Cover image for Cheating monkeys and citizen bees : the nature of cooperation in animals and humans
Cheating monkeys and citizen bees : the nature of cooperation in animals and humans
Dugatkin, Lee Alan, 1962-
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New York, NY : Free Press, [1999]

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xi, 208 pages : illustrations ; 23 cm
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QL775 .D84 1999 Adult Non-Fiction Central Closed Stacks

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Here biologist Lee Dugatkin outlines four paths to cooperation shared by humans and other animals: family dynamics, reciprocal transactions (or "tit for tat"), so-called selfish teamwork, and group altruism. He draws on a wealth of examples--from babysitting among mongooses and food sharing among vampire bats to cooperation in Hutterite communities and on kibbutzim--to show not only that cooperation exists throughout the animal kingdom, but how an understanding of the natural history of altruism might foster our own best instincts toward our fellow humans.

Author Notes

Lee Dugatkin is Professor of Biology, University of Louisville

Reviews 3

Booklist Review

Behavioral ecology studies of the evolution of cooperation in animals are useful, Dugatkin argues, because they supply "a stripped-down version of what behavior in a given circumstance would look like without moral will and freedom." He describes "four paths to cooperation" identified by behavioral ecologists--family dynamics, reciprocal transactions, selfish teamwork, and group altruism--and then discusses each in terms of dozens of examples, from ants and guppies to squirrels and lions. In analyzing animal behavior, Dugatkin endeavors to isolate the factors that make cooperation an appropriate and appealing behavior for a particular species--in order to consider whether the same factors may apply to human situations. He also discusses the history of the science and philosophy of cooperation, with comments on Locke and Hobbes, Darwin and Wallace, Huxley and Kropotkin, among others; in his last chapter, "Possibilities and Pitfalls," he discusses attempts to apply these animal patterns to human behavior. Dugatkin's chatty, accessible style should appeal to fans of PBS nature specials. A thought-provoking approach to the cooperation-competition battle. --Mary Carroll

Publisher's Weekly Review

Evolutionary biologist Dugatkin (Cooperation Among Animals) is unabashed in his belief that "the study of evolution and animal behavior can be used to foster and enhance cooperation in humans." Without resorting to simple minded biological determinism, he argues forcefully that the behavioral predisposition of humans may be predicted by evolution. Thus, he asserts that research in animal behavior can provide baseline information about parallel behavior in (admittedly more complex) humanity. Such investigations may ultimately help us better understand the underpinnings of human behavior and allow us to restructure our environments to promote more cooperation. Dugatkin explains that cooperation arises through four pathways, "family dynamics, reciprocal transactions, selfish teamwork, and group altruism." He devotes one chapter to each pathway, clearly explaining the underlying evolutionary theory and providing myriad animal examples. His fascinating instances range widely from vampire bats willing to regurgitate blood for starving neighbors to mongooses who take turns baby-sitting. Each chapter concludes with an attempt to tie the lessons learned from animals to suggestions for public policy issues as diverse as class size in elementary schools and partnering in police departments. These applications, however, are the weakest part of an otherwise startling and eye-opening glimpse into the evolution of behavior. (Feb.) (c) Copyright PWxyz, LLC. All rights reserved

Choice Review

In light of the Darwinian imperative, cooperation seems out of place. Can cooperation between people, widespread but still cherished when it occurs, be explained using evolutionary biology? The short answer is yes, the long answer is "read this book." Expanding on his own research involving guppies (but not belaboring it), Dugatkin reviews our current understanding of the evolution of cooperation. He draws meaningful parallels from other animals to the human species. These comparisons are not overly ambitious, nor do they stretch credibility by interpreting all current human behaviors as being shaped by our evolutionary history. Nevertheless, the extensive chapter notes give a flavor for modern behavioral science, with all its grandstanding and heated debates, and they offer the reader an annotated bibliography that encourages further exploration. Though advanced students will find many of the examples a bit threadbare, those just being introduced to the field will find them classic and described in a casual manner. An excellent introduction to a fast-changing field. All levels. G. Stevens; University of New Mexico



Chapter One All in the Family And the man knew his wife Eve; and she conceived and bore Cain ... and again she bore his brother Abel.... And in the process of time it came to pass, that Cain brought of the fruit of the ground an offering to the Lord. And Abel, he also brought of the firstling of his flock and of the fat thereof. And the Lord had respect unto Abel and his offering, but unto Cain He had no respect. And Cain was very wroth, and his countenance fell. And the Lord said unto Cain: "Why art thou wroth? and why is thy countenance fallen? If thou doest well, shall it not be lifted?" --Genesis 4: 1-7        According to the Old Testament, the first human siblings were not particularly fond of one another. Fair enough--we all know of cases of sibling rivalry, not to mention the animosity that we might hold for some of our more distant relatives who pop in for the occasional holiday dinner. But, in general, blood really is thicker than water despite tales of murderous brothers. Later in the story of Cain and Abel we encounter a poignant question: "And the Lord said unto Cain: where is Abel thy brother? And he said `I know not; am I my brother's keeper?'" The answer, at least in terms of how we behave, is a qualified yes--you are indeed your brother's keeper.     