Moffett elegantly expresses the need to understand what keeps societies together and what tears them apart. This need is very clear as several nation states are fighting over territories and arguing over who belongs to what society. Meeting this requires a clear, interdisciplinary understanding of what is a society. This is what the paper provides. I applaud the breadth of Moffett's effort as he presents information from insects to nation states. In trying to bridge between humans and other animals, he relies heavily on primate examples. I will follow this lead.

Moffett places considerable emphasis on territories, using the term 37 times. In the first sentence of the introduction, he states “I broadly address societies, in the sense of enduring territorial groups whose members recognize each other as belonging.” In his formal definition, he drops the term territory, and states a society “regulates access to a space or spaces it ultimately controls.” Thus, he uses the term territoriality in a very broad sense to include any area the society controls access over through either aggression or avoidance, allows for extensive spatial overlap, and mobile territories that simply involve groups defending the space they occupy at a specific moment and avoidance-based spacing, which has been well described for baboons. The definition even allows for cordial visits of non-members. What the definition does not allow for is a set of individuals coinhabiting a physical location to be considered societies; for example, primate matrilines with a social group, religious groups, and any diasporas living in a larger society. This definition is considerably more general than that traditionally used in ethology (Brown & Orians, Reference Brown and Orians1970) and differs from classical dictionary definitions. For example, the Oxford dictionary defines a territory as “an area that one person, group, animal, etc. considers as their own and defends against others.” The broader definition is valuable, but the repeated use of the term territoriality is potentially damaging as it emphasizes the defense of space which many readers will view in a traditional sense as a demarcated geographical area. Considering the current actions of human nation states this is undesirable.

In this commentary, I would like to focus on the regulation of access to space as it is central to distinguishing one society from another. However, boundaries can be blurry both with respect to space and time. Blurry spatial boundaries are well known as many species do not have areas of exclusive use. Gorillas and red colobus monkeys provide illustrative examples with respect to boundaries changing over time. Gorillas (Gorilla gorilla and G. beringei) are considered to be non-territorial due to the high degree of spatial overlap among groups, limited inter-group aggression, and large home ranges (Morrison, Dunn, Illera, Walsh, & Bermejo, Reference Morrison, Dunn, Illera, Walsh and Bermejo2020a), however there is avoidance-based spacing (Seiler & Robbins, Reference Seiler and Robbins2020) and aggression increases toward the group's core area (Morrison et al., Reference Morrison, Hirwa, Mucyo, Stoinski, Vecellio and Eckardt2020b). Thus, a single group would be considered a society based on Moffett's definition. However, groups that have previously split exhibit affiliative behavior when they encounter each other within the peripheral areas of their home ranges (Morrison et al., Reference Morrison, Hirwa, Mucyo, Stoinski, Vecellio and Eckardt2020b). This affiliation blurs the definition of a society, as the dividing groups are one society and gradually become two. Thus, at what point are they considered separate societies?

Similarly, while red colobus monkey groups (Piliocolobus tephrosceles) (Kalbitzer, Golooba, & Chapman, Reference Kalbitzer, Golooba, Chapman, Reyna-Hurtado, Melletti and Chapman2023) and would be considered by Moffett to be societies. A large group of red colobus that overlapped ranges with approximately 10 other groups and interacted aggressively with them gradually split into two groups (Chapman unpublished data). These new groups interacted amiable for the next eight years before starting to exhibit intergroup aggression. These amiable interactions involved grooming, play, mating, and extended visits (weeks) by members of the other group. The behavior of gorillas and red colobus could be viewed as the process of forming new societies; however, defining a realistic cut-off point, which would be similarly appropriate for human societies, would be extremely difficult.

Moffett considers nation states to be societies, yet the creation of the boundaries of these societies has little in common with the gradual division of a single primate group into two groups/societies. For example, many European borders were demarcated following World War II by the victors of the war (WorldEconomicForum, 2021) and the size and shapes of African states were largely determined by colonial power (Green, Reference Green2012). This has resulted in extensive informal cross-border trade, as borders have little relevance to local people (Golub, Reference Golub2015) and in Asia (Abd Hair Awang, Bakar, Abdullah, & Liu, Reference Abd Hair Awang, Bakar, Abdullah and Liu2013) and Africa (Titeca, Reference Titeca2020) and cross-border traders have family and friends on both sides of the border. I question whether understanding how the formation of new primate groups will help us understand the formation of nation states.

The black and white colobus monkeys (Colobus guereza) challenge our understanding of what is being defended. Groups have extensive home range overlap, exhibit intense inter-group aggression, and use loud calls to facilitate avoidance-based spacing (Harris & Chapman, Reference Harris and Chapman2007). I have seen an encounter where two groups were aggressively interacting and a female left her group, traveled to the neighboring group, mated, and returned. If the defense is not over space, food, or mating, what is its function? Similarly, one could ask what is the function of national borders? Possibly, the establishment of the European Union offers a good discussion point for understanding their function.

What I view as the beauty of Moffett's contribution is that it provides a starting point for debate over what are the differences between societies, how are societies created and fall apart, what makes societal dissolution peaceful or not, and so on. Moffett does not pay much attention to elements within societies (e.g., primate matrilines, religious groups). Yet, these elements show many of the characteristics of a society, thus understanding how these elements form, organize, disband, and will also be an exciting area for research that could shed light on the process operating at a societal level.