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Forthcoming in The Journal of Bioeconomics March 18 2013

Agriculture as a Major Evolutionary Transition to Human Ultrasociality

John GowdyRittenhouse Professor of Humanities and Social ScienceDepartments of Economics and Science & Technology StudiesRensselaer Polytechnic Institute, Troy New York, 12140 USA gowdyj@rpi.edu

Lisi KrallProfessor of Economics, Department of EconomicsState University of New York at CortlandCortland, New York 13045 USA lisi.krall@cortland.edu

AbstractThe adoption of agriculture was one of the most momentous transformations in human history. It set into motion forces that changed our species from living in small numbers within the confines of local ecosystems into one that is now changing the biophysical characteristics of the entire planet. We argue that this transformation can be understood as a leap to ultrasociality—a type of social organization rare in nature but wildly successful when it occurs. Several species of ants and termites made a similar leap in social organization and the broad characteristics of their societies are remarkably similar to post hunter-gatherer human societies. Ultrasocial species dominate the ecosystems they occupy in terms of sheer numbers and the scale of ecosystem exploitation. We argue that the drivers for the ultrasocial transition to agriculture are economic. These societies operate as superorganisms exhibiting an unparalleled degree of division of labor and an economic organization centered around surplus production. We suggest that the origin of human and insect agriculture is an example of parallel evolution driven by similar forces of multi-level selection. Only with the evolution of expansionist agriculturalist societies did humans join ants and termites in the social domination of Earth. Viewing agriculture as an ultrasocial transition offers insights not only about the origins of agriculture and its consequences, but also about the forces shaping the current demographic transition and the modern global socio-economic system.

The authors would like to thank the following for comments on an earlier draft: Jennifer Fewell, Marina Fischer-Kowalski, Dan Franks, Michael Ghiselin, Mason Inman, Clark Spencer Larsen, Doug Price, Peter Richerson, Simron Singh, Arild Vatn, David Sloan Wilson and Ulrich Witt. The comments of four anonymous reviewers were particularly useful. They are not, of course, responsible for the opinions expressed in this paper. We would like to acknowledge the contribution of discussions with the participants in a series of workshops funded by the National Evolutionary Synthesis Center (NESCent) at Duke University and a workshop “Evolution and Bioeconomics” at Ringberg Castle, Germany sponsored by the Max Planck Institute at Jena.

Keywords: Bioeconomics, Division of labor, Economies of scale, Evolutionary economics, Neolithic demographic transition, Origin of agriculture, Ultrasociality

JEL Codes: B52, N5, Q1, Q5

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Agriculture as a Major Transition to Human Ultrasociality

Both human civilization and the evolution of extreme insect superorganisms were attained by agriculture, a form of mutualistic symbiosis of animals with plants or fungi. Human agriculture, which originated about 10,000 years ago, was a major cultural transition that catapulted our species from a hunter-gatherer lifestyle to a technological and increasingly urban existence, accompanied by an enormous expansion of population. Humanity thereby turned itself into a geophysical force and began to alter the environment of the entire planetary surface…Approximately 50 to 60 million years before this momentous shift, some social insects had already made the evolutionary transition from a hunter-gatherer existence to agriculture…The most advanced agricultural insect societies, like their human counterparts, rose to ecological dominance. (Hölldobler and Wilson 2009, 408)

1 Introduction

The human impact on planet earth is so unique that the current era in earth history has

been dubbed the Anthropocene, the age of humans (Jones 2011, 133). Humans now dominate

basic atmospheric and biophysical processes on every part of the planet and are likely

influencing the course of evolution on earth for millennia to come (Barnosky et al. 2012; Gowdy

and McDaniel 1995; Steffen, Crutzen, and McNeill 2007). Anatomically modern humans lived

sustainably on the planet in small groups for something like 200,000 years. In only a few

thousand years after the widespread adoption of agriculture, quite suddenly in geological time,

the human population exploded from around 6 million to over 200 million by the beginning of

the Common Era (CE) 2000 years ago (Biraben 2003; Bocquet-Appel 2011; Cox et al. 2009).

Today the human population exceeds 7 billion and is projected to reach 9 billion by 2050.

Understanding the transition to agriculture may be a key to understanding the forces behind past

and current episodes of explosive population growth, the expansionary tendencies of many

human societies, and the extreme material inequality present in post hunter-gatherer societies.

We advance the thesis that the development of agriculture was a major bioeconomic change in

human evolution that drove the emergence of ultrasociality in our species in a manner closely

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analogous to its development in social insects. Complex human societies are not sui generis but

rather are products of co-evolutionary processes that are entirely consistent with the principles of

biological evolution, and especially recent work in multi-level selection (MLS) theory.

The characteristics of post-agricultural human societies are a consequence of a major

evolutionary transition to ultrasociality. Defining humans as “ultrasocial’ is controversial,

reflecting in part the lack of consensus among evolutionary biologists as to how to classify social

behavior. Following Campbell (1982, 1983), Richerson and Boyd (1998, 1999) and Turchin

(2010), among others, we use the term ultrasocial broadly to include humans as well as social

insects. Campbell (1982, 160) writes: “Ultrasociality refers to the most social of animal

organizations, with full time division of labor, specialists who gather no food but are fed by

others, effective sharing of information about sources of food and danger, self-sacrificial effort in

collective defense. This level has been achieved by ants, termites and humans in several scattered

archaic city-states.” E.O. Wilson (2013) uses the word eusocial to include humans and also

singles out ants and termites as being unique among insects in their numerical success:

Of the two dozen independent lines, just two within the insects—ants and termites—globally dominate invertebrates on the land. Although they are represented by fewer than 20 thousand of the million known living insect species, ants and termites compose more than half of the world’s insect body weight.

We define ultrasociality to include Campbell’s characteristics but we emphasize 1) its economic

orientation directed toward the production of agricultural surplus through the expropriation of a

larger part of the photosynthetic potential of land, and 2) a lock-in where non-producers and

producers form an articulate whole. Class divisions divide producers and appropriators. We

wish to stress the biophysical consequences including the enormous expansion of population and

biomass capture, and the emergence of the humans and other ultrasocial species as a geophysical

force (as in the opening quote by Hölldobler and Wilson, and as documented by E.O. Wilson in

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The Social Conquest of Earth). With ultrasociality the group functions as an organic whole with

individual activity organized to enhance the survival and growth of the superorganism.

We use “eusocial” to refer to social insects and a handful of other species as defined by

E.O. Wilson and Hölldobler: “…an evolutionarily advanced level of colonial existence, adult

members belong to two or more overlapping generations, care cooperatively for the young, and

are divided into reproductive and non-reproductive (or at least less reproductive) castes” (Wilson

and Hölldobler, 2005, 13368). Although E.O. Wilson (2012) also refers to humans as eusocial

because they share with ants and termites the characteristic of the “social domination of earth”,

we wish to avoid the controversies surrounding the genetics of eusocial insects (Abbot el al.

2011; Nowak, Tarnita and Wilson 2010).

The ultrasocial transition, like other major transformations, is not simply a matter of the

right ingredients (preadaptations). The sequence in which they are put together is also an

essential part of the evolutionary story. The chance that these ingredients and the right sequence

will create an ultrasocial species is unlikely but once it does the species may be very successful.