If your brother and a stranger were drowning, who would you save first? The reply most of us would expect--"my brother, of course " suggests, but certainly does not demonstrate, the importance of the role of kinship in structuring human cooperative acts. Is it possible, however, that the reason we might choose our brother has nothing to do with the fact that he is a blood relative per se? Is it merely the fact that we have spent so much time with our siblings that drives our actions? A simple thought experiment might help us to understand whether this true. Imagine it is not a stranger with your brother there in the water, but your closest friend. How would you feel in that case? How would most people react? If kinship was the overarching theme, most should still reply "my brother, of course." This train of thought has led behavioral ecologists to appreciate just how important kinship is in the human social dynamic. Family Accounting Schemes The scenario in which you save your brother or someone else functions as a nice illustration of how kinship affects human cooperation, but it is not something one envisions as a starting place for significant scientific breakthroughs. Or is it? Evolutionary biologists' first introduction to the notion that blood relations affect social behavior actually came in the 1930s in a form similar to this example. It was then that J. B. S. Haldane, a founder of modern evolutionary theory, suggested that he would risk his life to save two (but not one) of his brothers and eight (but not seven) of his cousins. Haldane, quite versed in mathematics, made this rather bold statement by counting copies of a gene that might code for cooperative behavior. Such a gene-counting approach to kinship and the evolution of cooperation has been extended by theoreticians but in its most elementary form is the heart and soul of kinship theory. Let us see how this idea works and how it has been formalized into what is known as kin selection or inclusive fitness theory.     The evolutionary biologist's definition of relatedness and kinship may strike many as surprising, if not odd. In such a definition, relatedness centers on the probability that individuals share genes that they have inherited from some common ancestor (parents, grandparents, etc.). A jargon phrase summing up this approach in behavioral ecology is "identity by descent." For example, you and your sister are kin because you share some (in this case many) of the same genes and these have been inherited from common ancestors, mom and dad. Similarly, you and your cousins are kin, because you share genes in common (not as many as siblings)and common ancestors, your grandparents. Common ancestors are the most recent individuals through which two (or more) individuals can trace genes that they share in common.     Once we know how to find the common ancestry of two or more individuals, we can calculate their relatedness, which simply amounts to the probability that they share genes that are identical by descent--genes that have been inherited from a common ancestor. In the literature on kinship, this probability is often labeled r (for "relatedness"). For example, you and your brother are related to one another by an r value of 1/2.     From a "gene's-eye" perspective, calculating relatedness is the first critical step in understanding how kinship can favor cooperative behavior among individuals. Genes' survival depends on the number of copies of themselves that they get into the next generation. This is often thought of in terms of what effect a given gene has on the individual in which it resides, but relatedness suggests that this is a myopic view. If relatives have a high probability of sharing a given gene, then that gene can potentially increase its chances of getting more copies of itself into the next generation by coding for some behavior that helps relatives. Again taking a gene's-eye view, relatives are just vehicles who are likely to have copies of you (the gene in question) inside them as well. But, and this is a big "but," relatives only have some probability ( r ) of having a copy of, for example, a gene for cooperation. A gene in sibling 1 "knows" that a copy of itself may reside in sibling 2, but only with a 50 percent probability. The more distant the relative, the less likely a copy of the gene resides in them as well. So, phrased in the cold language of natural selection, relatives are worth helping in direct proportion to their relatedness. This is because relatedness is a measure of genetic similarity, and genes are the currency of natural selection.     Behavioral ecologists are not so foolish as to assume that animals are able to calculate relatedness in the manner described above. We only assume that natural selection favors individuals who act in ways that make it appear as though they are able to make such calculations. How animals determine who is kin and who isn't is a matter of some debate these days. For example, one theory suggests that animals determine relatedness by matching a suite of traits (a template) that they possess against the same suite of traits in another individual. Depending on the degree to which traits match up, individuals are treated as full siblings (if many matches occur), half siblings (if fewer matches occur), cousins, and so on, down to the category "unrelated individual" (if, for example, no matches occur). Such "matching games" have their flaws; mistakes can be made in determining the level of overlap, and some relatives may erroneously be treated as nonrelatives, while some nonrelatives may be viewed as relatives. Often, however, rather than a suite of traits, a single characteristic is used to determine whether another individual is kin and if so, what type of kin. In many insect species, for example, kinship is assessed by odor. Individuals who smell like you (or your nest) are relatives, and how closely related they are is determined by how similar their odors are to yours.     