The Illustrated Insectopedia - Hugh Raffles (2010)


When I wish to attract some bees for training experiments I usually place upon a small table several sheets of paper which have been smeared with honey. Then I am often obliged to wait for many hours, sometimes for several days, until finally a bee discovers the feeding place. But as soon as one bee has found the honey many more will appear within a short time—perhaps as many as several hundred. They have all come from the same hive as the first forager; evidently this bee must have announced its discovery at home.



Karl von Frisch won a Nobel Prize in 1973 for his discovery of “the language of bees.” It was the year of ethology, and along with von Frisch, the prize in Physiology or Medicine was also awarded to Konrad Lorenz and his Dutch colleague Nikolaas Tinbergen. There was nothing recondite here, no obscure fiddling at the margins of theory. The 1973 prize was awarded for populist research that illuminated the mysteries of animal existence and promised profound and far-reaching truths about the human condition.

Honeybees, said von Frisch, though so tiny and so different, possessed language, the capacity long definitive of humanity. Through a series of elegant experiments carried out over nearly half a century, he showed that they communicated symbolically, that, in a manner more complex than that of any other animals apart from humans, they drew on experience and memory to convey information to each other and to their fellows.

More than ninety years after his first reports, these discoveries are still exciting. And they are made more so by von Frisch’s way of telling. By inclination and early training a naturalist, he offered nature not in today’s technical language of genomics but in his own deeply personal language of bees, a remarkably affective language that imbued his subjects with purpose and intentionality, that made them appealing and familiar.

Von Frisch offered a science of “what animals do, and how and why they do it” that was as comfortable with ontological difference and abiding mystery as it was with the more familiar scientific impulse toward revelation.1 Unashamed in his confessions of affinity, he made readers believe—just as he did himself—that they could understand bees, psychologically and emotionally. He turned his public into animal analysts. And in doing so, he gave new impetus—though, perhaps, despite himself—to the Darwinian notion that not only the morphological but also the behavioral, moral, and emotional basis of human existence could be found in the lives of nonhuman animals.2 Von Frisch spoke for honeybees. And he made them speak. He didn’t just give them language; he translated it. Is there anything that is more irresistible?

Nonetheless, these affinities were deeply fraught in a discipline barely born yet already haunted by the specter of fallibility. Ethology’s ghost was Clever Hans, the celebrity horse whose cleverness unfortunately lay not in mathematics but in an uncanny sensitivity to the nonverbal cues of his unwitting trainer. Clever Hans’s much-publicized debunking by the psychologist Oskar Pfungst in 1907 pushed questions of animal cognition to the very margins of scientific legitimacy and made it clear that ethology was at mortal risk from the allure of its subjects.3

It was a foundational temptation to which the resolutely anti-psychological behaviorists would not succumb. But it was the seduction to which von Frisch, caught between affect and object, preoccupied, as he himself wrote, by the interplay between “psychological performance and the physiology of the senses,” would forever be in thrall.4

Because von Frisch loved his bees. Loved them with a gentle passion. Tended and nurtured their generations. Warmed them in his cupped hands when the brisk air stiffened their wing muscles. Held them as his “personal friends.”5 They were his bees in the way that anthropologists of the past may have fancied the remote tribes among which they lived to be their tribes. That same heady mix of science, sentiment, and proprietorial pride, the same willingness to assume responsibility for another’s fate.

So even as he took such care over the tiny creatures’ welfare, von Frisch would lovingly (with another love), painstakingly (with a professional patience), and delicately (with such safe hands) snip their antennae, clip their wings, slice their torsos, shave their eye bristles, glue weights to their thoraxes, and carefully paint shellac over their unblinking eyes, modifying their bodies, mutilating their senses, manipulating their behavior according to the experiment’s requirement, reconciling his will to suture the yawning gap that separated human from insect with his unspoken assertion of a natural sovereign power.


In April 1933, the Nazi-dominated Reichstag passed the Law for the Restoration of the Professional Civil Service. Jews, spouses of Jews, and political unreliables could now be legally dismissed from the universities.6

By then, von Frisch was director of the new Rockefeller-funded Institute of Zoology at the University of Munich and a leading figure in German science. Years before, in the landscaped and columned courtyard of the institute, he had, as he recalled in his memoir, fallen “irresistibly under the spell of the honey-bee.”7

His enchantment by those he would come to call his little “comrades” had in fact begun even earlier. In 1914, with a magician’s flair, he publicly demonstrated what now seems the rather unsurprising truth that honeybees—whose livelihood, after all, depends on their identification of flowering plants—are able to discriminate by color (despite being red-blind). Using the standard behavioral method of food rewards, he trained a group of bees to identify blue plates. He then showed them small squares of colored paper and watched delightedly as they congregated “as if by command” for his skeptical audience.8

But it was in the garden in Munich that the bees first danced for him: “I attracted a few bees to a dish of sugar water, marked them with red paint and then stopped feeding for a while. As soon as all was quiet, I filled the dish up again and watched a scout which had drunk from it after her return to the hive. I could scarcely believe my eyes. She performed a round dance on the honeycomb which greatly excited the marked foragers around her and caused them to fly back to the feeding place.”