The creation of an ultrasocial species is a major transition—key phenomena that have only

happened a few times in evolutionary history but with momentous consequences (Maynard

Smith and Szathmáry 1995). For example, one major transition is the creation of RNA. The

“RNA World” hypothesis suggests that life itself originated when RNA evolved the ability to

organize self-replicating molecular systems (Ĉech 2011). Another higher level transition

occurred with the evolution of DNA from RNA precursors, possibly because of its enhanced

stability and information storage capacity. It is increasingly accepted that currently existing life

forms, from bacteria to multicellular organisms, are the result of such major evolutionary

transitions (Margulis 1970, 1998; Maynard Smith and Szathmáry 1995).

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We acknowledge that human sociality as it emerged in early hunter-gatherer cultures is

also a major evolutionary transition. Humans are unique in their manipulation of cultural

symbols, language, and their ability to cooperate with non-kin (Hill et al. 2009). We choose not

to characterize these developments as ultrasocial but rather preadaptations that paved the way for

ultrasociality. The gradual evolution of human culture, language, cognitive ability, and capacity

for abstract thought that took hold in Africa by about 200,000 years ago was a momentous and

unique occurrence.1 But agriculture, compared to the evolution of human hunter-gatherer culture

which apparently evolved in the genus Homo over a period of several million years (Pringle

2013), was a sudden, sharp, and decisive break in human history and it created a decidedly

different social dynamic with profound biophysical consequences: explosive population growth

and dominance of ecosystems. Ants, termites, and humans, after agriculture, achieved a level of

dominance of the earth’s ecosystems not seen in any other evolutionary lines.

A major theme in this paper is that humans are driven by the same evolutionary forces

that shape the characteristics of other species. Attributing agriculture, civilization, and human

domination of the planet to evolutionary processes common to ants and termites—not to the

superiority of the human intellect—may be hard to swallow. But the question of whether or not

humans have a special place in evolutionary history goes back to the debate between Darwin and

Paley (Gould 1990). The great successes of biology came about by moving from supernatural to

scientific explanations. We are sympathetic to Campbell’s (1974, 183) warning against the

presumption of human uniqueness: “I also join [Dobrowski] in distrusting that value-salvage

1There is clear evidence for complex symbolic culture from a site in South Africa dating 70,000 years ago (Henshilwood et al. 2002). Another site in South Africa (Pinnacle Point) contained red ochre pigments dating back to 164,000 years ago (Marean et al. 2007). Whether the development of human symbolic culture was gradual or a sudden revolution is still a matter of dispute but opinion is turning against the idea that human culture arrived in a sudden burst some 40,000 years ago. See the discussion in Knight (2010) and Pringle (2013).

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route which lies in worshipping man’s conscious experience, as by saving mind as separate from

body, or making body all mind, for these too commit the ancient sin of man worshipping himself

instead of the purposes and forces beyond any individual’s transient being.”

The dramatic demographic and socio-economic watershed that occurred in human society

after the widespread adoption of agriculture is termed the Neolithic Demographic Transition

(NDT). Although there were certainly population fluctuations before as the earth’s temperature

oscillated between warm and cold period, the magnitude of population growth during the NDT

was unprecedented in human history (Biraben 2003; Bocquet-Appel 2011, 560; Cox et al. 2009;

Gignouxa, Henn, and Mountain 2011). Biraben (2003) estimates that between 6000 BCE and

4000 BCE the Earth’s population increased from about 7 million to over 30 million and may

have reached 100 million by 2000 BCE. To appreciate the suddenness of the population increase

after agriculture, regional population numbers are more revealing than the total human

population. Complex agricultural societies developed first in the Huang Ho river basin in China

where the population grew from 0.8 to 20 million between 4000 and 2000 BCE and in the Indus

Valley where the population expanded from 0.7 million to 20 million during the same period—

4000 to 2000 BCE (Biraben 2003). Early population estimates are imprecise, but clearly

something momentous changed the relationship between the resource base and human

productive capacity.

Before the NDT, humans lived as hunter-gatherers in societies without permanent

settlements or agriculture. Compared to agricultural societies, it can generally be said that

hunter-gatherer societies were stable, egalitarian, and except for age and gender, largely

undifferentiated in terms of division of labor (Bird-David 1992; Boehm 1993; see the essays in

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Gowdy 1998).2 Importantly, the division of labor was not rigidly articulated around intensive

modes of production. Hunter-gatherers lived by extensively exploiting a wide variety of plants

and animals and living off of the indirect flow of solar energy. The purpose of economic activity

was subsistence, not the production of surplus (Woodburn 1982). By contrast, agriculturalists

collectively mobilized efforts toward intensively exploiting land to grow only a few crops (Price

and Bar-Yosef 2011). Unlike hunter-gatherers who lived off the flows of nature’s services3,

agriculturalists tapped into the stock of fertile soil and dominated the photosynthetic product of a

given place. Within a few thousand years after agriculture, a number of human societies were

characterized by (1) explosive population growth (2) the domination of ecosystems (3) relentless

exploitation of resources for the purpose of producing agricultural surplus (4) complex division

of labor, and (5) evolutionary success of the group at the expense of individual autonomy and

well-being.4

2 The 1968 conference “Man the Hunter” (Lee and DeVore 1968) overturned the conventional wisdom of the time that hunter-gather existence was “nasty, brutish and short.” Richard Lee, Marshall Sahlins and others made a convincing case that hunter-gatherers had ample leisure time and lifestyles that were “affluent” in the sense of having everything they needed for a satisfying life. In a predicable counter-reaction Kelly (1995) and others argued that hunting and gathering was a spectrum of lifestyles and questioned Lee’s calculations of working hours among the Ju/ ‘hoansi of the Kalahari and his description of their peaceful lifestyle. For a response and a discussion of the current state of hunter-gatherer studies see Lee and Daly (1999).

3This is not to deny that hunter-gatherers had dramatic impacts on local environments through hunting big game animals and by the use of fire to modify ecosystems (Murray 2003; Rule et al. 2012). The extent to which humans were responsible for megafaunal extinctions is still unclear. Current opinion leads toward the explanation that megafaunal extinction after the last ice age was a case of “coevolutionary disequilibrium” triggered by climate change and hunting pressure.

4The parallel evolution of human societies after agriculture is nothing short of astonishing. Wright (2004, 50-51) describes the results of parallel evolution from hunter-gatherers to civilization in Europe and the Americas: “What took place in the early 1500s was truly exceptional, something that had never happened before and never will again. Two cultural experiments, running in isolation for 15,000 years or more, at last came face to face. Amazingly, after all that time, each could recognize the other’s institutions. When Cortés landed in Mexico he found roads, canals, cities, palaces, schools, law courts, markets, irrigation works, kings, priests, temples, peasants, artisans, armies, astronomers, merchants, sports, theatre, art, music,

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In spite of the momentous importance of the agricultural revolution to population

dynamics, social organization, and the relationship between humans and the biosphere, and in

spite of the intensive scholarship on this transition and its importance in human history, basic

questions about the transition and its significance remain unanswered. “Why do hunters become

farmers? Why is agriculture so dynamic in changing human adaptation and behavior?” (Price

and Bar-Yosef 2011, S164) Why did people adopt agriculture even though it made the average

person worse off? What is the significance of this transition to human evolutionary history and

the future prospects of our species? We believe that our model of ultrasociality through group

selection answers these questions.