While most behavioral ecologists accept that such matching (either of many cues or a single cue) is important, they believe that there is another, simpler explanation for how animals determine who qualifies as kin, an explanation that I'll refer to as the "no place like home" hypothesis. Under this hypothesis, animals simply treat all others that grew up in their nest (territory, burrow, etc.) as relatives. This very simple rule is often quite powerful. With the exception of some species that try to trick other species into raising their offspring, the odds are quite strong that those who grew up in your nest are in fact your siblings and parents.     The details of how animals evaluate relatedness are fascinating, but all we really need to know to examine kin-selected cooperation is that many animals do in fact behave in ways that allow them to distinguish between kin and nonkin and even to distinguish between different degrees of relatedness. Once we have calculated relatedness, we are very close to reaching a general rule for when cooperation among relatives should be favored and when it should not. We need only consider two more factors: the cost of the action to the individual cooperating and the benefit to the recipient of such a cooperative act. Let us call the cost of a cooperative act to the donor c , and the benefit to the recipient b . In 1964, W. D. Hamilton (now at Oxford University) showed that cooperation among relatives should evolve when the following holds true: r X b [is greater than or equal to] c .     In other words, cooperation among relatives is favored if, and only if, the benefit of the act multiplied by the relatedness of the actors is greater than or equal to the costs ([is greater than or equal to] is read "is greater than or equal to"). This equation, r X b [is greater than or equal to] c , has become known as Hamilton's Rule. Essentially, Hamilton's Rule says the following: There is some cost ( c ) that "must be made up for" if the gene for cooperation is to evolve, as cooperating with others is often a risky business. One way to make up for this cost is through the benefits ( b ) a relative receives, because relatives may carry the gene for cooperation as well. But, relatives have only some probability of carrying the cooperation gene and so the benefits received must be devalued by that probability. If I pay a cost for undertaking an action, but there is only a probability that I will receive indirect benefits (in this case through my relatives), I need to factor that into my equation and that is just what r does.     We can illustrate the use of relatedness to predict cooperation among kin with a simple chart. Consider an action that you take that reduces your chances of survival by 50 percent (a very serious cost) but increases the probability of survival of the relative(s) you are trying to save by 50 percent each (a considerable benefit). Such extreme costs and benefits might, for example, mimic a situation in which you scream out when a gang of armed thugs is approaching. This serves to announce the presence of thugs to the relatives around you, but at the same time the scream draws the marauders' attention your way, a dangerous action indeed. Based solely on kin selection theory, the table shown here outlines the number of relatives that need to hear your scream before natural selection alone would favor such dangerous behavior on your part. IS SCREAMING AT THUGS WORTH IT? Number of relatives who must hear Relative Degree of relatedness your screams Sibling 1/2 [is less than or equal to] 2 Parent 1/2 [is less than or equal to] 2 Grandparent 1/4 [is less than or equal to] 4 Grandchild 1/4 [is less than or equal to] 4 Uncle/aunt 1/4 [is less than or equal to] 4 Cousin 1/8 [is less than or equal to] 8 Spouse 0.0 --     The table illustrates the fundamental point of inclusive fitness theory: the greater the degree of relatedness between individuals, the more likely that kin-selected cooperation is selected. There need only be two (or more) siblings around for you to make that scream, but you'd need eight or more cousins present (a much less likely event), if they were the only relatives in the vicinity! How exactly, though, do we use Hamilton's Rule to come up with the correct number of relatives in the table? Consider the case for siblings. If a single sibling hears an alarm call, then r = 1/2 and b and c are still each 1/2. In that case r multiplied by b is not greater than c , Hamilton's Rule is not met, and cooperation via kinship is not favored by natural selection.     Suppose, however, that three siblings hear the alarm call. Now b is tripled (three recipients), but c is the same (the alarm call still draws the predator's attention), so r X b = 1/2 X (1/2 X 3) for a total of 3/4, which is greater than c , and Hamilton's Rule is satisfied. The same logic can be applied to any relative in the table (or for that matter, any relative not in this table). Take note, as well, that the relatedness of an individual to his/her spouse is 0 (with the exception of marriages among relatives). Although one's spouse is kin in the everyday usage of the term, we don't generally share genes inherited from a common ancestor with our spouses and hence this category of relative is in effect removed from kin selection theory.     Of the four paths to cooperation that we will focus on, kinship is the best understood, most accepted, and least controversial. It is in every legitimate textbook on evolution and is cited in more papers in the field than any other set of theories. There is even a belief among some evolutionary and behavioral biologists that Hamilton's work in this area marks the start of the modern discipline of behavioral ecology. But even kin selection theory is not without its controversies.     One area of contention with respect to kinship and cooperation centers on whether it really matters where the genes we are counting are located. Kin selectionists correctly argue that blood relatives are more likely to carry the same gene than are individuals drawn at random from a population. But what if some other mechanism besides kinship could create groups in which individuals were all likely to carry one or more genes coding for cooperation? Does it really matter that such individuals don't share other genes, like kin do? After all, we are interested in the gene(s) coding for cooperation, and everything else is in some respects background material for that gene. Who cares whether individuals carry the same genes because of kinship or for some other reason--shouldn't the process by which cooperation is selected for work just as well in both cases? The answer to this question, as Hamilton himself noted, is yes, the process works the same; whether individuals share the gene(s) for cooperation because of relatedness or some other factors is irrelevant.     