Although beekeepers and naturalists had known for centuries that honeybees communicated the location of a food source among themselves, no one knew how. Did they lead one another to the nectar? Did they diffuse scent trails? “I believe,” von Frisch wrote more than forty years later, that this “was the most far-reaching observation of my life.”9

Under the civil service law, von Frisch and his academic colleagues—as well as all other civil servants in the Reich—were required to produce documentary proof of their Aryan ancestry. Already suspect for his willingness to sponsor Jewish graduate students even when their theses were far from his own specialties, von Frisch found himself in an even more dangerous dilemma.10 His mother’s mother, now deceased, the daughter of a banker and the wife of a philosophy professor, was a Jew from Prague. At first the university protected its star zoologist, arranging for his safe classification as “one-eighth Jewish.” But imagine the virulent mixture of ideology and ambition that began to ferment, fed by a rigid institutional hierarchy and the lack of opportunity for advancement among scholars locked out of academic privilege despite their years of training. In October 1941, the campaign against von Frisch succeeded in forcing his reclassification as “second-grade Mischling”—one-quarter Jewish—and securing the order for his removal from his post.

As we know, von Frisch survived the Nazis. Inevitably, though, it was far from straightforward. Influential colleagues mobilized on his behalf, arranging a platform in Das Reich, a new weekly in which Goebbels contributed the editorials. Von Frisch wrote about the national-economic contribution of the Zoological Institute and how its work was vital to the resilience of the home front.11 Eventually, though, if in somewhat tortuous fashion, it was the bees that saved him. For two years, an outbreak of the parasite Nosema apis had ravaged German hives. Both the national honey crop and agricultural pollination were threatened. Through the intervention of a highly placed ally, von Frisch was appointed as a special investigator, and a panicked Ministry of Food was induced to defer his dismissal from academia “until after the war.”12

The indifference of the honeybees to politics did not prevent their recruitment to the National Socialist war effort. The ministry soon expanded the Nosema remit to include a search for ways of persuading bees to rationalize pollination by visiting only economically desirable plants. Years before, von Frisch had experimented with scent guidance—training bees to respond to a particular odor before freeing them to visit the associated flower—but he had been unable to generate commercial interest. This time, galvanized by looming calamity, national enthusiasm, and news of a large-scale Soviet research project along similar lines, the Organization of Reich Beekeepers rushed to sponsor his work.

Exhausted by the intensifying air war on Munich, von Frisch and his lifelong co-worker, Ruth Beutler, evacuated to the village of Brunnwinkl on the shore of Lake Wolfgang, southwest of Salzburg. This was where von Frisch had spent his childhood summers, and attached to the family house was the natural history museum he had founded as an eager seventeen-year-old. It was here, pursuing adolescent obsessions, that young Karl had enrolled relatives and family friends in scouring the nearby woods and shoreline for local fauna. It was here, at the old mill on the edge of Lake Wolfgang, under the quiet hand of his uncle, the prominent Viennese physiologist Sigmund Exner, that he developed the classical skills in observation and manipulation that would characterize his experimental research.

And it was also here, here among the animals, that von Frisch found his “reverence before the Unknown,” less a formal religious conviction than a commitment to a pantheistic relativism. “All honest convictions deserve respect,” he insisted, “except the presumptuous assertion that there is nothing higher in the world than the mind of man.”13 And it was here, as he tells it in straightforward yet often lyrical prose, that his liberal Catholic family—doctrinally liberal in an era when Austrian biologists were routinely dismissed for espousing evolution—created a bourgeois haven, a home for science and the arts, for the gentle satisfactions of polite culture far from the upheavals of early-twentieth-century Mitteleuropa: his spirited mother and his caring if reserved father, his three older brothers, all preparing merely for the uneventful unfolding of long and distinguished academic careers.

And it was here, in the cocoon of family memory, as the Allied bombs rained firestorms on Munich and Dresden and as the air thickened over Auschwitz, that von Frisch and Beutler took advantage of their Reich permits to revisit the work on bee communication that he had laid aside some two decades earlier.

In those long-ago studies in the courtyard of the Institute of Zoology, von Frisch had identified two “dances”—he named them the round dance and the waggle dance—and concluded that bees used the former to indicate a source of nectar and the latter to indicate a source of pollen. Beutler had continued this work in the intervening years but had begun to doubt the hypothesis. Resuming their experiments together in 1944, they discovered that when they positioned the feeding dishes more than 100 yards from the hive, it didn’t matter what substance the bees were carrying: on their return, they all performed waggle dances. Rather than a descriptor of material, the variation they observed in the dances must be the bees’ way of communicating the far more complicated information of location. This ability to accurately describe distance and direction “seemed,” von Frisch wrote, “altogether too fantastic to be true.”14

It was the complexity of the bees’ behavior that was so arresting. Making connections between the intricate sociality of honeybees—which live in self-reproducing “colonies” of thousands of individuals—and the development of sophisticated forms of communication is commonplace now. But early-twentieth-century animal studies were dominated by the conviction of biologists and psychologists that animal behavior was fully explicable in terms of a range of simple stimulus responses, such as reflexes and tropisms. And von Frisch’s bees were doing something that leading behaviorists such as John B. Watson and Jacques Loeb considered impossible: they were communicating symbolically, representing information through a form—a predictable pattern of physical movements—that was tied to its object “by social convention, tacit agreement, or explicit code.”15 What was more, this representing could take place several hours after the flight it described. It relied on registering the details of that flight, recalling its content, and, of course, translating and performing the significant information. Moreover, it also required an audience able to interact effectively in its interpretation. To Donald Griffin, the tireless advocate of animal consciousness and the sponsor of von Frisch’s 1949 lecture tour of the United States, this was “the most significant example of versatile communication known in any animal other than our own species.”16 Von Frisch went further. It was, he believed, an accomplishment “without parallel elsewhere in the entire animal kingdom.”17

Contemporary bee researchers have refined von Frisch and Beutler’s wartime revisions of the dance theory. There is, most now believe, no difference between the types of information contained in the two main dances.18 Both use waggling to communicate distance and direction, and in both it is the enthusiasm of the performance that conveys the quality of food. Similarly, in both, the type of flower is revealed by the scent clinging to the insect’s body.