The debate among anthropologists surrounding the origin and rapid dominance of

agriculture has been structured around external and internal factors. The discussion has been

enhanced by evolutionary biology and in particular by emerging theories of the evolution of

human sociality through multi-level selection (Boehm 2011; Bowles 2012; Bowles and Choi

2012; Campbell 1983; Corning 2005; Haidt 2012; Nowak and Hightower 2011; Nowak,Tarnita

and , 2010; Richerson and Boyd 1999, 2005; Sober and Wilson 1998; Tomasello 2009; Turchin

2003; van den Bergh and Gowdy, 2009; Wilson 2012). We extend this discussion by arguing that

agriculture set in motion an ultrasocial transition in human evolution not unlike that which

occurred with agricultural ants and termites. When looked at in terms of economic and

biophysical consequences, the similarities in the organization of human societies and those of

advanced eusocial insects are so striking it is likely that similar evolutionary forces are at work.

We argue further that the ultrasocial transition was fundamentally economic. What happened

with humans in the transition to agriculture is not simply another variation on cooperation and

and books. High civilization, differing in detail but alike in essentials, had evolved independently on both sides of the earth.” As we argue throughout this paper, such results cannot be explained by chance. There must exist basic evolutionary forces driving human ultrasociality.

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group-level selection already present in hunter-gatherer societies, but instead an evolutionary

transition where society takes on the characteristics of a superorganism within which individuals

become cogs harnessed to further a higher-level goal which may or may not be in the interest of

individual well-being.

2 Ulrasociality as a major evolutionary transition

As with humans, the transition to ultrasociality (eusociality) by insects was a watershed

in their evolutionary history. Eusociality in insects is rare (13 or so species originally made the

transition) but hugely successful when it appears. This form of social organization is so

successful that it out-competes all rivals leading to a massive population explosion and

dominance of ecosystems in which it occurs. Indeed one of the most striking characteristic

patterns in eusocial insects is their domination in terms of sheer numbers compared to non-

eusocial species. In the Brazilian rainforest ants and termites comprise about 30 percent of the

entire animal biomass and 75% of insect biomass (Hölldobler and Wilson 2011, 6). Worldwide,

ants and termites comprise less than 2% of the earth’s insect species, but they account for 50% of

the earth’s insect biomass (Wilson 2013). Ants have roughly the same global biomass as humans

(Hölldobler and Wilson 2009, page 5, footnote 4).

Somewhere between 40 and 60 million years before humans practiced agriculture, two

unrelated eusocial lineages—ants and termites—independently began to grow fungi for food

(Mueller and Gerardo 2002). It has been suggested that, as with humans, complex ant societies

evolved from groups that practiced cooperative hunting. (Mueller et al. 2001; O’Donnell,

Kaspari, and Lattke 2005). One of the most complex insect societies, leafcutter (or attine) ants,

live in large cities of millions of individuals devoted to a single purpose—the cultivation of a

specific kind of fungus of the family Leucocoprineae. The entire ant society is organized toward

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the goal of maintaining or increasing the flow of fungus to feed the colony. Remarkable

similarities exist between ant and human agriculture (Diamond 1998; Mueller, Rehner and

Schultz 1998). Leafcutter ants (1) are monoculturalist, focusing on production of a specific kind

of fungi in their nests, (2) add manure to stimulate growth, (3) eliminate weeds mechanically and

with carefully manufactured antibodies, and (4) trade crops with other ant colonies, sometimes

with different species. Like humans, the insect farmers became dependent on cultivated crops for

food and developed carefully articulated task-partitioned societies cooperating in gigantic

agricultural enterprises. Agricultural life ultimately enabled the insect farmers to rise to major

ecological importance (Mueller and Gerardo 2002, 15247).

Central to the discussion of ultrasociality is the debate about MLS—whether natural

selection can work at more than one level (Wilson 2010; Wilson and Wilson 2007). Eusocial

insects exhibit high degrees of “altruism” as it is interpreted by biologists (the reduction of

individual reproductive potential for the sake of others). The question for evolutionary biologists

is how altruism survives under the forces of natural selection. There are two different

approaches to this question. One school of thought maintains that natural selection takes place at

multiple levels---the level of the individual and the level of the group and that total evolutionary

change is the sum of both. Wilson and Wilson (2007, 328) argue “…for the social group to

function as an adaptive unit, its members must do things for each other. Yet, these group-

advantageous behaviors seldom maximize relative fitness within the social group. The solution

according to Darwin is that natural selection takes place at more than one level of the biological

hierarchy.” E.O. Wilson (2012, 243) continues: “Nevertheless, an iron rule exists in genetic

social evolution. It is that selfish individuals beat altruistic individuals, while groups of altruists

beat groups of selfish individuals.” From a MLS perspective, an “invisible hand” guides both ant

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and human economies but it works by suppressing within-group competition, not encouraging it

(Frank 2011; Wilson and Gowdy, under review). The other school of thought maintains that all

selection is individual and that altruism can be explained through inclusive fitness and kin

selection—on closer inspection group selection is just a strategy to extend your own genes after

all (Dawkins 1976; Hamilton 1964). This approach brings selection back to the individual and

relieves evolutionary biologists from the complications and ambiguities associated with group

selection.5

D.S. Wilson (2010) argues that a consensus is emerging among evolutionary biologists

that natural selection may take place at many levels. Multilevel selection theory can explain the

struggle for relative fitness advantage within groups, as well as competition among groups that

might be detrimental to individuals. MLS can be extended downward to include competition and

cooperation among genes, or upward to include selection among groups (Wilson, Ostrom and

Cox 2013). Campbell (1982, 161) provides guidance as to how group selection works in human

societies:

Much hypothesized cultural evolution must achieve a kind of "group selection" precluded among vertebrates at the purely biological level and achieved by invertebrates only through caste sterility. The models of cultural evolution of Boyd & Richerson (1980) help here. Non-linear, multiple-social-parent transmission, with a majority amplifying effect, pushes face-to-face groups to internal unanimity in the absence of selection. This provides the raw material of ingroup homogeneity and group-to-group heterogeneity prerequisite for group selection. Such selection would come through differential group success, differential growth, conquest with cultural imposition, voluntary attraction of converts, imitation etc.

We argue below that competition between groups after agriculture favored those groups

that harnessed and directed a greater proportion of photosynthetic productivity. This engendered

5 For an illuminating discussion of the current state of the group selection controversy in biology see D.S. Wilson’s “Truth and Reconciliation for Group Selection” available at http://evolution.binghamton.edu/dswilson/wp-content/uploads/2010/01/Truth-and-Reconciliation.pdf

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an economically oriented and complex division of labor, a division between producers and

appropriators, and an imperative to increase output and capture increasing returns to group size

and the scale of production.6 This constituted a fundamental reordering of economic life.

Ultrasociality as a bioeconomic force emerges in the light of multi-level selection.