Yet kin selection advocates are not so fast to roll over. Sure, mathematically speaking, you are right, they say, but in practice the distinction we are arguing about is still real and important. Give us, they say, a good example of how individuals sharing a gene are brought together, if relatedness (which automatically brings them together) is not in force. The answer typically given by kin selection-critics is that individuals that share a gene for cooperation may gather together specifically to be near other cooperators, because cooperators do particularly well when around others like themselves and so should choose this option, when it becomes available. "Be specific," say kin selectionists, "give us a real example." And this is where the kin selectionists start looking a bit better than they did after losing the mathematics argument, because behavioral ecologists are usually stopped in their tracks when it comes to finding a good animal example to answer this question.     Although such examples may be hard to uncover in animals, those interested in enhancing human cooperation argue that the evidence for cooperators choosing other cooperators as partners in our own species is anything but scarce--even when kinship is not in play. How others will act is one primary means by which we choose with whom we will interact. So, for humans then, while kinship is an extremely important force selecting for cooperation, there are many other ways cooperators may cluster together aside from kinship. We need to recognize this in our behavioral studies and our conjectures about human cooperation.     The above controversy is admittedly a semantic one in part, but semantic arguments can be quite illuminating. Hamilton's Rule--which in words roughly translates to "all else equal, cooperation should be most common among close relatives"--is as close as behavioral ecologists get to a "law of nature." It is an underpinning of all modern evolutionary approaches to social behavior and is, in many ways, as much an approach to behavioral biology as it is a theory. The data gathered to date certainly support the claim that Hamilton's Rule is extremely powerful. It is not a "law" in the sense that gravity is, but it is about as near to one as behavioral biologists can hope to come, given the astonishing complexity and variability that is an inherent part of the subject matter they tackle.     From the standpoint of reputation, Hamilton's Rule was quite good for the field of behavioral ecology, at least in one sense. While solid mathematical theory has been part of evolution since the seminal work of J. B. S. Haldane, Ronald Fisher, and Sewall Wright in the 1930s, it was not truly a centerpiece of evolutionary approaches to behavior until Hamilton's Rule. For many in the field of behavior, there was an unspoken envy of the hard sciences (physics, chemistry, even other parts of biology) that had steadfast "rules" that could be written out for skeptics (not to mention funding agencies). Hamilton's Rule provided such ammunition to behavioral ecologists. Let's take a look at some examples of why this is so, with a few cases from the animal kingdom, before moving on to how such scenarios can help us foster human sociality. The Insect Police The so-called social insects have been a godsend for advocates of kin-selected cooperation. The reason lies, at least in part, with the bizarre genetics of social insects such as bees, wasps, and ants (collectively known as hymenopteran insects). Humans (and most other animals) are diploid organisms, which means that we have two copies of each of our chromosomes. Our forty-six chromosomes are twenty-three matched pairs. The only stages of human life that are not diploid are sperm and egg, as they have only a single copy of each of our twenty-three distinctive chromosomes. Sperm and egg then are called haploid rather than diploid. Of course, sperm and egg later fuse to form diploid animals.     Much of life on earth, such as bacteria and viruses, is always in the haploid phase. Why some life on earth is diploid and some haploid is a fascinating question, but not one critical to the issues we are examining. What makes bees, wasps, and ants so bizarre is that females are diploid and males are haploid--a genetic system known as haplodiploidy. What this means is that when a male fertilizes a female, only daughters are produced because the sperm and egg fuse to produce a diploid creature, and in most social insect species diploids are female. Females, however, produce sons from unfertilized eggs (eggs that have not fused with sperm)--which means that sons never have fathers!     Haplodiploidy creates some very strange scenarios. In diploid and haploid creatures, relatedness between two individuals is symmetric; that is, if a father is related to his daughter by an r of 1/2, then a daughter is related to her father by the same value. Not true for the social insects. To see why, focus your attention on the father/daughter relationship. Fathers are haploid and give a copy of each chromosome they have to their daughters. Hence fathers are related to daughters by a value of 1. Daughters, however, are diploid, in that they get one copy of each chromosome from each parent, both mom and dad; so a daughter's relatedness to her father is 1/2 (half her chromosomes come from dad)--fully half of her father's relatedness to her.     