In Munich, von Frisch had placed feeding stations directly alongside the hive to facilitate communication between his assistants observing the dances and those stationed at the feeders. However, in the round dances that the bees perform to indicate nearby food, the waggles are abbreviated, occurring just as the dancer turns to begin her new circle. Von Frisch and his team failed to observe those subtle cues, and it is likely that the bee audience doesn’t take much notice of them either, relying instead on sense of smell to locate such proximate feeding places. But when food is further away—the transition occurs at a point between 50 and 100 yards for the Carniolan bee, the bee favored by von Frisch—bees returning to the hive interpose an additional sequence of steps, a straight run that contains a “vigorous wagging” of the abdomen, a side-to-side movement they may repeat thirteen to fifteen times per second.19 It is this distinctive stretch that contains the critical information. Gyrating in darkness amid the crush of bodies on what von Frisch called the hive’s “dance floor,” the returning forager is closely shadowed by three or four followers, who receive the dance information with their antennae, utilizing scent (to identify the type of flower), taste (to gauge the quality of its product), touch, and an acoustic sensitivity that allows them to pick up the near-air movements produced by the dancer’s wings.20

The dancer uses the sun as her reference point. Illuminated by daylight on the horizontal platform at the hive entrance, her movements are indexical, pointing directly ahead, “just as we point to a distant goal with raised arm and outstretched finger.”21 Dancing in the open, she orients herself by angling her body so that the sun is at the same angle relative to her body as it was during her recent flight to the food source.22

But the vast majority of dances take place inside the hive, in total darkness, on the surface of a vertical comb. Those conditions present the bee with a significant set of problems, which she resolves by reconfiguring the indexical association between the dance and the food source. This interior dance involves a temporal and spatial displacement as the bee converts the angle of the sun, which has permitted her to mime her flight during the outdoor dances, into gravitational terms. To succeed, the bee must note optically the angle between the direction of the sun and the food source on her outbound flight, remember that information, accurately transpose it to an angle that relates to gravity, and in doing so, include a calculation that corrects for the movement of the sun in the time that has passed between her outbound flight and the dance.23

If food is located in the direction of the sun, the bee runs upward along the comb; if the feeding place is away from the sun, she runs down. If the material is located at, say, eighty degrees to the left of the sun—as in feeding table II in the diagram—she points her waggle run eighty degrees to the left of vertical (II’), and so on.24 Even if the sun is obscured by clouds, she can locate its position by recognizing patterns of polarized light invisible to humans.25

Von Frisch tracked bees foraging some seven miles from their hive and discovered that they convey distance through some combination of number and rate of waggles, velocity of forward movement, and length and duration of the straight section.26 However, distance is a “subjective” quality, which bees measure in terms of the amount of effort they expend on their outward flight. Von Frisch demonstrated this by appending weights of different kinds to various parts of the animals’ bodies, exposing them to head winds, and forcing them to walk. In each case, they reported a greater distance than they did without the handicap.27

Von Frisch liked to work with “calm and peaceful” bees.28 They were cooperative, and he was responsive, designing experiments and apparatus around their needs and desires. The bees were affected by wind and temperature. They revealed astonishingly subtle senses of smell and touch. They responded actively to changing light conditions. They grew to recognize individual field-workers. Alert to their sensitivities, he could never be certain that their observed behavior was not symptomatic of the artificiality of experimental conditions and so allowed them to force him into exhaustive (and exhausting) replications of his tests as he struggled to find ways to repeat controlled experiments in natural conditions. When his discoveries were too astonishing, he wondered whether his attention had created “a sort of scientific bee.”29

He began by building an observation hive. This was a standard beekeeping hive fitted with glass windows, through which the bees could be watched relatively undisturbed. But he soon realized that bright sunlight and the visibility of patches of sky distorted the dances, so instead he developed his own range of hives with removable panels that allowed him to manipulate external conditions.

He designed feeding stations and special food dispensers. And he invented an automatic counting apparatus disguised as a flower to record bee visits when it was impractical or unnecessary to use volunteers.

He devised a coding scheme—an ingenious one—that allowed visual identification of hundreds of individuals. And he used a fine brush to number each bee with spots of colored lacquer while they fed from his sugar water.

But his true gift was in the design of simple and effective experiments of exceptional elegance. (He initially translated the dance language, for example, by systematically removing to progressively greater distances from the hive the food source that his bees had been trained to seek and then closely observing the dances performed by the returning foragers.) And what underlay this—in addition to patience, self-criticism, and a creatively methodical practice—was his natural historical eye for bee ecology, temperament, and habit, and a deep affinity with bee ontology, the being of a bee.

It was all this that enabled him to recognize the individuality of the members of the hive, their characteristic predilections and temperaments, their shifting moods, and their subtle variations of activity. It was, without doubt, a profoundly anthropomorphic commitment. His bees are “shrewd,” “eager,” and “phlegmatic”; at one point they even exhibit “class consciousness.”30 But it would be a mistake to think that anthropomorphism, which we could think of here as the impulse to understand other beings by reference to human interiority, is a sufficient framework through which to make sense of his work. For von Frisch, the honeybees were personal friends, but they were also profoundly mysterious in their difference. And it is this gap and its crossings that permit both reverence and subjection, both the relentless search for some kind of redemptive communion and the willingness to brutalize along the way.