Societies already predisposed to cooperation and the division of labor reached a tipping point

with agriculture where the group takes on the character of a superorganism. The language hardly

exists to describe the ultrasocial superorganism. It is not a conscious organism but rather a blind

evolutionary process organized around a specific economic “goal”, whether the production of

fungus in leaf cutter ant colonies or the production of agricultural surplus in human agricultural

societies. The superorganism is directed by the unconscious process of natural selection, not by

conscious choice.

Among insects the steps to eusociality are described by Hölldobler and Wilson (2009),

Nowak, Tarnita and Wilson (2010) and Wilson (2012). The transition involves both necessary

ingredients and a unique sequence of events. Wilson (2012, 187) describes the process for

eusocial insects: (1) formation of groups (2) acquisition of pre-adaptations that form the

ingredients that reinforce eusociality (3) the appearance of mutations that reinforce group

persistence (4) emergent group traits that are reinforced through natural selection, and (5)

multilevel selection (group selection) that drives the evolution of superorganisms. Wilson and

Hölldobler (2005, 13368) lay out clearly the point in the evolution of insect colonies where they

move to the level of a superorganism. This point “comes very early…in particular when an

anatomically distinct worker caste first appears, hence when a colony can most meaningfully be

called a superorganism.”Most likely this occurs between steps 3 and 4. The key ingredient, an

6For example, more surplus supports larger groups in warfare. Those groups with institutions that led to cohesiveness (religion for example) could out compete others (Wilson 2002; Turchin 2003)

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anatomically distinct worker class, is reinforced in a positive feedback process by the increased

efficiency in production arising from the division of labor (Hölldobler and Wilson 2009, 47-48;

Beshers and Fewell 2001).

The claim that Homo sapiens is an ultrasocial species implies much more than the casual

observation that human society resembles a beehive or an ant colony. As with ants and termites,

ultrasociality in humans coincided with the adoption of agriculture as a primary means of

subsistence. The characteristics of hunter-gatherer societies were shaped by group selection--

within-group competition was suppressed because this favored the survival of the group. The

interest of the group was aligned with the interests of individuals within the group. But with

agriculture and production for surplus, the suppression of individuals went into overdrive. E.O.

Wilson tells us that the pull between individual and group selection (between selfish individuals

and altruists) “can never be complete; the balance of selection pressures cannot move to either

extreme. If individual selection were to dominate, societies would dissolve. If group selection

were to dominate, human groups would come to resemble ant colonies” (Wilson 2012, 243).

This view is controversial but it seems clear that with agriculture there was a profound shift in

the balance between the levels of selection, a tipping point was reached, and ultimately this shift

toward the dominance of group selection encouraged the formation of a human superorganism.

Humans did not become ants but the scale certainly tipped in that direction. With agriculture,

cooperation for “the good of the group”—favoring in some sense the lot of the average

individual within the group—became cooperation for “the good of the superorganism” in which

the well-being of individuals was secondary. In fact, when considering the formation of society

as an adaptive unit, and specifically in light of the transition to ultrasociality, the evolutionary

implications of the dichotomy between selfish individuals and altruists is not entirely clear cut.

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Aggressive individuals are clearly essential to the defense of property which itself is a necessary

part of the ultimate success of agricultural groups (Bowles and Choi, 2012). Presumably,

aggressive individuals would have the within group selection advantage in that they might get

more than their share of resources, yet they are inextricably and interdependently tied to their

group-social roles and understanding this interdependence is most important from the

perspective of social evolution (Witt and Schwesinger 2013). Altruistic individuals, who are

probably more likely to toil for the good of the group may not, in fact, be reproductively

disadvantaged by their efforts.

The large-scale agricultural societies that took hold some 8000 years ago came to

resemble eusocial insect colonies. Out of the many human groups that adopted agriculture, some

grew into agricultural civilizations that were hierarchical, differentiated into strikingly unequal

social classes based on a division of labor, and were aggressively expansionary. These societies

dominated the regions in which they were located. As well, the massive human population

increase and the ecological impacts from human productive activity were unlike anything

previously seen in human societies. Although there are obviously important differences between

eusocial insect species and humans, there are enough similarities to suggest that the agricultural

revolution may have been a transition not unlike that made by ants and termites.

We believe that the unprecedented evolutionary success of Homo sapiens after

agriculture was bioeconomic—the emergence of a superorganism designed around a narrow

economic purpose and in particular the production of economic surplus in agriculture. The

mechanisms for organizing ant and human superorganisms are different but the ultimate causes

are found in the same evolutionary dynamic, a dynamic reinforced by MLS and the force of

group selection and manifest in the rigid and attenuated division of labor around a specific

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material goal. Thus the emergence of ultrasociality must be understood as a bioeconomic

process; a co-evolution of biology and the economic production of the material existence of

society. With the organization of human society around the imperative of producing agricultural

surplus, a sort of economic “virtuous circle” emerged and reinforced itself and the system

became self-referential. This resulted in an ultrasocial transition with large scale economic and

ecological implications, including an explosion in population, domination of ecosystems, and

extensive and intensive resource exploitation. These biophysical results occurred in non-human

organisms—especially ants and termites—without human specific culture and consciousness.

In the transition to agriculture, it would appear that the play of selective pressures in

human society ushered in a new social and economic dynamic. This was not a matter of degree,

it was a matter of kind. In humans the navigation between genes and culture is complicated and

so too is the balance between levels of selection and the traits at play. In the end their

evolutionary play must speak to the outcome. In terms of their biophysical characteristics, human

agricultural societies and ant and termite colonies are strikingly similar.

Although the mechanisms are different for insect and human agricultural societies, the

evolutionary processes and their outcomes are similar. Acknowledging that human society

became a superorganism in the transition to agriculture runs counter to the current preoccupation

with individualism, selfishness and aggression and our bias in viewing the benefits of

civilization uncritically. And in terms of E.O. Wilson’s view that human evolution exhibits an

unresolved multilevel selection oscillation between the individual and the group, acknowledging

that humans are like ants means that this oscillation went decidedly in one direction. It is

possible that in the complex dynamic of multilevel selection, the evolution of complex societies

co-opted part of our humanity (our propensity for empathy, cooperation and altruism) and

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reduced it to something mechanistic and problematic. With the agricultural transition, individuals

began to function as cog-like parts to serve the perpetuation of the superorganism, not unlike the

ants that so fascinate us. We like to believe that we are different, that our power of rational

thought, imagination, creativity, and our penchant for cooperation keep us from this outcome but

a more critical look leads us to entertain the possibility that this isn’t necessarily so.

3 The agricultural transition to ultrasociality

Anthropologists have long studied the transition to agriculture yet this transition and its

meaning for humans remains enigmatic. Price and Bar-Yosef (2011, S168) summarize: “There is

as yet no single accepted theory for the origins of agriculture, rather, there is a series of ideas and

suggestions that do not quite resolve the questions.” There is every reason to believe humans

knew the principles of agriculture long before its widespread adoption (Cohen l977; Zvelebil and

Rowley-Conway l986). In fact it is inconceivable that there was not experimentation with

planting for a long stretch of hunter-gatherer history. It simply required that someone observed a

seed dropping to the ground, germinating, growing a plant, and producing more seeds. Flannery

(1968, quoted in Bowles and Choi 2012) observes: “We know of no human group on earth so

primitive that they are ignorant of the connection between plants and the seeds from which they

grow.” Knowledge was not the limiting factor in the adoption of agriculture.

