points that separate Wilson from most sociobiologists. In particular, Wilson must be congratulated for recognizing weakness in sociobiologists’ efforts to explain away group selection as a distinct evolutionary force. One key point is that eusocial societies evolve from populations of life forms that have often already formed large clusters for collective protection (without a division of labor), and thus eusociality does not emerge anew but represents ‘a next step’ in the evolution of sociality among many species. And, as he emphasizes, very few species have taken this ‘next step’ in the evolution of eusociality, and the interesting question Wilson poses is: Why is organizing by a division of labor so rare? A related point is that the pillars of sociobiological refutations of group selection – that is, kin selection, reciprocal altruism, and inclusive fitness – do not precede eusocial organization but, instead, evolve after the next step to task specialization and stable group formations has occurred. That is, he believes that ‘close kinship within the group typically followed from the origin of eusociality’ (p. 120). Thus, Wilson rejects his own earlier assertions and those that many sociobiologists still invoke to explain away group selection on individuals driven by bio-programmers to favor kin and to reciprocate actions by others that enhance individual and group-level fitness. But, Wilson now holds that individual selection alone cannot explain the evolution of eusociality.
More generally, Wilson presents a devastating critique of J.B.S. Haldane’s and later advocates’, such as George Williams (1966) and Robert Trivers (1971), efforts to explain away group selection by merely individual selection. Unfortunately, the nature of group selection is not well conceptualized in biology, even the original formulation by Wynne-Edwards, and even the current version offered by Wilson and Wilson (2007). There is, it seems, not just an empirical ‘entanglement,’ as E.O. Wilson suggests, but a conceptual vagueness as to what is being selected upon in human societies: phenotypes of human organisms, levels of social structure, or levels of culture.
In fact, the notion of ‘group’ as conceptualized in biology is, itself, too limiting because, in sociological analysis, groups are just the building blocks of much larger social structures and their cultures – organizations, communities, institutions, societies, and inter-societal systems – that typify human social organization. Thus, there are many levels of sociocultural organization upon which selection works, and moreover, in the sociocultural realm, it is not the population of humans that is evolving (as emphasized in the Modern Synthesis) but, instead, the levels and types of sociocultural formations created and used by humans to adapt to their environments. Thus, there will always be ambiguity in theoretical work by biologists or even sociologists unless they expand the notion of ‘group’ to incorporate the many levels of social organization and culture that humans have created and, equally important, unless theorists recognize that Darwinian selection on individual phenotypes (and underlying genotypes) cannot explain the evolution of complex sociocultural formations because Darwinian selection assumes that selection is ‘blind’ and non-teleological, whereas almost everything about humans and their creation of patterns of social organization is teleological.
Thus, in the human social universe, the unit on which selection works can, of course, be individual phenotypes but more often it also includes various levels and types of sociocultural phenotypes created by humans; and moreover, it is not the population of humans that is evolving but the sociocultural phenotypes that human have created and that they can re-create under selection pressures. And, these sociocultural phenotypes – from groups to organizations to institutional systems to whole societies and even inter-societal systems – can be viewed as additional layers of protection of the human genotype and, indeed, they are generally more important than the phenotype of the individual.
The evolution of human societies
Problems in Wilson’s evolutionary scenario
Wilson selectively cites literatures to develop his argument that human societies represent a form of eusociality. Indeed, some arguments supporting this assumption are obviously incorrect. For example, Wilson argues that ‘grandmother helpers,’ volunteers for social causes, homosexuals, monastic orders, and the like are the functional equivalent of castes of altruists who forgo reproduction in eusocial societies. Yet, grandmothers have been reproductive, or they would not be grandmothers, most volunteers still reproduce, homosexuals engage in a form of reproduction through adoption, slaves were encouraged to reproduce so as to provide more labor to exploit; and in general, then, humans seek to reproduce – if not, there would not be 7 billion of us polluting the world’s ecosystem. Thus, Wilson seems to be grasping at straws to make humans act like ants.
