Scale Models? What Insect Societies Teach Us ... - Semantic Scholar

Scale Models? What Insect Societies Teach Us about Ourselves JAMES T. COSTA Department of Biology, Western Carolina University

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NE BRIGHT sunny day in 1846, Henry David Thoreau happened upon a savage battle. The unfolding butchery didn’t disturb the peace of rural Concord, however, for the conflict was noiselessly conducted by ants just a stone’s throw from his cabin. “The legions of these Myrmidons covered all the hills and vales in my woodyard,” Thoreau reports, “and the ground was already strewn with the dead and dying.” This was internecine war, Dresden or Austerlitz in miniature, “the red republicans on the one hand, and the black imperialists on the other.” Musing on the considerable myrmecine carnage before him, Thoreau comments, “I was myself excited somewhat even as if they had been men. The more you think of it, the less the difference.” And therein lies a lesson for humanity, which is why these astute observations from Walden were often reprinted as an essay in their own right under the title “Battle of the Ants.” As he was a keen observer and critic of nineteenth-century America and one of the most eloquent philosophers of the relationship between nature and human society, it is unsurprising that Thoreau should join in a long tradition of observers who have seen parallels between insect and human societies. A great many social thinkers of the past would concur with him—the more you think of it, the less the difference— which is why it is not coincidental that these insect colonies have come to be called “societies” by entomologists to begin with. But, interestingly, the society that is seen often bears an uncanny resemblance to that of the social organism peering. What do people see when they view insect societies? Besides repulsion or fascination, what sense do they make of teeming masses of incessantly-probing ants, swarms of armed-and-dangerous hornets, or legions of termites marching under cover of darkness? These are the social insects that come most readily to the minds of entomologists and PROCEEDINGS OF THE AMERICAN PHILOSOPHICAL SOCIETY

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non-entomologists alike—ubiquitous, abundant, conspicuous groups that are forces to be reckoned with. No mere masses of bugs, these colonies seem to be highly integrated functional units whose members communicate, cooperate, and coordinate their efforts at such readilyappreciated activities as defending their group, finding food, building homes, and caring for their young much as do people. People have long looked to the natural world for guidance and instruction, seeking analogy, metaphor, and messages in all quarters of the animate and inanimate worlds. It was thus inevitable, perhaps, that these remarkable insects should have insinuated themselves into our lore, legend, and very conceptions of ourselves as social beings. The nineteenth-century Natural Theology tradition of seeking sermons in stones captures the essence of the human tendency to find meaning in nature. But what sort of meaning? Typically, meaning for a particular philosophy of life, for how people should live. The noted myrmecologist C. P. Haskins asks, in his 1939 book Of Ants and Men, “Can we as we gaze at the ant colony, discern any social pitfalls which menace [ants and humans] alike, into which ants, perhaps, have fallen more deeply than men . . . ?” (p. 5). History suggests that any attempts to draw parallels between humans and other organisms, and most especially attempts to moralize from nature, are doomed to failure or worse. There is no shortage of misguided social policies and doctrines in our century alone, born of a woeful ignorance or selective reading of “nature.” It is precisely for this reason that it is instructive to consider what sense has been made of social insects, and the ways in which these organisms have provided grist for the mills of social engineers, commentators, and poets since time immemorial. Protagoras said that man is the measure of all things, arguing that human knowledge is relative to the observer. This relativity is starkly evident when the observer is projected onto the observed, and when it comes to insectan societies humans are indeed the measure, using themselves as the meter stick. Consider the following sampling of social insects as model and metaphor, exploring how remarkably divergent social views are seen written upon the same hives and anthills. Go to the Ant . . . Industrious ants and busy bees are familiar staples of instruction. In this regard, King Solomon may have been the first on record extolling the virtues of ants: Proverbs 6 exhorts us to “go to the ant” and “consider her ways and be wise.” Aesop’s famous fable repeats this myrmecine model of industry and wisdom: it seems the grasshopper is doomed