Bettinger, Richerson and Boyd (2009) argue that the origin of agriculture can be

explained by a combination of external and internal factors. A period of unprecedented climate

stability beginning after the last glaciation can be seen as an external trigger that ushered in

agriculture. Evidence that the Holocene is unique in providing a long period of climate stability

provides a convincing case that climate was a key factor in this transition. Richerson, Boyd and

Bettinger (2001) demonstrate that the possibilities for agriculture were severely limited before

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the Holocene because of unpredictable climate fluctuations. Climate data indicates that prior to

the Holocene, changes in temperature as great as 8⁰C occurred over time spans as short as two

centuries (see Bowles and Choi, 2012, supporting on-line material, page 4).

Richerson, Boyd and Bettinger (2001) argue that the social institutions necessary for the

widespread adoption of agriculture probably took thousands of years to develop. This means

that, due to the constant abrupt interruptions because of recurring dramatic climate change before

the Holocene, any incipient attempt at large-scale agriculture was doomed to failure. Ice core and

pollen records indicate that centuries-scale abrupt climate events occurred regularly during the

Pleistocene and that it was not until the Holocene that a protracted period of warming occurred.

The abrupt climate events before the Holocene could have terminated incipient attempts at full-

fledged agriculture. Bettinger, Richerson and Boyd (2009, 628) also point out that late

Pleistocene plant productivity was low because of reduced CO2 levels (about 190 ppm compared

to 250 ppm at the beginning of the Holocene). Beerling (1999) estimates that the total amount of

stored organic land carbon was 33% to 60% lower in the late Pleistocene compared to the

Holocene.

Yet the argument against the climate change hypothesis—that several warming periods

occurred during the last 2 1/2 million years (the Pleistocene) but agriculture did not appear until

about 10,000 years ago—must be considered if only to note that warm and stable climate clearly

wasn’t sufficient for the transition if other characteristics of social organization and evolution

were not in place. Surely these must be part of the evolutionary story. As well climate change

might explain the success of agriculture but is not a sufficient explanation for the evolution of

agriculture from small scale farming to the overwhelming social and ecological force that it

became with the rise of imperialistic agricultural theocracies. In spite of the new stable climate of

18

the Holocene, agriculture did not arise in Australia, and in other places where it took hold—

Papua New Guinea for example—it did not lead to large scale civilizations (Diamond 1997,

Bowles and Choi 2012).

Other theories emphasizing external factors have focused on population pressure. Yet

population pressure does not fit neatly into the category of external factors especially given the

complex dialectic at work with population and agriculture. Binford (1968) and Cohen (1977)

invoked population pressure as the factor that pushed the shift to agriculture. In part, their

emphasis is supported by the fact that agriculture emerged at around the same time in numerous

parts of the world so it was not place dependent. Cohen (1977) tells us that population pressure

did not mean that carrying capacity was reached but rather that the type of food stuff people

could access shifts, and they had to settle for less desirable food and work harder to tap into what

they needed—therefore the push for agriculture. Yet population changes in the distant past are

empirically difficult to demonstrate and Price and Bar-Yosef (2011) point out that there is little

evidence for population pressure in the areas where agriculture first appeared. In fact, agriculture

appeared first in the most abundant natural environments (Diamond 1997). The population factor

is further complicated by the fact that agriculture resulted in a more sedentary life and shifted the

population dynamic by increasing body fat and total fertility rates. The gradual and marginal use

of cultivation might have altered population dynamics toward a positive feedback path

reinforcing the need for more agriculture. A larger population might have put pressure on wild

food sources and made hunting and gathering less productive. Changes in population size clearly

involve a dialectic process where cause and consequence are not easily disentangled.

The problem of explaining the transition to agriculture is further complicated by two

problems. First, there is unequivocal evidence that human health declined as a result of

19

agriculture and second, it is not clear, at least in its early stages, that agriculture was more

efficient than hunting and foraging. We take these two problems in turn. There is much

evidence that agriculture, compared to hunting and gathering, was detrimental for individual

humans as shown by the widespread decline in human health after its adoption (Larsen 2006).

Archeological data indicates that lifespans of people engaged in early agriculture were shorter,

they had a decline in stature and they suffered more debilitating diseases from leprosy to arthritis

to tooth decay than their hunter-gatherer counterparts (Cohen and Crane-Kramer 2007; Lambert

2009). Larsen (2006, 12) writes: “Although agriculture provided the economic basis for the rise

of states and development of civilizations, the change in diet and acquisition of food resulted in a

decline in quality of life for most human populations in the last 10,000 years.” An argument that

the transition to agriculture was ultrasocial is that individuals were worse off even though the

group flourished—individual well-being was sacrificed for the success of the “superorganism.”

Diamond (1987) has called the adoption of agriculture “the worst mistake in the history of the

human race” citing the “gross social and sexual inequality, the disease and despotism that curse

our existence.” One can argue about the positive and negative social aspects of the agricultural

way of life, but again the physical consequences, in terms of human health and longevity, are

clear—the average person was worse off compared to hunters and gatherers. Obviously this

leaves us to consider that once in motion it is a system with a life of its own irrespective of the

negative effect on individuals.

The fact that agriculture (certainly in its early stages) may not have been more productive

than farming is an enigma highlighted by Bowles. Bowles (2011) compared the productivity of

foragers and hunters and gatherers to early agriculturalists and found that the productivity of

labor (as measured by caloric return per hour of labor) was lower for early agriculturalists (where

20

farmers use hand tools) compared to hunter-gatherers. This leads Bowles to claim that “the one

case that should be the poster child for the better mousetrap theory turns out not to be. The

agricultural revolution 11 millennia ago did not occur because the first farmers were more

productive than the foragers they displaced.” Yet these societies somehow gained the

evolutionary advantage and in the context of human evolutionary history quickly displaced the

hunting and gathering way of life. Again, we wish to emphasize competition between groups as

well as the changing within-group dynamic that took hold with agriculture. Even though output

per unit of labor may be lower in larger post-agricultural societies, total output per group was

greater.

Bowles and Choi (2012) present an argument for the adoption of agriculture that places

“behavioral-institutional” factors front and center. We believe their argument dovetails with our

ultrasocial explanation. Their story goes something like this. Holocene climate conditions made

farming more favorable but it was also necessary to have property rights in order for farming to

outcompete foraging, especially in its early stages when farming was not as productive as

foraging as measured by caloric investment per unit of output produced. In fact, Bowles and

Choi hold that farming only made sense as an alternative to foraging if farmers were assured the

benefits of their efforts. Thus the rights to those benefits, property rights, were not derivative of

farming but were necessary for farming to be a competitive productive strategy with foraging.