The most relevant effect of the strange genetics of the social insects is its impact on average relatedness within insect colonies. Before seeing this in detail, keep in mind that in many social insect colonies a single queen produces all the offspring for a group. This means that the vast majority of individuals in such colonies are sisters and brothers. Haplodiploidy has the twofold effect of making sisters "super-relatives" and making the relatedness between brothers and sisters only half of what it is in diploid brothers and sisters. Sisters end up with a relatedness value of 3/4 (50 percent greater than the same relationship in diploid species), and sisters are related to brothers by a value of 1/4 (half the value found in diploid creatures). So, a clear prediction from kin selection theory is that since females are much more related to their fellow colony members than are males, when colonies have more females than males (as in most social insects), cooperation should occur predominantly in this sex. And of course it does, as "workers" in insect colonies are almost always female! It is females that sacrifice their lives by stinging folks and ruining an otherwise pleasant summer day. It is also females that undertake virtually all of the everyday activities that keep a colony functioning--food gathering, care for the young, and so on. One particularly interesting and unique behavior found among female social insects is "policing" behavior.     Bee colonies are rightly thought of as models of both efficiency and harmony. It is mind-boggling what a colony of tiny insects can accomplish in a short period of time: regulating the temperature of a hive, caring for young, defending against many predators, finding food, recruiting others to join in bringing back the booty, and a myriad of other activities. Some of this efficiency (and harmony) has been attributed to a single queen often producing all of the eggs for a colony, thus allowing worker females to spend their time on other hive-related necessities. Queens accomplish this enviable task by using a barrage of chemicals to inhibit other females--the workers from reproducing. Yet, as with any chemical inhibition system, it is inevitable that some workers will escape these anti-aphrodisiacs and thus will have a much greater chance of reproducing than their subdued sisters. Once this fascinating new door is opened, we can ask whether kinship theory can guide us with respect to a rather nasty question: should the eggs laid by workers that ignore the queen's chemical castration cues be left alone by their sisters or vigorously attacked? The answer to this question is rather personal, if you happen to be the queen, as it depends on how many males you opt to mate with.     The relatedness between individuals in an insect colony depends on how many males inseminate the queen. The more males the female mates with, the more different lineages there are in a colony--each line's ancestry going through the queen and a given male. Once again, however, the strange genetics of such insects creates a novel situation. Rather than showing a family tree more complicated than that of the British monarchy, it can be shown that if the queen of a colony mates with a single male, then female workers in the colony turn out to be more related to nephews than to brothers. If the queen mates with numerous males, however, that situation reverses itself and female workers in a hive are now more related to brothers than to nephews. We shall focus on the second scenario because of the fascinating kin-selected cooperation emerging from it.     Whether females in a social insect colony are more related to brothers than to nephews can have quite serious implications about when and whether we should see kin-selected cooperation, and if it exists, what form such cooperation should take. To see this, first recall that brothers are those individuals produced by the queen, while nephews are those produced by sisters that have somehow managed to avoid the queen's chemical anti-reproduction agent. A conflict of interest then arises between sisters that have managed to escape and those that have not.     Aside from the queen, females who can reproduce (i.e., those that do not fall victim to the queen's attempt to monopolize reproduction) are always selected to do so. When females reproduce they always produce males, since such females are almost never inseminated. This creates a problem, however, for those females who can't reproduce, as they are more closely related to the queen's offspring (their brothers) than to their sisters' children (their nephews). Kin-selected cooperation on the part of those nonreproducing female workers then favors any action that increases the odds of the queen's offspring surviving at the cost of nephews.     There is little a female can do to stop one of her sisters from reproducing, if her sister has avoided the queen's attempt to do so already. But there are options available. Once a worker has laid eggs behind the queen's back, her sisters could, for example, refuse to care for and help nephews. Or they could take more drastic action--they could eat eggs destined to be their nephews! Francis Ratnieks and Paul Visscher examined this possibility in honeybees, where females mate with ten to twenty different males. Their results were astonishing. Those honeybee females who did not produce offspring "policed" the reproductive actions of their sisters. If their sisters produced eggs on the sly, policing females destroyed the eggs. Ratnieks and Visscher found that honeybee workers showed remarkable acumen in discriminating between sisters' eggs and the queen's eggs. In a controlled laboratory setting, after twenty-four hours, only 2 percent of the sister-laid eggs remained intact, while 61 percent of the queen-laid eggs remained unharmed! But, given that the actual act of egg laying is rarely observed, how could honeybees know which eggs were laid by sisters and which by the queen? The answer appears to be that eggs are chemically "marked; such that queen-laid eggs smell different from worker-laid eggs. Why eggs should be marked so is still unclear, but one tantalizing possibility is that the queen marks her eggs to encourage workers to police the activities of their sisters.     Kin-selected policing is qualitatively different from the other types of cooperation so often found in animals. Rather than having individuals form a cooperative unit to accomplish some task, cooperation in honeybee police work takes the form of stopping others from cheating--a more subtle and complex action. At a more fundamental level, policing is powerful, because it provides a direct deterrent to cheating, whereas in many other cases, we simply rely on cooperation being somehow more profitable than cheating, and this holy grail is often difficult to obtain.     