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as much for its imprudent frivolity as for laughing at the industrious ant. These tried-and-true exemplars focus on the individual insect, though, and not the group. The moralizing gets more interesting when colonies as a whole are considered, for of course it is the colony that seems to resemble human society. The book of Proverbs notwithstanding, rulers have been viewed as integral to the structure of bee and ant societies. Aristotle counted bees and ants among the “political” animals, and many entomologists have in a sense concurred; the genus of the common paper wasp, Polistes, says it all: the name means “founder of cities.” Moreover, the idea of a diminutive city-state or polity is built into the very language entomologists use in describing the structure of insect societies and the roles played by their members. These insectan civilizations are based upon a division of labor: the queens, kings, workers, soldiers, and castes entomologists speak of are now technical terms with specific definitions, but they obviously originate in the human sociopolitical realm. In some cases the transfer of terms is not accurate; for example, there are many so-called “slave-making” ant species that raid the nests of other species, stealing larvae and pupae away to their own dens. There the helpless brood develop into adult ants and commence working for their kidnappers, defending their new colony-mates and collecting food as if they had never left their natal colony. This phenomenon suggests slavery on casual inspection, but is more closely analogous to animal husbandry since members of another species, not the same species, are pressed into service. But in general, terms like “worker” and “soldier” accurately reflect a division of labor within the colony. This division of labor naturally extends to colony leadership, the essence of political structure. The Royals The descriptive terms now applied to the organization of insect societies have their historical roots in a class-structured monarchical model that surely predates the ancient Greeks. Aristotle was one of the earliest thinkers to try to make sense of the social insects. Although his lengthy treatises on animals and their habits, reproduction, and classification are a curious mixture of keen observation and anecdote, and some of his observations of social insects are fraught with inaccuracies (one curious argument made in De Generatione Animalium 3.10 asserts that worker honeybees cannot be female because “nature does not assign defensive weapons to any female creature”—ironic in that sting-bearing bees and ants are exclusively female, as those structures are evolutionarily modified egg-laying structures), they provide instruc-

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tive glimpses into how he and his contemporaries conceptualized the natural world in relation to the human world. His Historia Animalium uses royalist language to portray bee societies: “rulers” and “kings” are standard descriptors of the head of the hive. This structured monarchical portrait was to dominate the Western conception of insect societies for many centuries, probably because that was the nature of Western human societies for many centuries. A snapshot of the monarchical view of social insects in the fourteenth century is found in Henry V. In Shakespeare’s play the archbishop of Canterbury exhorts the young King Henry in act 1, scene 2 to engage the French. Of concern is whether the Scots will attack as soon as the English turn their backs, and the archbishop argues that these enemies can be held at bay through a division of labor like that of the honeybees, splitting the army into an invading force abroad and a defending force at home. Therefore doth heaven divide The state of man in divers functions, Setting endeavour in continual motion; To which is fixed, as an aim or butt, Obedience: for so work the honey-bees, Creatures that by a rule in nature teach The act of order to a peopled kingdom.

The analogy is not perfect since honeybees, unlike some social insects, do not divide colony defensive tasks, but his point is that both societies are marked by a division of labor and are bound by the glue of obedience to the ruler. The implication that bee societies are in this regard models for our own is clear from the remainder of the archbishop’s speech: [The bees] have a king and officers of sorts, Where some, like magistrates, correct at home, Others, like merchants, venture trade abroad, Others, like soldiers, armed in their stings, Make boot upon the summer’s velvet buds. . . .

Surely both the portrait of the structured bee society and its use as a model for human society would have resonated with Shakespeare’s audience in 1598. Divine-right monarchy and class-structured division of labor was only natural. The insectan version of divine-right monarchy is also found in a remarkable work published in 1609 by Charles Butler, The Feminine Monarchie, or, A Treatise Concerning the Bees, and the Due Ordering of Them. This treatise, one of the earliest comprehensive treatments of beekeeping and the habits of honeybees, was published in the reign

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of James I, the first Stuart monarch of England. Its portrayal of honeybee societies as perfect monarchies seems to go beyond the flattering ornamental statements often prefacing works published under the watchful eye of patron sovereigns: in his opening chapter, after extolling the many virtues of honeybees, Butler marvels that “all this [is found] under the government of one Monarch, of whom above all things [the worker bees] have a principal care and respect, loving, reverencing, and obeying her in all things.” Butler is serious about the virtues of monarchy, as he goes on to explain why, should the queen “bring forth many princes,” the new royals will either leave the colony in a swarm or be killed by the workers: “For the bees abhor as well polyarchie, as anarchie, God having showed in them . . . an express pattern of a perfect Monarchie, the most natural and absolute form of government” (chap. 1, emphasis added). In other words, the bees will not abide more than one leader in the hive, driving off or killing off would-be oligarchs till one ruler remains; God has here provided a perfect monarchical model for people. Butler’s observation has a basis in fact: many social insect species do indeed have a single dominant reproductive, the queen, and will kill or drive off other individuals who attempt to reproduce within the colony. This is readily understood in terms of kin selection theory, first intimated by Darwin in his Origin of Species and later mathematically elucidated by the evolutionary geneticist W. D. Hamilton in the early 1960s: although these colonies remind us of city-states, they are actually large families in which the sibling workers are the queen’s offspring. By a quirk of sex-determination genetics in bees and their kin, sisters are more closely related to each other than to their brothers, nieces, nephews, or even their own daughters. Hence natural selection favors a female-biased colony sex ratio in which sisters defend their mother’s reproductive dominance. When honeybee colonies swarm, however, it is the old queen that leaves the hive with a retinue of workers to establish a new hive, leaving one of her daughters behind to become the new queen. (Establishing a new colony is a risky business, so it makes evolutionary sense for the old queen, who probably does not have very many more years to live anyway, to assume this risk and leave the tried-and-true nest site for her daughter heir.) The expelled oligarchs of The Feminine Monarchie were in all likelihood the old queen and a swarm of her worker daughters. Leviathan and Commonwealth The fifty or so years following publication of The Feminine Monarchie witnessed a sea change in how monarchy as an institution was viewed,