Perhaps it is just the moment in which all this is taking place (the terrifying, dehumanizing political-historical moment that is also the thrilling moment in which all these discoveries are entirely new). Or perhaps it is the revived ethological determination to find the human in the animal. But it is clear that both in von Frisch’s estimation and in the unfolding of his study, the bees were his collaborators as much as his subjects. He tests them—and makes no effort to hide his disappointment on the rare occasions when they fail to demonstrate their acuity. But they also test him: challenging him to devise experiments sufficiently sensitive to approximate their enigmatic way of being.

Von Frisch plunged into the Brunnwinkl research as into the lustrous depths of another world. “I tried to bury myself in it completely,” he recalled, “taking as little notice as I could help of the events around me.” Life outside Brunnwinkl was beyond control. In Munich, the Institute of Zoology lay in rubble, his house, too, “a gaping hole.” The hostilities of professional life confounded him. He persuaded his wife to burn her diary.31 Who could be trusted? Who was reading? Who might be listening? But the bees … The bees spoke, but they were indifferent to politics. Theirs was a language unsullied by the corrupting jargon of the Third Reich. The bees had a purity. The bees had an intelligible rationality. The bees offered refuge.

We don’t know how Ruth Beutler felt, but Martin Lindauer, eventually von Frisch’s most distinguished student, describes returning severely wounded to Munich from the Russian front, expressing the desire to study science, and being sent by his doctor to attend a lecture on cell division by Karl von Frisch. Lindauer recalls the event as an epiphany that opened the prospect of a normal, meaningful life for a confused twenty-one-year-old who had refused to join the Hitler Youth and had been sent instead to dig the foundations of Dachau, who had volunteered for the German army after an earlier lecture—this by SS officers recruiting at his high school—and who found in von Frisch a stern mentor with a “zeal for science … [who] tolerated no fraud … [who was] an extremely exacting person.”32

Perhaps it’s no surprise that, like his teacher, Lindauer experienced a profound attachment to the bees. As the national authoritarian order descended into chaos and conditions for professional science crumbled all around, von Frisch created an island of calm on Lake Wolfgang, finding in his honeybees a regularity, an ordered way of being in which, as in all well-run institutions, none need fear the unpredictable, none need feel unmoored. It was again the Germany of the amateur museum beside the Austrian lake, the Germany before the 1918 revolution, before the Weimar inflation, before the irruption of the Nazis. “After experiencing the senseless regime of the Hitler time, which was malicious, dishonest, and wrong from all perspectives,” Lindauer told an interviewer half a century later, “I drew strength from having work based on absolute correctness, honesty, and objectivity. Out of this material and spiritual collapse, this hopelessness, I was able, with Karl von Frisch as a teacher, to build a new way of life. I found a new home with the bees. It was really a new home, the bee colony.”33


It is not hard to understand. The honeybee colony has tens of thousands of members whose everyday life is a wonder of self-regulated complexity, a productive order continuously brought into being through the intricate fluidity of its social relations, exchange practices, and division of labor. The very first thing von Frisch tells us in his 1953 work The Dancing Bees is that honeybees are obligate social beings, that the level of task integration and cooperative interdependence is such that a bee alone cannot survive outside the hive: “There is no smaller unit [than the colony].… One single bee, kept all by itself, would soon perish.”34

Like ants, termites, and the other social insects, honeybees live in what entomologists call caste societies, an analogy zoologists use to indicate the presence of morphologically distinct occupational groups: the egg-laying queen, the multitude of nonreproductive female workers, and the few hundred fat male drones with big eyes whose sole purpose—so far as we know—is to have sex with the queen on her single mating flight and who ultimately, as winter approaches and food resources dwindle, will be dragged from the hive by the workers, expelled to starve or, if resistant, stung to their death. “From that time onwards until the following spring,” wrote von Frisch, evoking the feminist utopias of writers such as Charlotte Perkins Gilman, “the females of the colony, left to themselves, keep an undisturbed peace.”35

Not surprisingly, it was the workers that attracted the researchers’ attention. Von Frisch and Beutler catalogued their dances, and they made far-reaching discoveries concerning their orientation abilities. Lindauer extended their findings to swarming, nest location, and the extraordinary process of nest selection, which I describe below. All three carried out detailed studies of workers’ division of labor and time allocation, although Lindauer pushed this furthest, by tracking the entire life history of a bee he called 107.

Below is Lindauer’s first schematic of worker labor allocation. It shows what Thomas Seeley has called a “division of labor based on temporary specializations” and comes from Lindauer’s classic 1961 account, Communication among Social Bees, a collection of lectures he gave at universities in the United States.36 The column of figures indicates age in days. The whimsical bee people on the left are carrying out the activity associated with a particular point in a bee’s life (cell cleaning, caring for the brood, building and repairing the hive, guarding the nest, foraging for nectar, pollen, and water). The sketches on the right show the corresponding development of glands in the animal’s head (the nurse, or feeding, gland) and abdomen (the wax glands). Despite this tight linkage of activity, physiology, and life cycle, Lindauer was fully aware that under critical circumstances—for example, a sudden food shortage—these relationships could be radically interrupted. In such a situation, the glands might stop developing and the bee begin foraging before its appointed day. A bee’s physiology and behavior were flexible, adaptive, and responsive to changing conditions.