Bowles and Choi (2012,10) make the claim that among foragers a tradition of property rights

was extant “when resources such as fishing sites or stands of wild grains were sufficiently

productive that hunter-gatherers could be substantially sedentary, both allowing the emergence

of ownership of homes, and warranting the subsequent demarcation and defense of these highly

productive non-residential spaces.” And it was this heritage from foragers that in the context of

21

the Holocene conditions gave early farmers an institutional advantage that was then culturally

disseminated. Again Bowles and Choi (2012, 3) state: “These property rights could have been a

template that was extended to cover cultivated crops and tended animals, allowing the

introduction of farming.” They claim that these rights would have minimized within group

conflicts among farming groups and given the advantage to the early farmers of inter-group

conflicts with foragers.

The work of Bowles and Choi is important in shifting the focus of the transition to

agriculture from external factors and purely technological arguments to internal (group)

dynamics although one can disagree with their property rights first hypothesis. We recognize that

production for surplus and the evolution of property rights may have been a co-evolutionary

process. But care must be taken not to transpose western bourgeois institutions and notions like

property rights to unknown and unknowable cultures. Again, we assert the importance of

understanding the complexities surrounding surplus production in the ultrasocial transition. Ants

and termites do not have human institutions and property rights yet their transition to agriculture

is broadly similar to the human transition. The Bowles and Choi approach certainly leads us to

think more fully about the complicated dynamic of MLS and the processes that form and

strengthen groups as adaptive units. Richerson and Boyd point out that the transition from the

beginning of agriculture to the development of state societies took around 2000 years. Thus a

dynamic that started out modestly and benignly locked early agriculturalists into a relentless

process that led to hierarchical state societies.

There is no question that agricultural surplus required the protection of property and set in

motion a dynamic of expansion; an autocatalytic process molded around a division of labor

derivative of the necessity for defense where the production of surplus became an imperative

22

rather than an accident of nature. Agriculture involved a more elaborate division of labor

integrating those engaged in agricultural production and those responsible for laying claim to the

output produced. In Agriculture, nonproductive members were integral to production and

production and distribution were more fully articulated. Judging from contemporary accounts

(Endicott, cited in Bowles and Choi 2012; Lee 1968), hunter-gatherer societies were

characterized by a sharp separation between production and distribution. These societies were

aggressively egalitarian with an emphasis on fairness in distribution.7 The greater articulation

between production and distribution set in motion with agriculture opened the possibility for

hierarchical organization to take hold, allocating the surplus according to power, not need

(Georgescu-Roegen 1977). It is counterintuitive yet crucial to understand that it is surplus, not

power, that drives the system. The tendency for this change can be seen in hunter-gatherer

societies. Woodburn (1982) distinguishes between “immediate return” and “delayed return”

hunter-gatherer societies. Immediate return societies with simple technologies are “aggressively

egalitarian” while delayed return societies with more complex technologies and the storage of

surplus show the beginnings of social hierarchies. Hunter-gatherer societies such as the Indians

of the northwest coast of North America, because of the abundance of natural resources

including spectacular seasonal salmon runs, even had chiefs and settled communities.

There is no question that agriculture, like other major transitions, represented an amalgam

of pre-conditions and an internal collection of “spring-loaded pre-adaptations” (Nowak, Tarnita

and Wilson 2010) that came together to reconfigure society as an ultrasocial group. Climate

stability for example, may be a precondition as would the existence of wild plants suitable for

cultivation but these are not sufficient to explain what was clearly a complex co-evolutionary

7 For a humorous account of leveling mechanisms in hunter-gatherer cultures see “Christmas in the Kalihari” in Lee 1993.

23

process, an altogether distinctive and different group dynamic. As with eusocial insects, the way

had to be paved for humans to reach the point of social evolution that enabled the development

of large agricultural civilizations.

Pre-adaptations had to be in place and then a tipping point was reached where it became

likely that large scale agricultural societies would emerge from small scale experimentation.

These pre-adaptations for humans would include—the campfire as a “nest” (E.O. Wilson 2012)

the beginnings of a “social brain” (Wexler 2006) and the predisposition to cooperation, a

division of labor, and language, among others. Human evolution is a product of both genes and

culture, dubbed “gene-culture coevolution” by Lumsden and Wilson (1981) and “biocultural

coevolution” by Jablonka and Lamb (2006), Richerson and Boyd (2005) and Wilson (2007). The

human capacity for open-ended behavioral and cultural change may be almost unique among

mammals but it is still the subject of evolutionary forces. Human genetic evolution and cultural

evolution have been influencing each other throughout the history of our species and our genes

are a product of our cultures, no less than our cultures area product of our genes (Wilson and

Gowdy 2013).

Culture itself is an artifact of the development of long term memory and the ability to

“construct scenarios and plan strategies,” as well as the capacity for cooperation, all likely

chiseled out of the interplay of multi-level selection. E.O. Wilson (2012, 224) argues that:

A group with members who could read intentions and cooperate among themselves while predicting the actions of competing groups, would have had enormous advantage over other less gifted. There was undoubtedly competition among group members, leading to natural selection of traits that gave advantage of one individual over another. But more important for a species entering new environments and competing with powerful rivals were unity and cooperation within the group.

In recent years convincing evidence has accumulated that the human brain is uniquely

designed for social interaction, and that extended parent-child and non-kin relationships ensure a

24

rich capacity for intimacy, trust and cooperation (Gowdy et al. 2003; Wexler 2006). Many

mammals are highly social animals with a variety of behavioral attributes that evolved to

facilitate social interaction, but humans seem to be unique in their degree of sociability. Two

related features of the human brain are particularly important to human sociality and to gene-

culture coevolution: brain plasticity and the existence of Von Economo neurons (Allman et al.

2005; Sherwood, Subiaul Zadiszki 2008). Von Economo neurons are almost unique to humans

and apparently evolved to enable people to make rapid decisions in social context.8 Our capacity

for culture that enabled agricultural society is a product of biocultural coevolution.We believe

there is an aspect of biocultural evolution broadly construed that should be explored more

carefully in explaining the transition to ultrasociality, namely, the detailed aspects of the material

reproduction of society---the bioeconomic dynamic.

4 The co-opting of cooperation

With the evolutionary preadaptations of culture, cooperation and sociality, the road was

paved for the evolution of the human superorganism long before the advent of settled agriculture.

But in order for the transition to be final this evolutionary trajectory had to reach a tipping point.

We are told by Wilson that the tipping point occurs in eusocial insects when “an anatomically

distinct worker caste first appears” and then selection pressures work to reinforce this phenotypic

expression. Thus the mark of advanced ultrasociality in insects is when the group becomes

entirely self-referential and allele plasticity gives rise to a specific phenotypic expression that

determines differential roles for group members. A lock-in (accommodated by this phenotypic

expression) reveals itself in the division of labor articulated around a common economic

purpose. When this occurs individuals within these groups can’t survive without performing

8The features of the social brain and other unique biological characteristics of humans may indicate that we developing phenotypical characteristics of eusociality (Foster and Ratnieks 2005; Shanley and Kirkwood 2001).

25

specific functions within the group (although these function may vary throughout an individual’s

lifespan). The phenotypic and age-specific division of labor in the ant colony is reinforced and

orchestrated through chemical signals.

