The discussion of the role of agriculture in the evolution of ultrasocial behavior in both social insects and humans is focused on cultivation of crops such as fungi (ants and termites) and cereals (humans). However, agriculture also involves domestication of animals. Animal husbandry plays a very important role in many human and ant societies. Numerous ants rear “cattle” (such as aphids, coccids [i.e., scale insects], and treehoppers) and use their secretions as an important source of food. They construct shelters for their “cattle,” transport them, and protect them against predators. Like humans, ants also sometimes eat their “cattle” to obtain proteinic food (Hölldobler & Wilson Reference Hölldobler and Wilson1990).
G&K discuss food storage largely in terms of storage of produced surplus, although they mention the evidence for pre-domestication granaries in the Jordan Valley (Kuijt & Finlayson Reference Kuijt and Finlayson2009). There exists further evidence showing that food and/or water storage preceded the advent of agriculture in both humans and ants. Bushmen hunter-gatherers from Kalahari store water in emptied ostrich eggshells, and the tradition of using such containers is at least 60,000 years old (Texier et al. Reference Texier, Porraz, Parkington, Rigaud, Poggenpoel, Miller, Tribolo, Cartwright, Coudenneau, Klein, Steele and Verna2010). Harvester ants do not culture plants, but forage for seeds and store them in underground granaries. Honeypot ants may act as “living larders” and store liquid food in their distended abdomens (Hölldobler & Wilson Reference Hölldobler and Wilson1990).
G&K extensively discuss morphological adaptations of ants to various functions fulfilled by them within the framework of the division of labor among the members of the same colony. Yet, in the vast majority of ant species (including some ultrasocial ones) workers are monomorphic and worker behavioral specialization is based on other factors than morphology – in particular on worker age and/or degree of physiological maturation (the so-called temporal polyethism) (Hölldobler & Wilson Reference Hölldobler and Wilson1990; Reference Hölldobler and Wilson2009). Interestingly, temporal polyethism also seems to play an important role in the organization of human societies. In the traditional Brahmin culture of India, each man had to pass through four successive life stages: Brahmacharya (student), Grihastha (married householder), Vanaprastha (hermit), and Sannyasa (wandering ascetic). Similarly, the Vestals of the ancient Rome had to pass through three stages (training, service, and training others) during their 30-year service at the Temple of Vesta.
According to G&K, ultrasocial ants and humans dominate the earth's ecosystems. However, raiding and domination inflicted by raiding robbers are known not only in humans, but also in ants. Army ants, the best known nomadic ants employing group raiding, form huge societies characterized by sophisticated caste polyethism (division of labor based on morphological differences) (Hölldobler & Wilson Reference Hölldobler and Wilson1990). This implies that among the ants neither dominance in the ecosystem, nor advanced division of labor are strictly correlated with the ultrasociality.
G&K also lay stress on the limited longevity of ant workers. However, ant workers may live up to several years (Godzińska et al. Reference Godzińska, Szczuka and Korczyńska1999; Hölldobler & Wilson Reference Hölldobler and Wilson1990), and they play an important role in the cultural transfer of information between older and younger nestmates. It's hard to disagree with G&K's statement that “one cannot reasonably argue that insects have culture in the way that humans do” (sect. 2.1, para. 2). However, cultural transfer of information seems to exist also in ants. According to Rosengren and his colleagues (Rosengren Reference Rosengren1977; Rosengren & Fortelius Reference Rosengren and Fortelius1986), red wood ants of the genus Formica form the so-called topographic traditions. The information about the localization of aggregations of aphids is stored throughout the winter in the brains of older workers. In the spring these “veterans” transmit that information to unexperienced young foragers, the “novices,” by guiding them to the places where the aphids will briefly appear.
G&K state that, in contrast to humans, the social life of insects is based on the reproductive division of labor (sterile and/or less fertile individuals take care of the offspring of more fertile ones), although non-reproducing individuals (namely, post-menopausal grandmothers) exist in human societies, too. However, these non-reproducing society members may also include young females (e.g., virgin priests and warriors, but also prostitutes, who may bear children but refuse to rear them) and males (e.g., eunuchs, celibate priests, unmarried slaves, servants, and soldiers). Ultrasociality and social organization based on city-states certainly promoted the role of these non-reproducing individuals in the overall organization of human societies, enhancing the similarities between advanced human and insect societies.