Further, Wilson is highly selective when he cites the very large literature on hominoid evolution to portray both great apes and humans’ hominin ancestors as hierarchical, as forming bands and even ‘gangs’ within larger communities, as driven by aggressive bio-programmers to make war on other communities. With these questionable conclusions, Wilson then argues that warfare was the force driving human evolution during the hunter-gatherer era. Yet, in contrast to Wilson’s claims, most studied hunter-gatherer populations have powerful norms against hierarchy in favor of equality, and moreover, it is well documented that warfare among nomadic hunter-gatherers is infrequent. As David Damas (1972: 33) writing on the Copper Eskimo phrased it, ‘Egalitarianism was the keynote of interpersonal relations,’ or as June Helm (1972: 80) writing on the character values of the Dogrib Indians emphasized, ‘The ethos ideal embraces generosity, emotional equanimity, egalitarianism, and the commitment to hard work and physical endurance as a . . . means toward the goal of group well-being.’ Indeed, in a review of the hunter-gatherer literature, Patrick Nolan and Gerhard Lenski (2018: 95) found that ‘Differences between individuals are so slight . . . that a number of observers have spoken of a kind of “primitive communism”.’ In addition, population densities for nomadic food collectors average only one person per square mile with population growth very low; and so, while conflict certainly exists, perpetual warfare (e.g., feuding, rioting, external or violent contact) for hunter-gatherers is non-existent, and warfare in general rare (Nolan and Lenski, 2018: 118; see also Leavitt, 1977).
Both the ethnological and archeological records also confirm that a higher incidence of warfare begins only with the horticultural era beginning about 10,000 years ago (Nolan and Lenski, 2018: 118). Indeed, sociologists as early as Herbert Spencer (1898 [1874–94]) or even earlier with Ibn Kaldun (1342–1406), and as contemporary as Peter Turchin (2003, 2006, 2011), have documented this finding for sedentary food producing populations (i.e., garden farming and beyond) but not for nomadic food collectors. Moreover, Wilson’s portrayal of early hominins (human or near human ancestors) is drawn from a narrow portrait of chimpanzees and, hence, is also problematic. For example, in the highly selective primate literature that he relies upon to make a case for early humans as warlike, great ape social organization is characterized as filled with intra- and inter-community conflict, which allowed him to surmise that the first hunter-gatherer populations also revealed these traits because they were inherited from the common hominoid ancestors of great apes and humans. Indeed, the fact that early human and human-like populations that lived contemporaneously –Neanderthal, Denisovans, and Homo naledi (as well as other fossils recently found in South Africa by Lee Berger and colleagues) – could survive for thousands of years at the same time suggests low population densities, coupled with geographical barriers, and little warfare.
Thus, Wilson’s imagery echoes Richard Ardrey’s well-known portrayal of early humans in his The Territorial Imperative as hierarchical and warlike – a portrayal that in 1966 was popularized but so obviously overdrawn that few social scientists embraced it. To be sure, Wilson’s focus on the frequency of human warfare is certainly in line with horticulturalists and all subsequent settled populations. But, horticultural societies did not evolve until very late in human evolution; indeed, for over 99% of humans’ time on earth (at the very least, 300,000 years), hunting and gathering was the only mode of subsistence for very low-density populations that did not manifest an overriding territorial imperative and that did not tolerate hierarchies or chronic conflict.