What is the equivalent of the existence of a phenotypically distinct worker caste in human

society? We know that the mechanism in humans is not simply a matter of phenotypic

differentiation giving rise to an anatomically distinct worker caste. Yet the process in humans is

not that different than in eusocial insects, it only appears so because we tend to attribute all

human social features to our culture, institutions, and human intelligence, and dismiss complex

non-human behavior as instinct. The transition to agriculture must be understood by analyzing

the profound alteration in the mode of production, the attendant division of labor, and the

changing group dynamics that happened with agriculture. The group, as an adaptive unit,

changed so profoundly that the biophysical impact of humans on earth was altered. It is here that

cooperation found a different institutional, cultural and we would add evolutionary expression.

The point of no return, the human equivalent of a phenotypically distinct worker caste in insects,

was found here in an altered group dynamic which centered on the production of agricultural

surplus. The forces and dynamic of group level selection moved into overdrive as the positive

feedback loops and the autocatalytic tendencies of the group coalesced around hierarchy and the

imperative of surplus production.

The basic evolutionary pattern can be seen as a series of “punctuations” from cooperative

hunting and gathering of wild plants capable of living independently from their cultivators, to the

development of a symbiotic relationship within which neither the cultivated plants nor their

cultivators are capable of living independently of each other.9 Mobile hunters-gatherers moved

9Ant ultrasociality may have also begun with cooperative hunting and ended in an unbreakable symbiotic relationship with a fungus (Mueller et al. 1998; Schultz and Brady 2008).

26

through places that were prime locations for production of cereal grains and they had the time to

experiment with planting since their days were not filled with never-ending toil. Studies of

contemporary hunters and gatherers indicate our hunter-gatherer ancestors had ample leisure

time (Lee 1968; Sahlins 1968). As the climate warmed and became more stable during the

Holocene their experiments were increasingly productive in terms of yield per unit of land

though not necessarily in terms of caloric output per unit of labor. The decline in the caloric

return per hour of labor was more than offset by laborers working longer hours, tapping into the

stock of biomass created from the distant past, and reproducing more workers. At some point the

caloric output from semi-cultivated plants must have been sufficient to hold humans for longer

periods of time in one place and they were probably increasingly inclined to do so as the caloric

return from big game hunting receded along with the glaciers.

Transitioning from early sedentary agriculture to large scale agricultural civilizations

took several thousand years (Bowles 2011; Rindos 1984) but eventually humans began to live

almost entirely from agriculture and their day-to-day activity became reoriented and centered

around the maximization of agricultural output. This is a fundamental alteration of society in

that economic life becomes the focal point around which all else is derivative. This change

marked a fundamental reorientation of humans away from a diffused interchange with each other

and a diverse nonhuman world and toward a concentrated orientation around a human made

world—that of maximizing agricultural production. With the shift to ultrasociality, economic

activity changed fundamentally from production for livelihood to production for surplus. This is

in line with Bettinger, Richerson and Boyd’s (2009) observation that hunter gatherers were “time

minimizers” while agriculturalists were “energy maximizers.” In a similar vein V. Gordon

Childe (1936) distinguished between “food gatherers” and “food producers.”

27

One can imagine that the transition was reinforced because the ability of the individual to

make a living independently of specific others became extremely limited. During the initial

stages of agriculture, foraging and hunting did not cease but more sedentary life with crops

necessitated more extensive foraging and hunting from the home base. This increasingly reduced

its practicality as a way to augment the production of food. Intensive cultivation resulted in

more dense living arrangements and a sedentary life. The greater population density further

diminished the viability of hunting and gathering. Over time the knowledge necessary for

individual autonomy in food production was diminished.

Sedentary life created both the opportunity for accumulation (simply because people did

not have to carry their possessions) and the imperative for accumulation, especially of food.

Certainly in non-tropical climates, there were long periods when plants were not growing and

there was no harvest. Because harvests could vary dramatically from year to year people were

entirely dependent on food that had been stored from one harvest to another. Thus the ability to

maintain and access the larder became all important in the reproduction of society. Institutional

changes accommodated this imperative. It is not accidental that the worship of fertility

goddesses and hierarchical social structures took hold and the importance of maintaining the

food machine became the focus of society. Agricultural societies also signified a more

precarious hold on the sustenance of daily life for its members, and more was at stake for

everyone if the system failed, thereby reinforcing greater group dependency and loosening the

hold of kinship dependency.

The final manifestation of lock in and the tipping point must be understood in the way

cooperation was coopted for purposes of maximizing agricultural output. The result was greater

productive potential for the group and the ability to produce surplus. Cooperation was a

28

consistent trait in the human population and had given the groups that held this trait an

evolutionary advantage over groups that did not (Boehm 1997; D.S. Wilson 1997). But in settled

agriculture the fact that individuals had no other option to secure the material necessities of their

lives other than to participate in agricultural production placed cooperation at the disposal of

coercion. Nascent property relations also furthered the institutional structuring of cooperation in

ways that were hierarchical. Defense was essential to the success of groups engaged in

agriculture. Expansion became increasingly necessary to accommodate population growth, the

growth of increasing numbers of nonproductive individuals (those engaged in defense) and to

counteract the loss of soil fertility associated with the growth of annuals.

The division of labor in early civilizations became more extensive, rigid, and detailed

around the maximization of agricultural output. Work was fundamentally reorganized extending

both the productive and social division of labor. Agricultural output necessitated many jobs

associated with production—planting, cultivating, harvesting, storing, organizing and processing.

The majority of the population was engaged in agricultural activity, and for the vast majority of

those the conceptualization of work had likely been disaggregated from the execution of work

altering the meaning of work and simply making humans more one-dimensional.10 This is a

common theme about the labor process under the capitalist mode of production in the economics

literature but there is no reason to believe a similar dynamic did not play out in early agricultural

society. Harry Braverman tells us: “... the detailed division of labor subdivides humans, and the

subdivision of the individual, when carried on without regard to human capabilities and needs, is

a crime against the person and against humanity” (Braverman 1974, p 73). Adam Smith

10As conscious beings with long term memory and the capacity for planning and forethought, humans can disaggregate the conception of work from its execution and further rationalize the process of production and organize economic life based on the accumulated knowledge of the past.

29

similarly comments on the effects of a detailed division of labor: “They whose life is spent in

performing a few simple operations, of which the effects too are, perhaps always the same…

generally becomes as stupid and ignorant as it is possible for a human creature to become”

(Smith 1776).11 The complex division of labor subdivides and controls humans in the same way

that phenotypic expression subdivides and controls ants. From the perspective of human beings it

signifies an unfortunate structuring of cooperation but there is no question this detailed division

of labor enhanced agricultural production. As well, the production of surplus extended the social

division of labor further reinforcing the dependency of the individual on the group and the

focused purpose of maximizing agricultural output. As previously stated, it was necessary to

support an ever larger group of people engaged in defense and expansion.

Our argument is that the co-opting of cooperation culminated in an extended and rigid

social and detailed division of labor. This was the tipping point and the equivalent of phenotypic

variation not unlike that which occurred in eusocial insects when they made the transition into

superorganisms. E.O. Wilson tells us that “group selection must be exceptionally powerful to

relax the grip of individual level selection.” (Wilson 2012, 55) We argue that this is how the

group selection force became so powerful in humans and clearly gave these groups the

evolutionary advantage and an imperative to maximize the production of agricultural output.