In section 2.3 (para. 4), G&K mention Smaldino's (Reference Smaldino2014) statement that “an ant colony represents the extended phenotype of the queen.” However, they do not point out that such a view is oversimplified, as ant colonies are often polygynous (contain numerous queens). Consequently, relatedness between nestmates is frequently very low, sometimes even close to zero. Nevertheless, ants from such colonies can successfully cooperate (Hamilton Reference Hamilton, Ito, Brown and Kikkawa1987). However, conversely to what was stated by G&K in section 5.1 (para. 4), important within-group conflicts do exist also in insect societies. Members of the same colony compete mostly for the access to reproduction (Heinze et al. Reference Heinze, Hölldobler and Peeters1994; Hölldobler & Wilson Reference Hölldobler and Wilson1990; Reference Hölldobler and Wilson2009). Ant workers are not completely sterile: they may reproduce by parthenogenesis and produce males from unfertilized haploid eggs and sometimes also females from unfertilized diploid eggs (Hölldobler & Wilson Reference Hölldobler and Wilson1990). Various forms of worker resistance against reproductive dominance of the queen lend support to the statement of G&K that “resistance has no analytical equivalent in ants (except perhaps attempts by workers to reproduce)” (sect. 6, para. 4). However, ant resistance may take yet other forms. In particular, enslaved ants may successfully rebel against their social parasites. Various forms of slave resistance include physical aggression against adult slave-makers, devouring of slave-maker's brood, attempts to reproduce in the slave-maker's colony, and slave emancipation (Czechowski & Godzińska Reference Czechowski and Godzińska2015).
In conclusion, all of these examples of not strictly adequate statements demonstrate that ant and human social behavior should be compared in a painstakingly careful way in order to draw entirely valid parallels.
The discussion of the role of agriculture in the evolution of ultrasocial behavior in both social insects and humans is focused on cultivation of crops such as fungi (ants and termites) and cereals (humans). However, agriculture also involves domestication of animals. Animal husbandry plays a very important role in many human and ant societies. Numerous ants rear “cattle” (such as aphids, coccids [i.e., scale insects], and treehoppers) and use their secretions as an important source of food. They construct shelters for their “cattle,” transport them, and protect them against predators. Like humans, ants also sometimes eat their “cattle” to obtain proteinic food (Hölldobler & Wilson Reference Hölldobler and Wilson1990).
G&K discuss food storage largely in terms of storage of produced surplus, although they mention the evidence for pre-domestication granaries in the Jordan Valley (Kuijt & Finlayson Reference Kuijt and Finlayson2009). There exists further evidence showing that food and/or water storage preceded the advent of agriculture in both humans and ants. Bushmen hunter-gatherers from Kalahari store water in emptied ostrich eggshells, and the tradition of using such containers is at least 60,000 years old (Texier et al. Reference Texier, Porraz, Parkington, Rigaud, Poggenpoel, Miller, Tribolo, Cartwright, Coudenneau, Klein, Steele and Verna2010). Harvester ants do not culture plants, but forage for seeds and store them in underground granaries. Honeypot ants may act as “living larders” and store liquid food in their distended abdomens (Hölldobler & Wilson Reference Hölldobler and Wilson1990).
G&K extensively discuss morphological adaptations of ants to various functions fulfilled by them within the framework of the division of labor among the members of the same colony. Yet, in the vast majority of ant species (including some ultrasocial ones) workers are monomorphic and worker behavioral specialization is based on other factors than morphology – in particular on worker age and/or degree of physiological maturation (the so-called temporal polyethism) (Hölldobler & Wilson Reference Hölldobler and Wilson1990; Reference Hölldobler and Wilson2009). Interestingly, temporal polyethism also seems to play an important role in the organization of human societies. In the traditional Brahmin culture of India, each man had to pass through four successive life stages: Brahmacharya (student), Grihastha (married householder), Vanaprastha (hermit), and Sannyasa (wandering ascetic). Similarly, the Vestals of the ancient Rome had to pass through three stages (training, service, and training others) during their 30-year service at the Temple of Vesta.