Wilson also appears to connect brain growth to the conflict among populations when, in fact, brain growth occurred, as he notes, very rapidly and in response to the movement of populations of later Homo erectus to ever more diverse ecologies where adaptations to new ecologies more than warfare increased selection for more intelligence. But such growth in the neocortex would be highly costly (in terms of calories and proteins needed to support it), and thus, selection pressures were no doubt intense, but we would argue warfare was not an important pressure because population densities were still very low and movement away from conflict would be an easier strategy than chronic warfare. The requirement for daily food collecting would also preclude frequent combat just as their nomadic way of life precluded the accumulation of possessions, including lethal weapons. Thus, there are so many problems with Wilson’s scenario that we feel it necessary to address the same issues as Wilson but with an evolutionary lens focused on the fossil, archaeological, and molecular records. We also need to use cladistics and network analysis along with data on hominoid evolution (i.e., ape and human) that can work as a corrective to Wilson’s scenario of ‘deep origins.’
Using cladistic analysis to discover the behavioral and organization propensities of the last common hominoid ancestor
Cladistic analysis in biology is a well-established methodology that allows us to look back in time by using contemporary great apes (i.e., chimpanzees, gorillas, and orangutans) as proxy representatives of the last common ancestors (LCA) that humans shared with the ancestors of contemporary great apes. Humans share an LCA of humans with each lineage of extant great apes: 5–6 million years ago (MYA) with chimpanzees, 8–9 MYA with gorillas, and 13–16 MYA with orangutans. Because humans are still so closely related to these lines of great apes – sharing 99% of our genes with chimpanzees, 98% with gorillas, and 96–97% with orangutans – a cladistic analysis can provide useful information by using their shared relational patterns, evolutionary novelties, organizational biases, and other systematic regularities to draw blueprints on the organization structures of their last common ancestors. Maryanski’s cladistic analysis (1987, 1992, 1993, 1995) of the great apes has been used in many articles and books (e.g., Maryanski, 2018; Maryanski and Turner, 1992; Turner and Maryanski, 2005, 2008; Turner et al., 2018) to fill in details about the behavioral and organizational patterns of those hominoids from which hominins and then humans evolved.
As noted above, in a cladistic analysis the endpoints of evolutionary lineages among closely related species such as the great apes are examined to determine the traits that they still have in common, with this commonality a good indicator of the traits of the last common ancestors that these clades inherited. Indeed, chimpanzees, on DNA grounds could be put in the genus Homo since chimps are more closely related to humans genetically than to either gorillas or orangutans. As Marie-Claire Kin and Allen Wilson lightheartedly phrased it (1978: 91), ‘the chimpanzee–human difference is far smaller than that between species within a genus of mice, frogs or flies.’ And, this is, of course, why Wilson focused on chimpanzees as his source of insight into the behavioral propensities of hominins (i.e., near or direct human ancestors) on the human clade. But a cladistic analysis reaches very different conclusions than Wilson does in his scenario on deep origins.
In performing such an analysis, Maryanski documents that the last common ancestor population of humans and great apes had very weak social ties to conspecifics and lacked permanent groups (aside from a mother and her dependent young), with the only stable structure being a community or home range within which members would move about. In addition to low-density networks and weak ties, a cladistics analysis also portrays the LCA as sexually promiscuous (precluding any paternal relations with offspring). After puberty, both female and male offspring emigrated from their mother never to return, thus curbing all extended kinship relations. The closest living great ape matching this hypothetical cladistic reconstruction of the last common hominoid to current great apes and humans is the present-day orangutan, a near solitary ape. The LCA population was, therefore, highly individualistic with very weak ties and a lack of any structural foundation for intergenerational kinship relations.
A discussion of why selection forces would favor such weak-tie communities is not possible here (but see Turner and Maryanski, 2005, 2008), but Wilson’s portrayal of humans’ hominin ancestors is certainly at odds once we look back to 10–16 million years ago and view great ape social organization through the ‘distant mirror’ provided by cladistics.
Today, while species of extant orangutans are still mostly arboreal, with individuals spending most days alone high in the rainforest canopies of Borneo and Sumatra, subspecies of present-day gorillas and chimpanzees are more socially inclined, spending considerable time on the forest floor. In the case of gorillas, their size and weigh by itself would justify a ground living lifestyle while chimpanzees, who are also large for an arboreal primate, still live in the forest canopy but spend time on the ground when traveling, food foraging, or simply maintaining a quiet co-presence.