There are many other cases of cooperation in highly related social insects. I'll mention one other curious example: honeypot ants. In one species of these ants, the largest individuals actually hang from the top of a colony and act as living storage tanks for water and sugar. These "honeypot" individuals have soft and elastic abdomens, and if you watch long enough you will see other individuals come up and "turn on the faucet" to drink the resources stored there. For significant periods of time, honeypot individuals do nothing but hang from the rafters and supply this service.     It is fascinating to find policewomen and living storage bins in the insect world, but how much of the cooperation we see is strictly due to the bizarre haplodiploid genetics of social insects? Can we expect anything so dramatic among mammals? "Eureka!" Naked Mole-Rats Physics is not the only discipline in science that has "Eureka!" stories. Just as physicists can recount the bathtub adventures of Archimedes and his famous exclamation when coming up with his theory of buoyancy (specific gravity), so too can the ardent student of behavioral ecology recite the story of Richard Alexander and Jenny Jarvis's discovery of extraordinary cooperation in a bizarre creature, the naked mole-rat. Alexander, a professor of biology at the University of Michigan, traveled to various universities in the 1970s, giving lectures on the evolution of social behavior, particularly cooperative and altruistic behavior. One of his themes was why, despite significant effort, extreme sociality (like that seen in insects) had not been uncovered in mammals. Alexander described the characteristics he believed a mammalian system would need for insect-like ultrasociality to exist. He outlined a hypothetical creature that would undertake altruistic acts for relatives who lived in a safe environment with lots of food. He went so far as to give details: the species would eat large tubers (potato-like foods) and live in burrows in a tropical spot that had clay soil.     One day in May 1976, Alexander presented these ideas to some folks at Northern Arizona University. Afterwards, he was approached by someone in the audience who told him he had given a perfect description of the naked mole-rat of Africa. On the advice of this fellow, Alexander contacted Jennifer Jarvis (at the University of Cape Town), who knew more about naked mole-rats than anyone in the world. After much back and forth, which included trips by Alexander and his colleague Paul Sherman to Africa to actually see the creatures, Jarvis and Alexander realized that they indeed had found the first eusocial (ultrasocial) mammal.     After all the attention this animal has attracted from both scientists and the media, it is almost disappointing to see how bland the native habitat of the naked mole-rat actually is and how ugly these creatures are, even by rodent standards! Naked mole-rats are hairless and blind, with crinkled skin and two large incisor teeth sticking out from their mouths. And those are the adults; the babies are even harder to look at for very long. First collected in Ethiopia in 1842, naked mole-rats (whose scientific name is Heterocephalus glaber ) live within groups averaging about seventy individuals (but ranging up to almost three hundred) in underground burrows, from which they rarely, if ever, emerge. Such burrows average about two miles in length. Naked mole-rats have been studied primarily in Kenya and are often found in arid areas covered with dust and brush. Typically found near dirt roads, colonies can be located by molehills that pock the landscape. But what naked mole-rats lack in beauty and scenic living conditions, they make up for in fantastic behaviors.     One female alone (among many in the colony) is responsible for all the reproduction in a naked mole-rat group (three or so males in the group are responsible for the male side of mating). No other mammal that we know of, except another species of naked mole-rats discovered later on, has a single "queen", and this finding sent shock waves through the behavioral biology community. Kin selection theory suggests that such extreme cooperation, wherein most individuals give up the opportunity to reproduce, should be limited to species in which individuals are somehow extremely related to each other, yet naked mole-rats are mammals and don't have the bizarre genetics that allow for the "super-relatives" we saw in the bees and ants. So how could such a bizarre system have evolved here? Before answering this question, let's get a more comprehensive sense of just how much cooperation goes on among these creatures.     The queen and the handful of males she mates with have a twofold advantage over others in naked mole-rat colonies: not only do they monopolize all colony reproduction, but they also live longer than their nonreproducing colony-mates. Yet in the relatively short time that nonreproductive males and females are around, they get a lot accomplished, and without their cooperation naked mole-rat colonies would surely come to a screeching halt. In fact, those individuals not specialized in reproducing take on virtually all of the everyday cooperative actions that are the very lifeblood of colony existence. They excavate new tunnels (an absolutely critical aspect of colony survival), sweep debris, groom one another as well as the queen, and take on the unenviable and dangerous task of defense against predators.     Why such dramatic examples of cooperation in a single species? What singles out naked mole-rats? The answer probably lies in kinship within colonies. As we mentioned earlier, naked mole-rats do not have the strange genetics of some insects, but they have managed to achieve the highest average relatedness on record for naturally occurring mammals. DNA fingerprinting (the same technique we read about in criminal cases) showed that the average degree of relatedness among individual naked mole-rats in a colony was a whopping 0.81 (out of a possible score of 1). To put this in some perspective, unrelated individuals have a value of 0.0 for this indicator, brothers score (on average) 0.5, and the most related of all individuals, identical twins, score 1.0. So naked mole-rat individuals on average fall between ordinary siblings and identical twins on a relatedness scale, and even lean toward the identical twins' side of the equation. What a wonderful finding in support of kin selection and cooperation! Question: Where do we find the highest recorded degree of cooperation among all mammals? Answer: Just where kin theory says we should--in a species with uniquely high degrees of relatedness among group members.     