According to G&K, ultrasocial ants and humans dominate the earth's ecosystems. However, raiding and domination inflicted by raiding robbers are known not only in humans, but also in ants. Army ants, the best known nomadic ants employing group raiding, form huge societies characterized by sophisticated caste polyethism (division of labor based on morphological differences) (Hölldobler & Wilson Reference Hölldobler and Wilson1990). This implies that among the ants neither dominance in the ecosystem, nor advanced division of labor are strictly correlated with the ultrasociality.
G&K also lay stress on the limited longevity of ant workers. However, ant workers may live up to several years (Godzińska et al. Reference Godzińska, Szczuka and Korczyńska1999; Hölldobler & Wilson Reference Hölldobler and Wilson1990), and they play an important role in the cultural transfer of information between older and younger nestmates. It's hard to disagree with G&K's statement that “one cannot reasonably argue that insects have culture in the way that humans do” (sect. 2.1, para. 2). However, cultural transfer of information seems to exist also in ants. According to Rosengren and his colleagues (Rosengren Reference Rosengren1977; Rosengren & Fortelius Reference Rosengren and Fortelius1986), red wood ants of the genus Formica form the so-called topographic traditions. The information about the localization of aggregations of aphids is stored throughout the winter in the brains of older workers. In the spring these “veterans” transmit that information to unexperienced young foragers, the “novices,” by guiding them to the places where the aphids will briefly appear.
G&K state that, in contrast to humans, the social life of insects is based on the reproductive division of labor (sterile and/or less fertile individuals take care of the offspring of more fertile ones), although non-reproducing individuals (namely, post-menopausal grandmothers) exist in human societies, too. However, these non-reproducing society members may also include young females (e.g., virgin priests and warriors, but also prostitutes, who may bear children but refuse to rear them) and males (e.g., eunuchs, celibate priests, unmarried slaves, servants, and soldiers). Ultrasociality and social organization based on city-states certainly promoted the role of these non-reproducing individuals in the overall organization of human societies, enhancing the similarities between advanced human and insect societies.
In section 2.3 (para. 4), G&K mention Smaldino's (Reference Smaldino2014) statement that “an ant colony represents the extended phenotype of the queen.” However, they do not point out that such a view is oversimplified, as ant colonies are often polygynous (contain numerous queens). Consequently, relatedness between nestmates is frequently very low, sometimes even close to zero. Nevertheless, ants from such colonies can successfully cooperate (Hamilton Reference Hamilton, Ito, Brown and Kikkawa1987). However, conversely to what was stated by G&K in section 5.1 (para. 4), important within-group conflicts do exist also in insect societies. Members of the same colony compete mostly for the access to reproduction (Heinze et al. Reference Heinze, Hölldobler and Peeters1994; Hölldobler & Wilson Reference Hölldobler and Wilson1990; Reference Hölldobler and Wilson2009). Ant workers are not completely sterile: they may reproduce by parthenogenesis and produce males from unfertilized haploid eggs and sometimes also females from unfertilized diploid eggs (Hölldobler & Wilson Reference Hölldobler and Wilson1990). Various forms of worker resistance against reproductive dominance of the queen lend support to the statement of G&K that “resistance has no analytical equivalent in ants (except perhaps attempts by workers to reproduce)” (sect. 6, para. 4). However, ant resistance may take yet other forms. In particular, enslaved ants may successfully rebel against their social parasites. Various forms of slave resistance include physical aggression against adult slave-makers, devouring of slave-maker's brood, attempts to reproduce in the slave-maker's colony, and slave emancipation (Czechowski & Godzińska Reference Czechowski and Godzińska2015).
In conclusion, all of these examples of not strictly adequate statements demonstrate that ant and human social behavior should be compared in a painstakingly careful way in order to draw entirely valid parallels.