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and there is a parallel shift in the way insect societies are discussed in relation to this political system. In particular, the 1651 publication of Thomas Hobbes’s Leviathan marks a shift in philosophy from conventional monarchy to an alternative theory of absolute authority. The philosophy of Hobbes (1588–1679) represents a milestone along a tortuous path that originates with the Magna Carta in 1215 and culminates in Rousseau’s social contract of the eighteenth century. But far from arguing for representational government, Hobbes argues that we very much need our kings. However, absolute sovereigns are made, not divine, Hobbes argues, and the voluntary relinquishing of personal rights and concomitant conferring of those rights on a designated sovereign is necessary for social stability. People must live under the thumb of a coercive power—the Leviathan—because humans cannot be trusted to live harmoniously without an absolute sovereign to keep them under control. The state of nature is “the war of the all against the all” (the Hobbesian War) and the commonwealth is formed by tacit agreement among citizens to relinquish rights to the sovereign to avert this horrific bloodbath. This view represents two fascinating and interrelated departures from earlier sociopolitical views of human society: first, that the human polity is fundamentally unnatural, and second, that, that being the case, there is no such thing as divine conferral of the right to rule, since such a right cannot exist if the polity itself is an artificial construct. Viewing human society as unnatural turned on its head a conception of human biology dating back to the ancient Greeks: Aristotle wrote in The Politics, for example, that “man is a being meant for political association, in a higher degree than bees or other gregarious animals.” This reversal also meant a shift in the way social insects were called into service as metaphor and example by Hobbes. For Hobbes, insect societies are not so much diminutive polities with authoritarian queens and dutiful workers, as they are harmonious functional commonwealths where colony members “have no other direction than their particular judgments and appetites.” Most importantly, their association is not an artificial construct of convenience. Hobbes devotes a sizable portion of chapter 17 of Leviathan to an explicit comparison between human and insect societies. He addresses, as he puts it, “why certain creatures without reason or speech [i.e., ants and bees] do nevertheless live in society, without any coercive power.” These insects, he argues, can live harmoniously without the rule of an iron fist. He presents six ways in which human society differs from that of ants and bees, all of which reflect his conception of the meddlesome and bellicose antisocial “nature” of people and underscore the need for a dictator to rein them in.

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It is worth paraphrasing Hobbes’s arguments: social insects, unlike people, are not continually in competition for dignity and honor, while “among men there arises on that ground envy and hatred and finally war.” For bees and ants the common good is indistinguishable from the individual good—one for all and all for one—but “man, whose joy consists in comparing himself with other men, can relish nothing but what is eminent.” He argues that social insects lack reason and so are unaware of the faults of their leaders, such as they may be. Among men, on the other hand, “there are very many that think themselves wiser and abler to govern . . . better than the rest . . . these strive to reform and innovate . . . and thereby bring it into distraction and civil war.” Hobbes further argues that social insects lack language and its use for guile and evil, while some men “can represent to others that which is good in the likeness of evil, and evil in the likeness of good . . . discontenting men and troubling their peace at their pleasure.” In this vein, too, non-rational creatures like bees and ants pay no attention to personal injury and damage and so are typically not offended by their colony-mates. Man on the other hand is “most troublesome when he is most at ease, for then it is that he loves to show his wisdom and control the actions of them that govern the commonwealth.” Hobbes sums up his point by asserting that bee and ant societies are intrinsically natural, and thus the “agreement” binding these societies together is natural. In stark contrast, “[the polity] of men is by covenant only, which is artificial; and therefore it is no wonder if there be somewhat else required besides covenant to make their agreement constant and lasting.” Hobbes concludes that that “somewhat else” is an iron-fisted ruler to keep people “in awe and to direct their actions to the common benefit.” A most unflattering portrait of humanity is painted here, but Hobbes knew whereof he spoke, having lived through the English civil wars, the beheading of Charles I in 1649, and Cromwell’s Protectorate. No friend of divine-right monarchy, Hobbes watched with what was probably no small unease as the Protectorate crumbled and the House of Stuart was restored in 1660 under Charles II. The portrayal of social insects in Leviathan is in some respects little different from that of Hobbes’s predecessors: these creatures exemplify a social hierarchy controlled from above, albeit harmoniously, and are natural polities. In terms of political philosophy, it is more significant that Hobbes sees human society as inherently unnatural, which in turn means that divine right to rule cannot exist. Some subsequent comparisons of insect and human societies restore the old monarchical model, but interesting twists reflective of the times arise. For example, it might seem curious that while Alexander