The result was that gorillas developed propensities for a loose group formation around a lead silverback. Daily life in these fluid groups is relaxed and casual, even as they evidence rather social weak ties and interactions among adult members, who, aside from the lead silverback, are not obligated to stay in the group. Given that, at puberty, both sexes depart and that all adults can come and go as they please, the leadership and dominance of the alpha silverback is subtle and does not constitute a hierarchy of authority. And, while a gorilla group can be stable for a time, it manifests a shifting collection of individuals over longer stretches of time (Bradley et al., 2005; Harcourt and Stewart, 2007a, 2007b).
The ancestors of chimpanzees evolved more social structure and continuity because present-day chimpanzee males are lifetime locals in their community while females depart and are replaced by immigrating females from neighboring communities. After puberty, chimpanzee males are self-reliant nomads, but adult males sustain strong relations with their mothers by periodic visits. Brothers can also develop strong ties but often prefer to socialize with unrelated male friends. Fathers are unknown because incoming females are promiscuous, thus precluding extended kinship along the male line and, because of female transfer at puberty, along the female line as well.
Chimpanzee communities evidence ‘long-lived cultures that are passed on conservatively’ through many generations (Luncz and Boesch, 2014: 656). Up to 150 individuals move about independently within a solid block of forest that can range in size from 10 square miles in a rainforest habitat to more than 70 miles in a savanna-woodland habitat (Watts, 2002). Community members may not see each other for weeks, if not months; but in coming into co-presence, they recognize locals, remember past interactions, engage in greeting rituals, and also role take (theory of mind). The image that Wilson presents of chronic conflict among roving bands and gangs within chimpanzee communities is simply not accurate. Mothers with dependent offspring are reclusive and usually forage alone (without fear of roving gangs) while male chimpanzees wander about alone or join small, daily impromptu parties for a few minutes or hours. Rarely, if ever, does an entire community cluster in propinquity to constitute a group (Goodall, 1986; Luncz et al., 2012; Stumpf et al., 2009; and see a full listing of citations documenting these details in Maryanski, 2018; Turner and Maryanski 2005, 2008; Turner et al., 2018).
Chimpanzee males do, as Wilson emphasizes, monitor the boundaries of their community in ad hoc and episodic patrols; and they will injure or kill alien males who enter their community. But Wilson’s portrayal of frequent inter-community warfare is exaggerated. There are reports of conflict between communities, but this is not commonplace (see Boesch et al., 2008 for a more representative empirical review about inter-community conflict among chimpanzees).
Implications of the behavioral and organizational propensities of the last common ancestor
The portrayal outlined above is far different than Wilson’s discussion of chimpanzees, but the thrust of our truncated discussion is a much better indicator of what humans’ early hominin ancestors were like around 5–6 million years ago. In contrast to Wilson’s scenario, the most notable feature of hominins was the lack of stable group structures as hominins ventured from a secure forest habitat into open-country. Kinship networks were meager, structured hierarchies did not exist nor did stable adult groups. How did they protect vulnerable mothers with dependent young? And how can nomads with few tools and little structure organizing their lives mobilize for war? Indeed, the biggest source of conflict was finding an effective defense against high predation? The weak ties and individualist propensities of great apes were well suited for a large-bodied primate in arboreal and semi-arboreal forest niches, but as the forests began to recede with global cooling during the Pleistocene epoch (about 2.6 million years ago), some species of primates were forced to take up open-country habitats. The imperative driving great apes on the savanna was to get sufficiently organized to survive rather than to conquer.