But that isn’t all. When Lindauer tracked 107, he realized that she was not only spending more of her time multitasking than attending to the one expected assignment, but she was also doing an awful lot of wandering around (“patrolling,” indicated in Lindauer’s diagram by the bowler hat and walking stick) and a considerable amount of what appeared to be nothing (40 percent of her time, in fact, “resting” on the chaise longue). Lindauer found explanations for these observations. Patrolling, he reasoned, was a form of site monitoring that allowed the bee to identify immediate needs and allocate her time accordingly. “Loafing,” he claimed somewhat less convincingly, maintained the “reserve troops,” who could swing into action as occasion demanded.37

Both of these unexpected activities suggested the importance of horizontal, peer-to-peer communication in a society organized without leaders or centralized decision making. The honeybees’ ability to maintain the hive’s internal environment—despite changes in external ambient conditions and the availability of critical resources—relies on contact between returning foragers and those already inside. The alacrity with which foragers are relieved of their loads, for instance, shows the degree of collective need for the substance in question. And not only is the recognizably sign-based language identified by von Frisch involved. Something more fundamental to social life is also going on. The bees are in constant physical contact, palpating each other’s head and antennae, sensing each other’s odor, passing compressed pollen to each other, sharing and exchanging the sugary contents of each other’s stomach, receiving each other’s near-field vibrations. Together, constantly, in the deep communal darkness, exchanging substances, sucking and regurgitating, touching, feeling, smelling, tasting, sensing. Together, touching, in the warm darkness, sucking, feeling, touching, smelling, tasting, touching. Another country. Another language of bees.

And this language is somehow tied to that other language that is all around us here: the language of colonies, of castes and races, of sisters and half sisters, of queens and workers, the language of dance. The language of language, for heaven’s sake! This language didn’t disappear with von Frisch and Lindauer either. Today’s bee scientists speak it too, even if they often bury it in a mechanical discourse of bioenergetics, a dissonance apparent in the distance between the anthropomorphic terminology and the machine-like organism it describes.

The new bee is an evolutionary bee for whom (as for all social insects) society is the individual and whose relationship to the hive is equal to that between the cell and the body. Out of these metaphors comes a compelling narrative of bee evolution in which selective pressures operate at the level of intercolony competition for food, foraging area, and other resources, a narrative supported by the absence of observable tension within the hive.38

But von Frisch suggests a supplement. It is not only—as all beekeepers know—the hives that exhibit different personalities (some tidy, some messy, some peaceable, some aggressive). In von Frisch’s story, the interplay between individual and collectivity leaves room for individual variability and for the role of varying bee capacities and talents in furthering collective success. In his version, the hive is the expression of a culture of cooperation among its thousands of distinct individuals.


Ernst Bergdolt, lecturer in botany at the Institute of Zoology in Munich, joined the Nazi Party in 1922, when he was just twenty years old. A presciently premature fascist, in 1937 Bergdolt became an editor of the Zeitschrift für die gesamte Naturwissenschaft (Journal for the Entire Natural Sciences), the most significant attempt to wrestle the biological sciences into conformity with Nazi ideology.39 It was Bergdolt, the leading light of the German National Socialist Lecturers’ League, who led the campaign to remove von Frisch from the Institute of Zoology. This is from a letter he wrote to the Ministry of Education, calling for the director’s dismissal:

Professor v. Frisch has an unusual ability to make propagandistic use of the results of his research, the sort of ability we know from Jewish scientists. In contrast, he lacks entirely the ability to survey his work from a broader point of view, let alone to find connections to the natural establishment of a volkish polity, something that seems so self-evident and would be so easy given his areas of expertise, bees.40

Bergdolt had already tried and failed to arrange von Frisch’s prosecution for cruelty to animals.41 His opening shot here is little more than a conventional invocation of “Jewish science.” But the second charge was more unusual. While the logic of the hive offered von Frisch and Lindauer refuge from what they experienced as the disorienting chaos of the Nazi Reich, for Bergdolt that same systematicity embodied the utopian promise of Nazism itself. The bees readily provided a mirror of the human. But they did so through lives that—despite the transparency of language—were sufficiently opaque to allow such apparently conflicting fantasies. Even if, in this instance, the fantasies were structured in the same feverish milieu.

This is only partly a matter of different ideas of order. For the Nazis, of course, order required and enforced a savage and exemplary hierarchy. In the hive, however, hierarchy was profoundly ambiguous. Not only were the gender relations of the bee world drastically at odds with the ideals of National Socialism, but the nominal leader—the queen—was a figure of doubtful autonomy, subject in almost all respects to the workers who serviced her. Yet such inconvenient details of bee order were as nothing in light of the allegorical possibilities of formal orderliness: the disciplined subjection to the well-being of the greater good, the self-sacrificial altruism of the nonreproducing workers, the dissolution of the individual in the anonymity of collective purpose, the efficient disposal of lives not worth living, the dedication to a civilizational temporality. And perhaps what also drew Bergdolt to the hive was the brute visuality of that bounded world, self-sufficient and regimented despite its teeming energy, so evocative of totalitarian aesthetics.