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Pope, in his celebrated 1752 poem Essay on Man, invokes the insectan monarchy for mankind’s instruction, he also mentions commonwealth and property rights: Here subterranean works and cities see; There towns aerial on the waving tree. Learn each small People’s genius, policies, The Ant’s republic, and the realm of Bees; How those in common all their wealth bestow, And Anarchy without confusion know; And these for ever, though a Monarch reign, Their sep’rate cells and properties maintain. (Essay on Man, epistle 3, emphasis added)

In this insectan commonwealth, individual “cells and properties” are maintained despite the rule of a monarch, understandable in terms of the evolving social contract that limits the formerly absolute power of sovereigns. The monarchy is becoming constitutional. Socialist States Many, if not all, insect societies do exhibit elements of hierarchical structure, predisposing them for a ready fit into the human monarchical model. But they also exhibit elements of cooperation, and so it is unsurprising that, in the trend away from absolute monarchy in human polities, hierarchy is de-emphasized and cooperation becomes the focus for some thinkers. This shift occurs concomitantly with the rise of socialist ideology. Of course, the ideal of cooperation is not exclusive to socialism: the cooperative nature of social insects can also be viewed in terms of social experiments aimed at communal living. A good example of this is the early Mormon church, which modeled itself on honeybees. (Utah is known as the Beehive State more for this reason than for any special affinity of its citizens for apiculture.) But socialism as an institution inevitably seems to invite comparison to the apparently cooperative and prosperous insect societies. In Capital, Karl Marx argues, like Aristotle, that humans are social animals. But for Marx, immersed in the industrial revolution, it is the tendency for collective productivity to exceed that of individuals that compels the formation of society. Cooperation and ownership in common has been the hallmark of human societies since the dawn of human development, Marx argues, as “each individual has no more torn himself from the navel-string of his tribe or community, than each bee has freed itself from connexion with the hive” (Capital, vol. 1, chap. 13). Here we see a shift in focus away from the head of the hive—the

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queen—to the worker and his/her dependence on the group. Marx, unlike Hobbes, does not present a detailed analysis of insect societies in relation to our own, but another socialist philosopher picks up this theme, seeing cooperation in insect societies where others before him saw only obedience to the queen. Petr Alexeyevich Kropotkin (1842–1921) was a prolific writer of social and scientific tracts. Although a prince born into an aristocratic Moscow family, Kropotkin early on became a self-avowed anarchist and, later, a socialist. Jailed repeatedly for his subversive political views, he lived much of his life in exile, returning to Russia only in the wake of the 1917 revolution. Kropotkin also had some training in zoology. He was disappointed in the competition-driven vision of Darwinian evolution promulgated by Spencer and Huxley. In response to what he saw as an overemphasis on relentless competition in nature, Kropotkin focused instead on cooperation, and produced in 1902 an acclaimed volume entitled Mutual Aid: A Factor of Evolution. Mutual Aid is a fascinating book that expressly sets out to challenge the dominance of the “struggle for existence” paradigm. The book’s opening chapter, “Mutual aid among animals,” presents cooperation as “a law of Nature and chief factor of progressive evolution.” Inevitably Kropotkin looks to the social insects to find lessons for humanity, but his conclusions differ radically from those of his predecessors. In a manner reminiscent of the scala naturae of old, Kropotkin recognizes a hierarchy of social perfection, the pinnacle of which, interestingly enough, is occupied not by humans, but by ants and termites! He writes of these insects that “self devotion . . . and very often self sacrifice for the common welfare are the rule. The ants and the termites have renounced the ‘Hobbesian War,’ and they are the better for it” (Mutual Aid, 1902, p. 14). Human society is fraught with strife and conflict, as pointed out by Hobbes. According to Kropotkin, human societies thus cannot be compared directly with those superior species—the ants and termites—that have renounced the Hobbesian War to achieve harmony and cooperation. Rather, humans are best compared with social insects somewhat further down the scale of social perfection, creatures that still exhibit some measure of discord. Curiously, these turn out to be bees in Kropotkin’s scheme. Why? Because among bees “predatory instincts and laziness continue to exist.” We are further told that “about the sugar plantations of the West Indies and the sugar refineries of Europe, robbery, laziness, and very often drunkenness become quite usual with the bees” (Mutual Aid, p. 17). Those are telling social attributes on which to focus. One interpretation of these statements is that, under conditions of plenty and opulence, antisocial “predatory” instincts rear their