How, then, did natural selection work around the lack of proclivities for group orientations and kin relations to make those species of hominins forced onto open-county more social, more kin oriented, and hence, more group oriented? Most, hominins like Australopithecus robustus did not survive in the long run. But somehow selection acted to fashion something like the universal nuclear family that could then be organized into nomadic hunter-gatherer bands for protection of mothers and young, for coordinated food collection, and for defense against predators rather than other hunting and gathering bands. Selection took an indirect route to making hominins more social and more organized at the group level.
The neurology of group and kin organization among hominins
In a number of works (e.g., Maryanski, 2018; Turner, 2000, 2002, 2007; Turner and Maryanski, 2008; Turner et al., 2018), we have outlined in more detail how selection got around the low sociality and unstable group formations of early hominins. For, on the predator-ridden African savanna, hominins had to organize into some type of stable groupings, or go extinct. Yet, selection can only act on distributions of existing traits among large, complex animals as hominins, since large random mutations would have harmful effects on the intricate complexity of hominin neurobiology after 25 million years of hominoid evolution (see Fisher 1930: 83; Stebbins, 1969: 105 on the unviable nature of large mutations). Our view is that it worked on the hominin brain as Wilson suggests, but not initially on the neocortex, which, as Wilson also recognizes, did not increase significantly until late Homo erectus 600,000 to 400,000 year ago. More cognitions do not promote bonding; instead, humans and hence ancestral hominins formed relations of solidarity in groups through the arousal of positive emotions, reinforced and intensified by ritual activities (Turner, 2000, 2007). We know this is the case because it is how humans today form and sustain relationships and groups.
Thus, selection worked first on the subcortical areas of the brain where emotions are generated in mammals, and this process probably began with early Australopithecines who may have ventured out for brief periods onto the savanna – as Wilson also suggests. Growth in the subcortex is not easily detected in fossilized cranial endocasts because of the low position of the subcortex below and inside the neocortex. But, selection began, we believe, by working on the already robust emotional capacities of primates to enhance the number, variety, intensity, and nuance of emotions, first by expanding the range of primary emotions (e.g., anger, fear, sadness, and happiness) that humans still share with chimpanzees and most other mammals; then, selection began to combine in some yet unknown ways (probably by configurations of simultaneous and/or phased activations of neurotransmitters, neuro-active hormones, hormones in general, and musculo-skeletal systems) the subcortex structures that generate emotions. And at the very end of this evolution of emotions came uniquely human emotions like shame and guilt that chimpanzees do not experience (Boehm, 2013; personal communication with Boehm on the chimpanzees at Gombe).
In this way, bonds of solidarity and attachment could be developed for male–female relations (and eventually more stable nuclear families) and for social relations in groups generating higher levels of solidarity. This route for enhancing capacities to form groups is, however, a double-edged benefit because three of the primary emotions are negative, and so enhancing negative emotions had to be balanced by even more enhancement of positive emotions and new types of consolidated and mixed emotions that mitigated the power of the negative by being blended with the positive. But, as is evident today, human emotions can easily turn to a darker side, a risk that had to be incurred to increase solidarity. And, no doubt, it is one of the reasons that hunter-gatherers try to avoid conflict, given the intense emotions that can be aroused (e.g., hatred, vengeance), and also why food collecting populations often have powerful normative constraints on individuals wanting to be recognized as better than others. Later hominins like Homo erectus surely learned that inequalities in power increase conflict and generate negative emotions and, ultimately, destroy the solidarity of groups.
Thus, unlike Wilson, we would argue that early hunter-gatherers relied upon positive emotions to forge the needed solidarities for a band-level coordination and an elementary division of labor by age and sex; and worked very hard at avoiding inequality and conflict, which could damage solidarity. And, for nearly all of humans’ time on earth, such was the predominate way of enhancing fitness among hominins without bio-programmers for nuclear families or high-solidarity groups.