Unlike his Nobel co-laureate Konrad Lorenz, who was not only an active member of the Nazi Party but also a key figure in its Office for Race Policy, von Frisch—as Bergdolt realized—had little interest in the larger analogy.42 Where Lorenz explicitly established racial-hygiene correspondences between the degeneration of domesticated animals and the decline of the civilized human races, von Frisch most often restricted his own editorializing to remarks on the majesty of the bees’ senses. In this period, instinct for Lorenz had a particular meaning, in which “instinctive action”—common to humans and other animals—is directed to the preservation of the species and the category “species” is homologous to the Volk. Evolution, he maintained, is imbued with moral purpose, selection operates at the level of the community, the subordination of the individual is a social good, and further, the elimination of individuals of “inferior value” is a social necessity. These ideas were also promoted by Alfred Ploetz and the Nordic strand of German Rassenhygiene that underwrote Nazi race policy. And Lorenz drew them even more directly from Ernst Haeckel’s 1874 idiosyncratic Anthropogenie oder Entwickelungsgeschichte des Menschen (The Evolution of Man), in which social insects in their hive are a model for the relationship between citizen and state.43 Lorenz’s eagerness to buttress such notions with the authority of science was appropriately rewarded.44

It’s no wonder that Bergdolt was dissatisfied with von Frisch’s bees. Instinct, which could so easily be cast as the motor of racial progress, was decidedly muted in their hive. Only rarely does genetics escape mediation by consciousness.45 In contrast to Lorenz’s persistent diminution of animal capacity—what appears to be intentional action is revealed again and again as merely, if complexly, mechanical—von Frisch’s work, focused principally at the level of individual behavior, is motivated by a spirit of valorization, in which the dominant registers are affinity and wonder. (Could we call it a humanism generous enough to include the nonhuman?)

Von Frisch’s status as a founder of ethology rests on his revelatory account of the sensory world of the animal. It was an account that called into question simplified stimulus-response models of animal behavior and reoriented debates on animal cognition toward sensory complexity.46 In contrast to his behaviorist antecedents, von Frisch directed attention to the mind of the animal, not merely to its external expression. His honeybees—“the most perfect insects with sheer incredible instincts”—are conscious, purposeful, able to learn, and capable of making decisions.47 His language of language is far from accidental. There is, he leaves little doubt, a species of subjectivity in this species. It’s a simple claim with complex implications. Perhaps the best way to follow them is through the famous research that von Frisch’s student Martin Lindauer carried out on honeybee nest selection.48

When the population has grown and the hive is crowded, when nectar is abundant, stores are full, and the foragers are unable to pass off their loads, the bees prepare to swarm. The queen stops laying eggs, and workers tending the larvae begin feeding royal jelly to those they have selected to take her place. The foragers, for their part, stop collecting foodstuff and start searching abroad for cavities, inspecting holes in trees, buildings, and any other likely spots. Within days, the old queen exits the hive, escorted by about half the workers, perhaps 30,000 bees, leaving “house, honeycomb, and food supplies behind for her successors,” as Lindauer wrote. They settle, all one living cluster, often on the branch of a nearby tree.49

The foraging workers fly missions from this temporary home, still scouring a huge area but looking now for potential nests that meet a set of precise criteria: appropriate size, small and well-positioned entrance, protection from wind, sufficient distance from the original colony, dry, dark, free of ants. And as they locate possible sites, they return to the cluster and—just as they do for food sources—they signal their discovery by dancing, though now they perform on the massed bodies of their assembled fellows.

Lindauer observed this behavior and realized that the returning foragers were dancing but no longer exchanging nectar or pollen. He identified and marked them, interpreted their dances, mapped the coordinates they were pointing to, and on reaching the designated locations found that, instead of collecting from flowers, the bees were “busily inspect[ing] holes in the ground, hollows of trees, or a crack in an old wall.”50 These foragers, he realized, were now “house hunters.” This is his description of what happens as they return to the cluster:

If one follows for some time the dances of the scouting bees in the cluster and records their announcements of location, one comes to a very surprising conclusion: not just one nesting place is reported, but rather announcements are given of different directions and distances, and this means that several possible dwellings are announced at the same time. For example: on 27 June 1952 I noticed a dance in a cluster, which reported a nesting place 300 meters to the south. A few minutes later, a second dance could be observed which announced another nesting site 1400 meters to the east. In the next 2 hours, five more announcements came in, from the northeast, north, and northwest, with varied reports of distance…. On the next day 14 new reports of locations of nesting sites were added, so that now there were 21 different possibilities to choose from. The scout bees showed at the first glance that they had inspected various quarters: some were powdered over with dried dirt, because they had burrowed in a hole in the ground; others came from a cave in a ruin and were covered with red brick dust; once these seekers of quarters were soiled with soot, having discovered a suitable nesting place in a narrow chimney that was not in use during the summer.51

So how are these options weighed? With just one queen—a queen who may be old and weak and have difficulty flying—the swarm must hold together. To avoid disaster, it must reach not only decision but consensus. Yet this does not always happen. If a suitable site cannot be found, the bees may simply nest in the open, consigning themselves to certain death, victims of predators or the first frost. If, on the other hand, two cavities generate more or less equal interest, the swarm may split, each group following a different faction but only one containing a queen. Ultimately, the other will have no alternative but to end its secession and rejoin the swarm, often while both are still in flight to their new home.52

At this moment of danger, the survival of the colony depends entirely on the scouts. Nest selection, Lindauer discovered, as well as location, is up to them. They are both dancers and followers. Yet how these scouts “elect themselves” from the other foragers and persuade the rest of the swarm to follow them is still not known.53

Just as for nectar and pollen, the intensity of the dance corresponds to the appeal of the resource. A lively dance, a dance that indicates a nest of superior quality, can last for hours, its length and energy making it evident to a large number of scouts. The overall dancing in the swarm continues for days—up to two weeks even—and as it progresses, the number of cavities performed declines. Finally, if all is well, an overwhelming majority of the dancers will be proposing the same location, and any remaining “dissenters” are then disregarded.54 A general excitement seizes the colony. With the queen at its center, the swarm takes wing for its new home.

