See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/304625948 Ultrasociality: When institutions make a difference Article in Behavioral and Brain Sciences · June 2016 DOI: 10.1017/S0140525X15001089 CITATIONS READS 0 5 3 authors, including: Petr Houdek Julie Novakova Vysoká škola ekonomická v Praze Charles University in Prague 11 PUBLICATIONS 5 CITATIONS 6 PUBLICATIONS 9 CITATIONS SEE PROFILE SEE PROFILE Available from: Petr Houdek Retrieved on: 20 September 2016 BEHAVIORAL AND BRAIN SCIENCES (2016), Page 1 of 60 doi:10.1017/S0140525X1500059X, e92 The economic origins of ultrasociality John Gowdy Department of Economics and Department of Science and Technology Studies, Rensselaer Polytechnic Institute, Troy, NY 12180 [email protected] http://www.economics.rpi.edu/pl/people/john-gowdy Lisi Krall Department of Economics, State University of New York (SUNY) at Cortland, Cortland, NY 13045 [email protected] Abstract: Ultrasociality refers to the social organization of a few species, including humans and some social insects, having a complex division of labor, city-states, and an almost exclusive dependence on agriculture for subsistence. We argue that the driving forces in the evolution of these ultrasocial societies were economic. With the agricultural transition, species could directly produce their own food and this was such a competitive advantage that those species now dominate the planet. Once underway, this transition was propelled by the selection of within-species groups that could best capture the advantages of (1) actively managing the inputs to food production, (2) a more complex division of labor, and (3) increasing returns to larger scale and larger group size. Together these factors reoriented productive life and radically altered the structure of these societies. Once agriculture began, populations expanded as these economic drivers opened up new opportunities for the exploitation of resources and the active management of inputs to food production. With intensified group-level competition, larger populations and intensive resource exploitation became competitive advantages, and the “social conquest of Earth” was underway. Ultrasocial species came to dominate the earth’s ecosystems. Ultrasociality also brought a loss of autonomy for individuals within the group. We argue that exploring the common causes and consequences of ultrasociality in humans and the social insects that adopted agriculture can provide fruitful insights into the evolution of complex human society. Keywords: agricultural transition; division of labor; major evolutionary transitions; multilevel selection; surplus production; totipotency; ultrasociality But the mere fact that organisms and societies evolve by various through similar selection pressures. We use the term ultra- selective mechanisms is not the whole (superorganism) story, for sociality to refer to these lineages, and we address the ques- it does not tell us the fundamental reasons that ants and termites tion of its origin through the fundamental question of live social rather than solitary lives. Those reasons are to be “ – evolutionary biology: Where did something come from found in economics in the science that concerns itself with and what were the selection pressures that favored its how resources are utilized and allocated. ” — Michael Ghiselin (2009, pp. 243–44) spread? (Blute 2010, p. 13). We follow Campbell (and Darwin) in insisting that the evolution of human ultrasocial- ity is a consequence of some of the same mechanistic (i.e., not consciously directed) evolutionary forces that govern 1. Introduction other species. Foley (2008, p. 164) calls the adoption of ag- riculture by ants and humans a case of “convergent selec- With the widespread adoption of agriculture some 10,000 tion.” In the struggle to survive, agricultural ants and years ago, human societies took on some important charac- agricultural humans face similar problems and selection teristics shared with social insects – ants and termites in – tends to favor similar solutions. We fully recognize that particular that also engage in the production of their the details of ultrasociality in humans play out in ways own food. These characteristics represented a sharp that are mediated by human intentionality and cultural break in the evolutionary history of these lineages and led norms. to two important outcomes: (1) Ecosystem domination as a product of a dramatic increase in population size and much more intensive resource exploitation; and (2) the sup- 1.1. What is ultrasociality? pression of individual autonomy as the group itself became the focus of economic organization. The evolution of agri- There is no general agreement in the use of the term ultra- culture in fungus-growing ants and termites, and in human sociality, partly due to the lack of consensus in the biolog- societies, is an example of convergent evolution – the inde- ical and social sciences in classifying social behavior. The pendent evolution of similar characteristics in species not many definitions of ultrasociality are conflicting, even closely related. In terms of genetics, ants, humans, and ter- among the same authors. For example, Campbell some- mites could hardly be more different. Yet, in all three line- times classifies ants, humans, and termites as ultrasocial ages similar patterns of economic organization emerge (Campbell 1982) but other times refers to ultrasociality as © Cambridge University Press 2016 0140-525X/16 1 http://journals.cambridge.org Downloaded: 01 Jul 2016 IP address: 195.113.56.251 Gowdy & Krall: The economic origins of ultrasociality large-scale cooperation among unrelated individuals tion.2 The extent of differentiation, collaboration, and co- (Campbell 1983; see also Turchin 2013). This seems to re- hesion of agricultural social species place them in a qualita- strict ultrasociality to humans, but remarkable examples of tively different category. This demarcation is, of course, non-kin altruistic behavior in non-human populations are somewhat arbitrary, and we recognize the antecedents of being documented, for example, the recent discovery of co- both managed agriculture and the specific characteristics operative brood raising by two different species of spiders of ultrasocial societies. We argue that the change from for- (Grinsted et al. 2012). E. O. Wilson (1975), although he aging wild plants and animals to the active management of does not use the term ultrasocial, considers humans to be agricultural crops was a particularly powerful impetus for one of the four pinnacles of social evolution, along with the human transition to ultrasociality.3 To clarify our use colonial invertebrates, social insects, and non-human of this term, by our classification leafcutter (attine) ants mammals. Richerson and Boyd (1998) use the term ultra- would be both eusocial and ultrasocial. Complex human so- social to describe humans after agriculture, but do not cieties with agriculture would be ultrasocial but not euso- include social insects in their definition. Following Wilson cial. Human hunter-gatherer bands are not ultrasocial, and Hölldobler (2005, p. 13368), we reserve the term euso- although the antecedents of ultrasociality are clearly cial to refer to social insects and a handful of other species present in these societies, as we discuss below. A striking having an advanced level of colonial existence and a sharp difference between insects and humans is the presence of division between sterile and reproductive castes.1 non-reproductive castes in the former.4 We recognize We begin with Campbell’s(1982, p. 160) definition of that the term ultrasocial is controversial, but we insist ultrasociality: that human and insect societies that practice managed ag- Ultrasociality refers to the most social of animal organizations, riculture are fundamentally different from the small-scale with full time division of labor, specialists who gather no food foraging societies from which they evolved. 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. 1.2. Agriculture and ultrasociality We confine our discussion of ultrasociality to human, ant, Mueller et al. (2005, p. 564) list the defining features of ag- and termite societies that actively manage food produc- riculture as: (1) habitual planting, (2) cultivation, (3) har- vesting, and (4) nutritional dependency on the crop (obligate in insects and effectively obligate in humans). The active cultivation of crops calls forth a similar configu- JOHN GOWDY is Professor of Economics and Pro- ration of production in dissimilar species. The production fessor of Science and Technology Studies at Rens- of crops is a physical process entailing similar kinds of eco- selaer Polytechnic Institute. He has authored or nomic efficiencies. The prime examples of agricultural coauthored 10 books and 180 articles, and his re- insect societies are the attine ants of the New World search areas include biodiversity preservation, tropics (comprising about 200 different species) and the climate change, and evolutionary models of the fungus-growing termites of the Old World tropics (com- economy and society. He was a Fulbright prising more than 300 species). All of the existing species Scholar at the Economic University of Vienna of fungus-growing ants and termites apparently arose