Only when humans began to settle down, about 10,000 years ago, did conflict over territory become common among human populations. And, as the human population has become organized into mega-societies – also typified as ultra society (Turchin, 2016) – they were not eusocial. Order was maintained by the elaboration of new institutional systems like kinship, polity, law, religion, and economies based upon complex exchanges of resources. Hence, rather than importing a referential model best suited for understanding how tiny creatures like insects generate ultra- or mega-societies, we are far wiser to turn to sociology, which for 200 years has sought to explain just how large-bodied humans with big complex brains that require high levels of nutrients for growth and survival could generate mega-societies.
Why did mega-societies evolve among humans?
The first scientists who viewed themselves self-consciously as sociologists often addressed this problem in biological terms. Herbert Spencer in his The Principles of Biology (1898 [1864–67]) and, then, in his The Principles of Sociology (1898 [1874–94]) outlined a theory of evolution on what he termed superorganisms, that is, the internal organization of a population of organisms that included not only insects but human societies as well. Spencer argued that to sustain population growth requires the differentiation of members (into task specializations) and, in the case of human societies, also the differentiation of diverse types of social units organizing human activities in this division of labor. Thus, there is a fundamental relationship in the biotic universe of differentiation of structure (and culture for humans) and size of the population to be organized.
Like Wilson, but arguing almost 150 years earlier, Spencer emphasized that, indeed, warfare had been a driving force in the evolution of societies, especially after hunting and gathering. When societies go to war, the larger, more organized, and more technological advanced society is likely to win the war. And, unless the conquered population is simply exterminated, the emerging blending of societies increases the size of the now consolidated societies (even in exploitive ways such as colonization), leading to further differentiation of a population that, in turn, provides the structural base for further growth. Of course, other forces could be in play, such as technological innovation, but evolution from simple to more complex and, eventually, to mega-societies or very large superorganisms is also driven by warfare, although by 1900, Spencer had begun to argue that world-level market forces might now make warfare less of a driving force because, he always argued, warfare increases concentrations of power which increase inequalities that, in turn, make societies less stable due to internal conflicts. Thus, evolution by warfare will beget more internal and external conflict within and between societies.
Today, many scholars studying world-system dynamics have highly sophisticated models, with the bio-ecologist Peter Turchin (2016) among those most relevant to Wilson’s argument. For Turchin, the period from horticulture or pastoralism and through early and late agrarianism to industrialism was typified by warfare and, hence, has been a driving force of societal evolution as simple hunting and gathering societies disappeared. Thus, for many decades, sophisticated models and theories have been developed in sociology and other social sciences that better explain what Wilson seeks to do with a rather selective interpretation of primate data and his implausible applications of models describing eusocieties among insects.
Human nature and mega-societies
Recently, Turner and Maryanski proposed that ultra or mega-societies are possible because humans are an evolved ape. Mega-societies require that individuals move about joining diverse groups and interacting with others, often strangers, without stress; and they require an orientation to the larger social context beyond self and kin. Such a social universe with millions of individuals is alien to chimpanzees, even though they evolved in communities but, at most, with only about 150 individuals. But Homo sapiens share a recent common ancestor with chimpanzees who are highly individualistic, who evidence more weak than strong ties and who are not locked into genetically controlled kin networks. Moreover, chimpanzees possess virtually all the interpersonal capacities of humans, often even in the human measure (Maryanski, 2018; Turner et al., 2018). They are thus able to role-take or evidence what biologists term Theory of Mind; they can interact with individuals whom they have not seen for a while; they can see themselves as objects in their environment and can evaluative themselves from the perspective of others and the larger community; they can read the emotions of others and empathize with others; they can form short impromptu parties and yet, they can also engage in emotionally-charged collective rituals when some members congregate in an apparent celebration of their larger community (see Maryanski, 2018; Reynolds, 1965).