But there is more. In the first place, as the debate unfolds, the scouts revisit and redescribe their cavities. And their opinions are likely to change. On return trips, they might find their site less appealing—it leaks in the rain, ants have moved in, a change in wind direction has made it vulnerable. If so, the enthusiasm of their dancing declines, and quite possibly they shift their support to a competing option.

Observing marked house hunters, Lindauer realized that those bees that dance with relatively little energy for one site are likely to eventually transfer their allegiance to another, more popular location. Flexible and open to persuasion, scouts invest decision making with appropriate seriousness. Rather than taking their fellow dancers at their word, they visit a number of sites in person, making inspections for themselves. Nor do they restrict themselves to the most popular options. Scouts attend a variety of dances and visit a range of proposed cavities. Only then, armed with comparative evidence and eyewitness authority, do they make a final decision on how to cast their lot.55

To James Gould and Carol Gould, this interaction illustrates “the basically democratic nature of certain colony activities.”56 For Donald Griffin, “These exchanges of dance communication resemble conversational exchanges.”57 They have the back-and-forth quality of a committee meeting, he suggests. And I, too, am impressed by the effectiveness and suitability of the process through which this life-or-death decision emerges, as well as by the mental subtlety it reveals. It is not easy to dismiss the insistence on determination and confirmation, the allowance for change, the hesitations and doubts, the willingness to reevaluate, the calculus of commitment and compromise, the comparative method.

But what kind of language is this? And what types of conversations does it make possible? We know the scientists will gladly speak on the honeybees’ behalf. But can these tiny insects truly speak for themselves?


Despite his eighty-seven years, von Frisch traveled to Oslo in the winter of 1973 to receive the Nobel Prize. In his lecture he recalled his life’s work—his science, his bees, his colleagues—but he said nothing about his language of language. It was only his title that offered a clue: “Decoding the Language of the Bee.”58

This was characteristic reticence. Along with his wonder at the bees’ capacities was a reluctance to move from documentation—from the tooled-up natural history in which his new bees could be simply displayed for admiration—to a more reflectively theoretical mode in which those capacities could be assessed, evaluated, perhaps found wanting. In fact, it was through this reserve that the bees’ linguistic lives became self-evident in his work. And it was through this silence that his analogy gained its effective concreteness—even if, more often than not, he took particular care to cover that special word language within the uncertain shelter of quotation marks.

So he is cautious. The bees have “language” but never speech. They do not talk (though he listens and understands). And when he describes Lindauer’s research in Asia and Africa on the evolutionary lineage of bee communication as a “comparative philology” of Apis “dialects,” he is pursuing the established plot. The terminology is descriptive, the comparisons will not reach beyond the honeybees, and the Latinate pretensions evince more than a little self-mockery.

But, although sometimes he seems like a scientist from a different era, he is also accomplished in the registers of theoretical biology, so distinct in tone and ambition, and he can wield them to address a different set of abstractions. In 1965, for example, he completes The Dance Language and Orientation of Bees, the summary statement of his research. Forced by the occasion to confront the ontological question in its fullness, he uses the preface to affirm unequivocally the limits of analogy: “Many readers may wonder whether it is proper to call the communication system of insects a ‘language.’ The use that is made here of this word must not be misunderstood, as though what bees inform one another of were to be regarded as the equivalent of human speech. In its wealth of concepts and its articulate mode of expression the language of man stands on quite a different plane.” The language of the bees, he concludes in his clearest statement on the matter, though “unique in the whole animal kingdom,” is actually a “precise and highly differentiated sign language.”59

But this may be less restrictive than it at first appears. Von Frisch was writing at a moment when sign language promised a key to the otherwise inaccessible nonverbal mind. In this spirit, he introduces to the hive a wooden bee—his own prosthesis—and manipulates its movements, hoping that if he speaks their language, his bees will respond. The object’s followers express curiosity, but they are not fooled. “The model,” von Frisch acknowledged, “evidently lacked some significant characteristic without which it could not be taken seriously.”60 The bees know it is an alien. They attack and sting it repeatedly.

Meanwhile, across the Atlantic, the psychologists R. Allen Gardner and Beatrix Gardner, preoccupied by the evolution of cognition, were preparing to welcome Washoe the chimpanzee into their Nevada home, to raise her as their human girl child and teach her American Sign Language. In an attempt to render vulnerable to empirical investigation Wittgenstein’s insight “If a lion could talk, we could not understand him,” the Gardners reversed von Frisch’s procedure and set out to prove that nonverbal animals can acquire human language and use it to communicate with one another and with their trainers.61

But Wittgenstein’s lion, as the animal philosopher and trainer Vicki Hearne pointed out, is not without language; he is just “not talking.”62 His muteness proposes an irreconcilable difference, an indifference that refuses to be tamed, a fullness, not a lack; a “consciousness that is beyond ours,” Hearne called it.63 And yet this phenomenological abyss is exactly what von Frisch dared to cross—though less by code breaking than by the projection of his (and Lindauer’s) most intimate longings. Because when forced to succumb to the language of science, to surrender the language of bees, he, too, was reduced to speaking in code.

Honeybees, like Wittgenstein’s lion, don’t talk to us. Instead, von Frisch taught us how to eavesdrop on them. And—in a whisper—he told us, too, that even if their “dance language” exhibits the automatic quality of the code, we should not assume that those signals we can access encompass their communicative world.