Cooperation, a human preadaptation to ultrasociality, became structured in a rigid and

11A loss of intelligence associated with the increasing division of labor may also be present in eusocial insects. Riveros, Seid and Wcislo (2012) tested the association between brain size and sociality across 18 species of fungus growing ants and found that increased colony size was associated with decreased relative brain size. In a study of human brain size, Geary and Bailey (2009) found that between 1.9 million and 10,000 years ago, when population density was low human cranium increased in size, but when population density increased beyond a certain point cranium size decreased. Average human brain size has decreased significantly, about 10%, since the Upper Paleolithic.

30

hierarchical way where individuals were left with little choice about their role in the cooperative

enterprises of society and those cooperative enterprises were dominated by and pivoted around a

narrow collective economic purpose. Finally cooperation in a locked-in and coercive form did

not entail a reproductive disadvantage within the group based on the division of labor. In the

realm of reproduction, all members of the ultrasocial group were reproductively fit. And clearly

groups that practiced agriculture could out-compete foragers because of their enhanced

productivity.

The human social system was transformed into a self-organizing, self-referential entity

whose “goal,” indeed imperative, was to maximize agricultural output. Although the

mechanisms of transformation were different than they were for social insects the results were

remarkably similar. Clearly the dominance of group selection became manifest and the chimeric

nature of human behavior tipped in the direction of ultra-group selection.

The subjugation of individuals for the good of the superorganism creates a problem for

those who (including the authors of this paper) believe that the worst aspects of civilization are

due to aggression and selfishness and the best to cooperation and altruism (Nowak and

Hightower 2011; Pagel 2012). There appears to be the very real possibility that evolutionary

forces reorganized the traits of cooperation and altruism to create a human society with an eerie

likeness to advanced eusocial insects. The effect of this change on human beings created

civilization but it also altered the felt experience of human lives in ways rarely acknowledged.

The social and individual consequences of moving from small-scale egalitarian social groups to

large-scale highly stratified agricultural theocracies where work was harder,more regimented and

more repetitious, requiring less of the creative and executive capabilities of the human brain.

Individual freedom (self-actualization) was more limited, and the orientation of the individual to

31

the non-human world became economically centered (Braverman 1974; Krall and Gowdy 2012;

Lee 1968; Sahlins 1968; Shepard 1998; Witt 2005). With human ultrasociality each individual

was required to gauge the rhythm and meaning of each moment of her/his life in a regimented

and restrictive way around the needs of the superorganism, now in its more evolved form. For

those engaged in agricultural toil, life was diminished by this transition.

The transformation affected the felt experience of living, the ability of people to fully

mature, and the ecological relationship of humans to the natural world. The plants he/she ate, the

animals encountered, the mental horizon, his/her dwelling and human access to the non-human

world shifted. The experience of living became more rigidly mapped to a larger economic,

social and domestic structure. The transition to ultrasociality in humans is their domestication,

their taming, the point where the tensions of multilevel selection shift and group selection

dominates. The loss in this transition is not easily understood for those of us now encapsulated

in civilization and its web of domestication. But if we can imagine for a moment that hunters

and gatherers resided in an interconnected world where the force of “all that was not them”

surrounded them completely and some would say resonated in them since the human genome is

the sum of our evolutionary history, we get a small glimpse of how this transition might have

diminished the felt experience of life.12

And finally the ecological consequences of the human ultrasocial transition must be

considered. After agriculture, human societies (with perhaps two major exceptions—China and

Egypt) were characterized by overshoot and collapse (Diamond 2005; Ponting 2009; Tainter

1988; Turchin 2006) because they were locked into an indivisible system incapable of looking at

the long-term consequences of their production system. An altered ecological dynamic is one of

12An extensive discussion of this is found in Paul Shepard’s Nature and Madness (Shepard 1982).

32

the hallmarks of the ultrasocial transiton. Evolution does not contemplate or account for the

future. The inability of human cultures to alter behavior in the face of impending collapse is one

of the hallmarks of human history and perhaps the most pressing problem we face today

Diamond 2005, Wright 2004).

5. Conclusions and implications for future research

We argue above that conceptual models of the transition to agriculture should include an

understanding of the self-organizing principles of the evolution of ultrasociality. Viewing human

societies as ultrasocial opens up a new research area drawing on the evolutionary history of other

species. One implication of MLS is particularly important. Researchers typically look for clues

to human behavior in our closest relatives, the primates. But if we focus on group processes we

can perhaps learn more from species that may not be closely related genetically but whose social

organization is similar to our own.

Biological analogies have had limited appeal to social scientists and economists (Gowdy

et al. 2013). The application of evolutionary biology to explain human society and in particular

to justify the problematic aspects of capitalism has often been crude and inappropriate. The use

of social Darwinism to justify tremendous inequality and the applications of sociobiology to

justify individualistic behavior and the market economy as the natural order of society leave

those who are critical of this form of economic organization and its outcome with little recourse,

after all it is natural. Biological arguments, in the form of “survival of the fittest” analogies,

have been used extensively by economists to promote reactionary social policies (Friedman and

Friedman 1980; Hirshleifer 1977). Yet multilevel selection theory and the evolution of

ultrasociality open a new door to understand our present circumstances.

33

The parallel evolution of human societies and social insects lends support to MLS and in

particular to the importance of group level selection. Through the lens of MLS we are forced to

look at the evolution of complex societies in a new light. Human society underwent an

ultrasocial transition with agriculture where the force of group selection became more

pronounced. This transition marks the beginning of the centrality of economic life in human

social organization and a reordering of the group as an adaptive unit. What emerges with

agriculture is the imperative of surplus fortified with the rigidly defined and hierarchical division

of labor. The social system becomes self-referential around this economic imperative. Thus a

different and now central economic impulse directs the ecological and social dynamic of society.

Cooperation is coopted and the biophysical impact of humans on earth begins its destructive

journey.

We may attribute our success on the planet to superior intelligence, technology, culture, or

our capacity for moral judgments, but the fact that social insects have achieved the same general

results—arguably better results since their systems have been sustainable for tens of millions of

years—should give us pause. We are left with a stark reality. Agriculture ushered in a unique and

unprecedented era in human history. It made possible civilization, advanced technology and

science, and the incredible success of the human species in terms of sheer numbers. But the scale

of human activity now threatens to disrupt the life-support systems upon which our survival as a

species depends. It would appear that the human evolutionary leap to ultrasociality that began

with the transition to agriculture has gone down a path that brings us to our present problematic

world order and our precarious historical moment. We might consider that evolution is not

without its unsuccessful experiments. Creating a sustainable and equitable human society will

require understanding and re-directing some basic evolutionary self-organizing principles of

34

ultrasocial systems. But in order to do so we must consider that under certain evolutionary

conditions our instincts for caring and cooperation, which we consider some of the most

distinctive aspects of our humanity, were co-opted in the evolutionary jump to agriculture and

ultrasociality. As early agricultural civilizations replaced hunter gatherer cultures, individuals

were made expendable and individual lives diminished. The ecological balance between humans

and the rest of the world was forever altered. Ten thousand years later, the ultrasocial forces

harnessing the economics of accumulation and exploitation ushered in the Anthropocene, the age

of human domination of the earth’s biophysical processes.

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