Without this suite of interpersonal capacities that allow for flexibility, fluidity, and mobility across diverse types of social structures, ultra or mega-societies would not be possible among a large animal like humans (see Turner, 2018a, 2018b). In fact, humans are probably better adapted to mega-societies than to earlier sociocultural formations revolving around horticulture and agrarian societies that limit freedom, mobility, and individualism. Humans evolved in hunting and gathering formations which are tiny compared to post-industrial societies, but unlike other societal formations since hunting and gathering (e.g., advanced horticultural and agrarian societies), post-industrial societies allow for more choice, freedom, mobility, and other opportunities that an evolved ape like humans clearly appreciates.
Comparative analysis in sociology
Comparative neuroanatomy
Over the last two decades, Turner has pursued a program in comparative neuroanatomy that compares the brains of great apes with those of humans. As great ape and human brains are essentially built on the same plan, any differences can be seen as evidence for the handiwork of selection over the last 6 million years. All the points made about emotions and interpersonal capacities are verified by this analysis. The subcortical portions of the human brain, where emotions are generated, are generally twice the size of those in great apes, controlling for body size, supporting the evolution of emotions (Turner, 2000; Turner et al., 2018). And such is even more the case with the septum, which is the area of the brain generating the pleasure associated with sex, which might be the ‘smoking gun’ of how selection enhanced emotional attachments among mating couples that could lead to more permanent conjugal arrangements and, eventually, nuclear families. Moreover, structural changes to the lateral amygdala (the center for fear and anger) also reveal selection for more sociality and awareness of social contexts (Barger et al., 2012, 2014). Also, the emphasis that Wilson puts on the neocortex is not misplaced, but the neocortex could not have grown without the prior enhancement of emotions since cognitions cannot be stored or remembered, or used in making decisions by the prefrontal cortex, unless they are first tagged with emotions (Damasio, 1994); and the more nuance, diverse, and complex are the emotions, then the more complex are memories and thinking. Thus, language and culture in the human measure could not have evolved in late hominins and early humans without a prior enhancement of emotion centers. Without emotion, the human neocortex would be a large, empty, energy/protein/calorie sapping warehouse, which would hardly promote fitness. Thus, Wilson’s discussion of the advantages of a large neocortex and of its rapid growth among late hominins is essentially correct but missing important neurological details, such as the necessity for a prior rewiring of the brain for emotions.
Comparative sociology
It is a curious fact that macro-societies are rare and restricted to two groups of remotely related organisms – the social insects and humans – who are separated by roughly 600 million years of divergent evolution. In 1992, Richard Machalek addressed this issue by comparing a variety of social forms and identifying the design features essential for an organism to evolve a macro-society with thousands and millions of individuals. More recently, he extended this analysis (see Machalek, 2018a) by comparing other dimensions among diverse types of superorganisms.
Machalek begins with the organismic constraints that hinder a population of organisms from evolving into a macro-society. For example, highly intelligent whales might qualify for macro-sociality except for the constraint of their very large bodies and the food requirement that they pose. Large-bodied humans also faced problems for organizing into macro-societies, until technologies for securing resources could be developed and until power could be consolidated to coordinate activities among the larger population.
In addition to body size constraints, there are ecological constraints revolving around food resources, competition for these resources, the density of other life forms in a habitat, rates of disease, and the like. Any of these can limit how large a population of any species can become. For example, larger populations of nomadic food collectors in woodlands, bushlands or savannas are not possible; only with migration to more plentiful habitats and more settled living could human populations increase at all, whereas social insects organized into societies of many millions in colony in most habitats probably consume fewer resources than a small band of hunter-gatherers. Moreover, ants and other insects are generally not in competition with larger life forms, as would be the case for early human populations.
Costs and benefits can pose either constraints or open up opportunities. The costs to large ant colonies (e.g., social parasitism) do not exceed the benefits of a complex division of labor, which allows ants to compensate for their small size in coordinated food gathering, defense, nesting, and reproduction. Among humans, costs for larger societies initially revolved around securing sufficient food and finding ways to organize more individuals, but as populations settled and grew, selection forces pushed such intelligent animals to find new modes of production (horticulture) and new forms of social organization revolving around expansion of kinship and eventually systems of power. So, animals that have large brains, language, and culture can invent new modes of social organization –structural and cultural differentiation and integration – to overcome costs, and increase benefits.