Cooperation in the naked mole-rat even exceeds the borders of a single colony. As in many species that live underground, founding a new colony, which entails coming above the surface, is a very dangerous activity. In addition to being away from the food source and potential mates, the brave mole-rat who tries to start a new colony may find numerous predators lurking above ground. In fact, for quite some time it was believed that new colonies formed when larger established colonies split in two, thus alleviating the problems associated with a single individual surfacing to hunt for a suitable place to start a new group. It turns out that this picture is not accurate, at least some of the time. M. J. O'Riain and his colleagues found that colonies did not undergo fission to form new groups, but that specialized individuals took on the dangerous task of colony founding, thus making it unnecessary for their kin to risk life and limb themselves. Such "dispersing cooperators" put on weight very quickly during development and were considerably larger and bulkier than other colony members, presumably making them fitter for the trials and tribulations of colony founding in a hostile environment.     Before labeling naked mole-rats as the perfect example of kin-selected cooperation, we must deal with one potential fly in the ointment. One might think that in such a cooperative system, the nonreproductives would voluntarily yield reproduction. This is, apparently, not quite true for naked mole-rats. Nonreproductives are coerced into yielding the act of producing offspring by aggression on the part of the queen; that is, reproduction by the masses is suppressed, not freely handed over to a single individual. Suppression, however, may not pose as great a problem to cooperation in this system as it seems. We would expect there to be tremendous selection pressure on the nonreproductives to breed if it would increase their fitness, suggesting that direct reproduction is not the best route to increases in fitness for such individuals. In essence, what appears to be suppression on the part of the queen may be the only mechanism available to increase the fitness of both queen and nonqueen, making such apparently suppressive acts really cooperation on the part of all. This remains to be seen, but the suppression = cooperation argument certainly is an intriguing possibility.     Even aside from its Eureka-like discovery, naked mole-rat ultracooperation is truly astounding. No other mammal species has a single queen producing offspring that undertake such a wide variety of cooperative and altruistic endeavors. But then, no other mammals are as closely related to one another either. In-House Baby-Sitters How many of us wish we could convince our older child that staying home to watch a sibling is a more noble and rewarding act than going to a friend's party down the street? Finding a person to watch your child while both parents work is an even greater dilemma. Dwarf mongooses, however, seem somehow to have solved this very problem in a rather cooperative fashion, based primarily on kin bonds.     Dwarf mongooses are small, social carnivores that typically occupy dens in the savanna habitat of Tanzania, in groups of approximately ten to twenty individuals. A pack of mongooses usually consists of a breeding male, a breeding female, and their young from a number of consecutive broods (i. e., young of various ages) as well as an occasional immigrant from elsewhere. Young "helpers" cooperate with their kin in a wide variety of activities, including feeding, nest defense, grooming, and transporting the young. But the most fascinating and interesting variant of kin-based cooperation may very well be "baby-sitting."     A typical day for a mongoose pack in the Serengeti begins with the adult male and female, as well as some of the immature adults, leaving the den to search for food. But what of the almost ever-present very young mongooses? Are they left alone at the burrow to take care of themselves, undefended? Not according to Jon Rood, who conducted field studies on dwarf mongooses for many years. Rood found that out of eighty-five observations recorded, in all but a single case there was at least one baby-sitter at the nest. Baby-sitting, almost always undertaken by kin (older siblings), provided a critical service as very young individuals left alone in the den are particularly susceptible to predation. Baby-sitters have been seen giving alarm calls and even chasing potential predators from the den. Cooperative kin then forsake foraging themselves to help watch over and defend their siblings.     Without the quintessential refrigerator to raid, what benefits are baby-sitters getting for their services? Clearly, helping and cooperating are primarily driven by kinship in this example, as most (if not all) young in a den are related. Save your sibling from being eaten and you save potentially many copies of genes that also reside in you (keep in mind again that this is how kinship is defined in evolutionary biology). Family ties, however, do not explain the whole picture. For example, Rood found that one pack he observed contained an immigrant, unrelated two-year-old female he named Carrie. Carrie not only undertook baby-sitting services, she was the group's predominant dispenser of child care. Rood suggests that one possible reason that unrelated individuals may baby-sit is that this act increases survivorship of all group members. The recipients of help today, even if unrelated, may be the alarm callers of tomorrow, benefiting the baby-sitter, albeit down the road a bit.     