Of course this isn’t merely a struggle over what the animal means; it is a struggle over the meaning of the animals themselves. And it is a battle long fought on the terrain of language. Though no philosopher, von Frisch understood this only too well. Language—the absence of language—continues to define the subor-dination (rather than simply the difference) of the animal in post-Enlightenment Western philosophy, a tradition remarkably Cartesian on this question.64 Could von Frisch be clearer? This “dance language” is restitutive, an appeal to an ethic of mutuality and recognition, a call for respect for the nonhuman animal—for the animal in general and the astonishing honeybee in particular.

“It took some ten years of patient observation,” wrote Jacques Lacan in the wake of the Brunnwinkl experiments, “for Karl von Frisch to decode … [the bees’] message, for it is certainly a code, or system of signaling, whose generic character alone forbids us to qualify it as conventional.”65 As code is to language, Lacan wants us to understand, so nature is to culture and animal is to human. Hardwired, hard-pressed, the bees stand for a programmed, mechanical nature in vivid contrast to the complex spontaneity of human culture.66 Indeed, they enable this line between animal and human, between nature and culture, to be drawn with some severity.

The argument is not new: animals can sign, but they cannot lie. They can react, but they cannot respond.67 They can communicate, but they cannot participate in the second-order metacommunication so familiar to humans. They cannot signify about signifying, think about thinking, and nor, for that matter, can they “dance about dancing.”68

It is a conventional claim, this humanist insistence on language lack in the animal. And its framing in such irreducibly human terms makes it impossible to disprove (though not to dispute: in the cooperative setting of the hive, for example, it is difficult to imagine why a bee should be moved to dissemble about the location of a feeding site, and, anyway, wasn’t it the bees’ “honesty” that so appealed to Lindauer?).

But the point is not to make the bees speak, to have them tell us their secrets as the Gardners would have liked poor Washoe to tell them hers. Nor is it to imagine that the little honeybees are somehow just like us, that their world somehow corresponds to ours, that to be a bee is somehow equivalent to being a human equipped with a different sensory apparatus. That somehow our shared evolutionary origins, our intertwined deep histories, provide us also with a shared ontology.

Instead, can it be enough to point out that the honeybees’ repertoire exceeds functional explanation and biochemical predictability, that the more researchers find out about honeybee cognition and behavior, the less appropriate and effective is the metaphor of the machine? In this instance, at least, it seems that language (or its absence) is an inadequate marker of interiority. And it seems that the assumption that language, human language, is “an unprecedented inferential engine” is itself a product of linguistic circularity—a product that tells us more about the making of the animal in language than about the living animal that is the ostensible subject of science.69

What, in such terms, can we make of the bees’ “spasmodic dance,” which is “more the expression of a dancing mood than an effective signal”? Or of the “trembling dance,” which according to von Frisch “tells the bees nothing” yet manifests in times of stress and appears to mark some kind of “neurosis”? Or of the “jerking dance” that he considers “an expression of joy and contentment”?70 Or, indeed, of the nest dances described by Lindauer, each of which intervenes in a larger social process of decision making?

But these are murky waters. Like von Frisch, I prefer to avoid this treacherous and much-debated question of language and cognition. The terms are too literal. The cards too stacked. The conflation of difference and deficiency too pervasive.

Like so many others, Lacan holds fast to the promise of that boundary between code and language, the promise of escape, a way to leave behind the animal to arrive as a thoroughly human subject. And in the inclusive corner, Griffin’s cognitive ethology, with its principled determination to restore dignity, agency, and consciousness to the animal through methodological and theoretical humility, arrives at a troubling humanism of its own, a “giving speech back,” conferring minority rights on the animal as on the thinking child, an uncanny recapitulation of the history through which colonial hierarchies were made.71

Such is von Frisch’s dilemma. He knows his bees do not speak as humans; he knows their “language” is less but also more than his own. And he knows his new discipline allows space only for the less. Where within the rationality of his science could he find a language to express the profound commonality of life and the irredeemable fact of shared mortality? Where could he find its double, a language to communicate a difference for which no words exist? And where could he find a language in which to understand the absence of language as something other than a lack?

(Pity the animal that lives only as a shadow of the human, the animal forced to react rather than respond, the animal whose task is to give flesh, spirit, and meaning to the human, the animal whose melancholy fate is to be humanity’s other.)


“There is really no reason to suppose,” says W. G. Sebald’s Austerlitz, “that lesser beings are devoid of sentient life.”72 Remembering childhood nights, he wondered, Do moths dream? Do they know they are lost when, misled by the flame, they enter a house to die?

What was von Frisch’s question: Can the honeybee speak? No, that wasn’t it. First he thought, There is really no reason to suppose that honeybees are devoid of language. And then he asked, My little comrade, what does she say?

Pity the honeybees. Pity and protect them. In this, too, their indifference is of no avail. So trapped in language. The honeybees and us, held together, pushed apart. Even von Frisch, even Lindauer, who loved them so dearly, who found in them redemption from the brutish horrors of their times … Well, do you remember what they’d do to prove their little ones’ capacities?

But enough paradox. To give them language was simultaneously to celebrate their difference and to doom them to impossibility, to condemn them to the merely imitative, at which they could only fail, to (mis)take “linguistic self-referentiality [as] the paradigm for self-referentiality generally.”73 But of course the failure is human (a specific scientific human, perhaps, but human nonetheless), this failure of being able only to imagine sociality and communication through something language-like and to grant ourselves its apogee. What foolishness to judge insects—so ancient, so diverse, so accomplished, so successful, so beautiful, so astonishing, so mysterious, so unknown—by criteria they can never meet and about which they could not care! What silliness to disregard their accomplishments and focus instead on their supposed deficiencies! What pitiful poverty of imagination to see them as resources merely for our self-knowledge! What sad, sad, sad sadness when language fails us.