Finally, cost-benefit analysis suggests sociological constraints that inhibit or facilitate population growth. Machalek emphasizes that (a) individuals must be able to engage in impersonal cooperation (which excludes only cooperation in kinship, once populations begin to grow dramatically), (b) the labor must be divided among distinct categories of individuals, and (c) the division of labor must be integrated and coordinated. Ants do this by creating genetically controlled types of ants engaged in differentiated functions, whereas humans achieve the same result (not as smoothly functioning, of course) through elaboration of new kinds of social structures with distinctive statuses and roles, integrated by culture (norms, values, and beliefs), interdependencies (mediated by markets), and of course, consolidations of power.
In many ways, Machalek offers a better strategy than that pursued by Wilson for using models of insect societies to help understand mega-societies among humans. Insects and humans are among the very few organisms that create mega-societies because they have evolved mechanisms to overcome organismic, ecological, cost-benefit, and sociological constraints, but these mechanisms are very different. Insects are the product of Darwinian selection on very small bodied organisms with high rates of constant reproduction, thus allowing small mutations to remake both the organismic and grouping phenotypes of insects more rapidly than long-lived mammals like humans. Thus, rather than apply the insect model to human societies, it is better to look at what sociology has been doing, and especially what those engaged in evolutionary sociology have been doing with respect to analyzing the biological evolution of hominins, to the point where their larger brains, language, and culture would allow them to make rapid changes as they overcame all the constraints outlined by Machalek. Humans do not evidence eusociality, but a unique kind of ultra sociality that allows them to be flexible to create and adapt to a complex division of labor and varying modes of societal integration.
Conclusion
In 1830 Auguste Comte, the titular founder of sociology, argued that sociology was now emerging out of the foundation laid down by biology but, he suggested, sociology would in the future be able to inform biology. This seemed like a pretentious prediction by Comte, but after reading much of Wilson’s work over the last decades, plus the efforts of many in the biological sciences to explain sociocultural processes, we are now convinced that a reading of sociology is needed, if biologists are to continue making sociological pronouncements. Being a member of a society does not make one an expert of sociocultural dynamics, but often scholars outside of sociology think that mere membership does. The implicit presumption is that sociology has nothing to offer a biologist, and such might be the case for biology, per se; but when efforts are made to explain not just human evolution but the evolution of social structures and cultures created by a species with culture and capacities for agency, these efforts inevitably fall short, as does Wilsons here and elsewhere, because there appears to be a rather complete ignorance of what sociologists have been doing for the last 200 years. The result is that obviously brilliant scholars in biology come across as naïve and simplistic when doing sociology. We and many of our colleagues are fascinated with evolutionary processes and have done due diligence in reading evolutionary biology. In our respective cases, we have read extensively in neurology, paleontology, and primatology in order to make sure that when we address issues like the evolution of human emotions, the evolution of hominins and humans more generally and, in particular the evolution of human societies, we have read the relevant literatures in our own fields (sociology and anthropology) but also in other relevant fields. We could list pages of relevant scholarly works on the evolution of human societies that those entering sociology from other fields should read (starting with Lenski, 1966, 2005; Nolan and Lenski, 2018; Turner and Maryanski, 2008), but alas, we assume that the present practice of ignoring the social science devoted to the study of humans and societies more generally will continue. The result will be seemingly scholarly work that, in reality, is not as scholarly as it can be, and should be. Yet, there could be very useful and important collaborative work when giants in their field, like E.O. Wilson, would at least make contact with evolutionary sociologists who have written extensively on the evolution of humans and their sociocultural creations.
Footnotes
Funding
The authors received no financial support for the research, authorship, and/or publication of this article.