Still, baby-sitters are temporary and not around all that much. Is anything more permanent in animal child care services available? Cooperative Breeding in Bee-Eaters In 1935 Alexander Skutch, naturalist and ornithologist extraordinaire, coined the term "helpers-at-the-nest" to describe a strange phenomenon he had observed in a number of species of birds. Skutch found that younger individuals, who were physiologically capable of reproducing, were not leaving their natal nests to find a territory and a mate. It was surprising indeed to find young individuals that could breed not cashing in on the opportunity. More fascinating, however, was the observation that such individuals were actually helping to raise the batch of babies (their new brothers and sisters) born during recent breeding cycles. Such helping often lasted significant periods of time and involved a suite of activities that included, but were not limited to, feeding the young and defending the nest against attack from predators.     Some sixty years after Skutch introduced the notion of helpers-at-the-nest, there is still controversy surrounding the question of why capable young birds remain at home and help raise their baby brothers and sisters. One possible solution was put forth by Jerram Brown, who argued that the evolution of helpers-at-the-nest is really a two-part story. He hypothesized that in many species, it is often quite difficult for young individuals to find suitable areas for starting their own new nests. Either there are no available slots in their environment or the usable area is sub-par, perhaps having too little food or too many predators. This situation creates pressure to remain at the home nest, past the age when individuals are physically able to breed on their own. Once the decision to remain home is in place, then such individuals help because it increases their inclusive fitness to do so (the critical comparison being with staying at home and not helping). So helpers do not stay at their natal nest because the inclusive fitness of doing so is higher than it would be on a good, new territory. Rather, they are forced by ecological factors to remain where they are, and as long as that is the case, the most productive activity for them to be involved in is raising their kin.     Lake Nakuru National Park, Kenya, is home to one of the better-studied species of birds with helpers, the white-fronted bee-eater. Stephen Emlen and Peter Wrege have been studying color-banded bee-eaters in various populations throughout this park for the last twenty-odd years, and they have come up with some remarkable findings. Bee-eaters are a particularly nice species in which to study the phenomenon of helpers-at-the-nest, because their ecology and population structure allow us to investigate two components of kin-selected cooperation that are often difficult to study: (1) whether individuals, when given the option, help those they are most related to, and (2) the effect of helpers on the survival of their younger siblings.     Bee-eaters exist in extended family groups, wherein a number of breeding pairs of adults produce offspring during a given season. Such "clans" typically number three to seventeen individuals, including up to five breeding pairs and many unpaired younger individuals. As such, potential helpers can choose not only whether to help but whom to help. Possible recipients range from siblings to more distant relatives to unrelated young. As predicted by kin selection theory, helpers consistently choose whom they will help based on relatedness. In the 108 cases measured by Emlen and Wrege, helpers provided assistance to the most related individuals in their nest 94 percent of the time. Helpers, then, are obviously helping the right individuals (according to kin selection theory at least), but do their actions really matter?     One initial critique of work on helpers came from researchers who did not think it mattered very much whether helpers were present or not. Whether helpers stick around the nest and appear to be helping their siblings is not the issue, the critical factor is whether one can demonstrate that they are helping--that is, increasing the number of young that survive. Bee-eaters may prove to be an extreme case on the continuum of how helpful helpers truly can be, because the effects of helpers in this species are dramatic. The average productivity of nests without helpers is doubled with the addition of each new helper.     Helpers have such a dramatic effect on the survival of their siblings, in fact, that parents have developed unique strategies to keep them around. By being particularly good at raising their brothers and sisters, helpers have made themselves a very valuable commodity, so valuable that parents would rather a helper remain at the nest than even attempt to breed elsewhere. As a result, older bee-eaters (almost always fathers) actively interfere with their sons' attempts at breeding somewhere else--surely not what one would initially expect if kin selection was a large player in this system. That is, one might think that kin selection theory would often favor having parents do whatever they can to increase the number of grandchildren they have. Grandchildren, after all, are kin as well (albeit not as close kin as offspring). But a father actually increases his inclusive fitness more if a helper stays home than if it leaves. That is, from the perspective of the parent, the number of copies of its genes making it to the next generation would be higher if helpers stayed at the nest and helped raise their sibs (the parents' offspring) than if they attempted to reproduce on their own (and produce grandchildren). Since dad is in control, in the sense of being larger and more powerful, he calls the shots, and kin selection favors disrupting junior's attempts at leaving home. The end result is that kin-selected cooperation will usually make it worthwhile for a helper to stay, but even if that is not the case, kin-selected aggression will see to it that he stays, regardless. (Continues...) Copyright © 1999 Lee Dugatkin. All rights reserved.

Table of Contents

Introduction: The Four Paths to Cooperation
1 All in the Family
2 One Good Turn Deserves Another
3 What's in It for Me?
4 For the Good of Others?
Conclusion: Possibilities and Pitfalls