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ugly heads, perhaps an attack on capitalist society with its emphasis on material wealth. The important point is Kropotkin’s stress on the cooperative characteristics of insect societies, and how degree of cooperation and peacefulness serves as ordination along his social hierarchy. Insect Societies in the Mirror Like the fabled descriptions of an elephant by seven blind men, visions of ant and bee societies and what they represent have turned on the perspectives and values of their observers. From Aristotle through the Enlightenment, these scale model societies were exemplars of singlerule monarchy, afterward slowly evolving into commonwealths and then cooperative socialist states, depending on whether their apparent hierarchical structure or their apparent cooperation is considered paramount. Curiously, only democracy seems to have been overlooked, perhaps because of the difficulty of grafting the idea of representation and limited-term leadership onto these colonies. Does the popular conception of social insects as mindless automatons fly in the face of the democratic ideal? The many interpretations given the same phenomenon, in this case insect colonies, likely reflect a certain subjectivity in our ability to rationalize the natural world. But is this observation anything more than an amusing curiosity? I would argue that the patterns sketched in this essay do offer something more than amusement: we can profitably look to the way we have looked to the ant to learn something about ourselves, and about the cautionary tale that “something” presents. If such a thing as human nature exists, it seems to include a tendency to seek reassurance or justification for our actions and institutions from a higher authority. With the accelerating rise of the sciences and concomitant understanding of the material basis for natural phenomena, that authority seems to have become nature itself. This has its benefits: much suffering has been alleviated and compassion inspired by recognizing a physiological or genetic basis for a host of maladies from epilepsy to addiction. But the practice of seeking lessons from the natural world also has its pitfalls. Firestorms of controversy continue to rage, for example, over the degree to which genes may influence our behavior and our abilities, the contentious arena in which the biological and cultural elements of our humanity are weighed against each other. One need look no farther than the 1994 book The Bell Curve, by Richard Herrnstein and Charles Murray, to see one of the most recent readings of nature—in this case our putatively inherent, genetically-determined cognitive abilities—readily translated into policy recommendation. We may have become too sophisticated to overtly analogize from animals, but that

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only means the moralizing has become more subtle, perhaps insidiously so, couched in terms of that which is considered natural. Consciously and unconsciously, our interpretations of the world are likely more shaped by cultural context than many of us realize, since by necessity what we observe is filtered through our own experiences, beliefs, desires, and expectations. We must therefore be cautious in our reading of the world and in what we make of the “natural.” What appears an eminently reasonable conclusion drawn from an objective assessment of what is, after all, “only natural,” may serve only to confirm a world view already firmly in place. All of which suggests we should bear in mind the most insightful parallel between human and insect societies, pointed out in 1930 by Harvard president Abbott Lawrence Lowell in a speech conferring an honorary doctorate upon the great myrmecologist William Morton Wheeler. Wheeler was a pioneer of the field of ant biology; the modern conceptual framework entomologists use to study the ecology and evolution of social insects has its origin in his work. Lowell cites Wheeler as a “profound student of the social life of insects, who has shown that they also can maintain complex communities without the use of reason.” His tongue-in-cheek remark may be truer than Lowell thought! Acknowledgments I thank Professor E. O. Wilson for kindly opening his personal library to me for researching this essay, and Ani Patel, Stefan Cover, Hal Herzog, and Scott Philyaw for helpful comments and criticisms on earlier versions of the manuscript. Special thanks are due Kathy Horton, for helping to track down the Lowell quotation. I dedicate this paper, in gratitude, to Judy Best, Distinguished Teaching Professor of Political Science at SUNY Cortland.