We question the need to explain the onset of agriculture by appealing to the second type of multilevel selection (MLS2). Unlike eusocial insect colonies, human societies do not exhibit key features of evolutionary individuals. If we avoid the mistake of equating Darwinian fitness with health and quality of life, the adoption of agriculture is almost certainly explicable in terms of individual-level selection and individual rationality.

It is uncontroversial that humans are extremely social, and that cultures have changed over time. But, the evidence shows that much of the social psychology underlying these phenomena (1) predates the agricultural transition, and (2) is not the result of group selection. Instead, this psychology appears intricately designed to capture social gains when possible in our complex ancestral social ecology.

Social insects show us very little about the evolution of complex human society. As more relevant literature demonstrates, ultrasociality is a cause rather than an effect of human social evolution.

The ultrasociality thesis proposes that the same “mechanistic evolutionary forces” may be at work in the evolution of insect eusociality and human ultrasociality in relation to agriculture. Wide variation in the reproductive division of labor among differing highly social phyla points to a resemblance of outcomes arising from very different selective environments and possibly different forces.

Gowdy & Krall neglect an important aspect of human societies, which is that class systems are not genetically based but rather based on threat. In turn, threat presupposes intentionality, so seems to distinguish human from arthropod ultrasociality. Class systems, in turn, seem to explain the distinctive mixture of persistence and instability characteristic of human, but not arthropod, ultrasocial populations.

The evolution of ultrasociality in humans may have involved the evolutionarily significant mechanisms that govern family social structure in many animal species. Adverse effects of ultrasociality in humans may be mediated by maladaptive effects of modern civilization on family groups, as many of the effects on both families and societies are especially severe in dense populations made possible by agriculture.

Gowdy & Krall describe the development of ultrasociality in terms of economics and the division of labor. We propose that the division of cognitive labor allows humans to behave in an ultrasocial manner without the radical evolutionary changes that are experienced by other species, suggesting that species may traverse different paths to achieve ultrasociality.

To counter human ultrasociality, alternative communities can arise (ongoing), and, unlike insects, lower echelons can unite and rebel. Examples include movements such as: “Black Lives Matter,” “Fight for $15,” “Occupy,” and the “Village Movement.” To strengthen ultrasociality, a surplus bottom echelon can be reduced: for example, by means such as imprisoning Blacks, deporting immigrants, wars, and the Holocaust. Alternatively, a new structure could be created, for example, ISIL (even more ultrasocial?).

Gowdy & Krall (G&K) essentially recapitulate Malthus's classic argument for ecological pessimism in modern biological dress. Their reasoning also reproduces Malthus's blindness to the implications of technological innovation. Agriculture might have suppressed human individualism as G&K insist, but technology has tended to foster it. This complicates human ecological prospects in a non-Malthusian way, and it might additionally provide the resources for deliverance from disaster.

We question the sequence of evolutionary transitions leading to ultrasociality in humans proposed by Gowdy & Krall. Evidence indicates that families are, and likely always have been, the primary productive units in human agricultural economies, suggesting that genetic relatedness is key to understanding when the suppression of individual autonomy to the benefit of subsistence groups, that is, extended families, evolved.

The evidence Gowdy & Krall (G&K) provide is more consistent with a biological markets explanation of human ultrasociality than a group selection explanation. Specifically, large-scale societies provide a better biological market for cooperation than do small-scale societies, allowing individuals to increase their fitness. Importantly, many of the quality-of-life costs G&K discuss (e.g., patriarchy) are not fitness costs.

We agree that human economic structures can be informed by comparative analyses; however, we do not agree with several of Gowdy & Krall's specific assertions, which may hinder the generative potential of their model. We discuss these limitations from both biological and economic perspectives, and offer an alternative explanation for the expression of human economic behaviors based on individual optimization strategies.

The study of cultural complexity is entwined with ancestors myths from contemporary cosmology. Gowdy & Krall expose this mythology by arguing that complexity emerged from economic changes following cultivation. There is more, however, to the development of cultural complexity. Complexity can emerge from abundant energy or from addressing societal problems, which compels still further energy production.

The target article is built on an analogy between humans and ultrasocial insects. We argue that there are many important limitations to the analogy that make any possible inferences from the analogy questionable. We demonstrate the issue using an example of the difference between a loss of autonomy in humans and in social insects.

Given the highly social nature of the human emotion system, it is likely that it subserved the evolution of ultrasociality. We review how the experience and functions of human emotions enable social processes that promote ultrasociality (e.g., cooperation). We also point out that emotion may represent one route to redress one of the negative consequences of ultrasociality: ecosystem domination.

The target article is a major step toward integrating the biological and human-related sciences. It is highly relevant to economics and public policy formulation in the real world, in addition to its basic scientific import. My commentary covers a number of points, including avoiding an excessively narrow focus on agriculture, the importance of multilevel selection and complex systems theory, and utopic versus dystopic scenarios for the future.

We appreciate the depth and breadth of comments we received. They reflect the interdisciplinary challenge of our inquiry and reassured us of its broad interest. We believe that our target article and the criticisms, elaborations, and extensions of the commentators can be an important contribution to establishing human ultrasociality as a new field of social science inquiry. A few of the commentators questioned our definition of ultrasociality, and we begin our response with an elaboration of that definition and a defense of our argument that human ultrasociality began with agriculture. We then respond to the second major area of controversy, namely, our use of group selection to explain the economic drivers behind the agricultural transition. We then focus on the issue of human intentionality raised by the phenomenon of collective intelligence. The intriguing question is to what extent can an entire culture change its own destiny? We then address the issue of the division of labor raised by a number of commentators. The complex division of labor was both a driver and a defining characteristic of ultrasociality, even though it was present in simpler forms in earlier societies. The remaining issues addressed include energy and complexity, expansion and sustainability, and the accelerating evolution of human ultrasociality. These were raised by only a few commentators, but their importance warrants further elaboration.

Neural reuse is a form of neuroplasticity whereby neural elements originally developed for one purpose are put to multiple uses. A diverse behavioral repertoire is achieved by means of the creation of multiple, nested, and overlapping neural coalitions, in which each neural element is a member of multiple different coalitions and cooperates with a different set of partners at different times. Neural reuse has profound implications for how we think about our continuity with other species, for how we understand the similarities and differences between psychological processes, and for how best to pursue a unified science of the mind. After Phrenology: Neural Reuse and the Interactive Brain (2014; henceforth After Phrenology in this Précis) surveys the terrain and advocates for a series of reforms in psychology and cognitive neuroscience. The book argues that, among other things, we should capture brain function in a multidimensional manner, develop a new, action-oriented vocabulary for psychology, and recognize that higher-order cognitive processes are built from complex configurations of already evolved circuitry.

Anderson (2014) uses an impressive, consolidating review of the literature to argue for major changes in cognitive science. Arguably, however, much of what he proposes is not particularly new. He also neglects important predictive coding approaches that call his perspective of the brain into question, and his misconstrual of evolutionary psychology devalues an influential paradigm that promises to complement his own.

The evidence that Anderson (2014) marshals in support of his theory of neural reuse is persuasive. However, his theoretical framework currently lacks a developmental dimension. We argue that an account of the fundamental aspects of developmental change, as well as the temporal context within which change occurs, would greatly enhance Anderson's theory.