Wednesday, September 30, 2015

2040. Book Review: The Amazing Inner Lives of Animals

By Tim Flannery, The New York Review of Books, October 8, 2015

Beyond Words: What Animals Think and Feel, by Carl Safina, 2015

The Cultural Lives of Whales and Dolphins, by Hal Whitehead and Luke Randall, 2015

The free-living dolphins of the Bahamas had come to know researcher Denise Herzing and her team very well. For decades, at the start of each four-month-long field season, the dolphins would give the returning humans a joyous reception: “a reunion of friends,” as Herzing described it. But one year the creatures behaved differently. They would not approach the research vessel, refusing even invitations to bow-ride. When the boat’s captain slipped into the water to size up the situation, the dolphins remained aloof. Meanwhile on board it was discovered that an expeditioner had died while napping in his bunk. As the vessel headed to port, Herzing said, “the dolphins came to the side of our boat, not riding the bow as usual but instead flanking us fifty feet away in an aquatic escort” that paralleled the boat in an organized manner.

The remarkable incident raises questions that lie at the heart of Carl Safina’s astonishing new book, Beyond Words: What Animals Think and Feel. Can dolphin sonar penetrate the steel hull of a boat—and pinpoint a stilled heart? Can dolphins empathize with human bereavement? Is dolphin society organized enough to permit the formation of a funeral cavalcade? If the answer to these questions is yes, then Beyond Words has profound implications for humans and our worldview.

Beyond Words is gloriously written. Consider this description of elephants:

Their great breaths, rushing in and out, resonant in the halls of their lungs. The skin as they moved, wrinkled with time and wear, batiked with the walk of ages, as if they lived within the creased maps of the lives they’d traveled.

Not since Barry Lopez or Peter Matthiessen were at the height of their powers has the world been treated to such sumptuous descriptions of nature.

Safina would be the first to agree that anecdotes such as Herzing’s lack the rigor of scientific experiments. He tells us that he is “most skeptical of those things I’d most like to believe, precisely because I’d like to believe them. Wanting to believe something can bias one’s view.” Beyond Words is a rigorously scientific work. Yet impeccably documented anecdotes such as Herzing’s have a place in it, because they are the only means we have of comprehending the reactions of intelligent creatures like dolphins to rare and unusual circumstances. The alternative—to capture dolphins or chimpanzees and subject them to an array of human-devised tests in artificial circumstances—often results in nonsense. Take, for example, the oft-cited research demonstrating that wolves cannot follow a human pointing at something, while dogs can. It turns out that the wolves tested were caged: when outside a cage, wolves readily follow human pointing, without any training.

Safina explains how an evolutionary understanding of the emotions helps us to see even humble creatures as individuals. The chemical oxytocin creates feelings of pleasure and a craving for sociality. So widespread is it that it must have originated 700 million or more years ago. Serotonin, a chemical associated with anxiety, is probably equally ancient: crayfish subjected to mild electrical shocks have elevated serotonin levels, and act anxiously. If treated with chlordiazepoxide (a common treatment for humans suffering from anxiety) they resume normal behavior.

The basic repertory of emotions evolved so long ago that even worms exhibit great behavioral sophistication. After a lifetime studying earthworms, Charles Darwin declared that they “deserve to be called intelligent,” for when evaluating materials for plugging their burrows, they “act in nearly in the same manner as a man under similar circumstances.” Emotions are the foundation blocks of relationships and personalities. Driven by the same complex mix of emotion-inducing chemicals as ourselves, every worm, crayfish, and other invertebrate has its own unique response to its fellows and the world at large.

Worms and crayfish may have distinct personalities and emotional responses, but their brains are far simpler than ours. Humans fall within a small group of mammals with exceptionally large brains. All are highly social, and it is upon this group—and specifically the elephants, killer whales, bottlenosed dolphins, and wolves—that Safina concentrates. The last common ancestor of these creatures was a primitive, small-brained, nocturnal, shrew-sized mammal that lived around 100 million years ago. The brains, bodies, and societies of these “animal intelligentsia,” as we might call them, are each very different, making it hard to understand their lives.

Safina sees and describes the behaviors of the animals he’s interested in through the eyes of researchers who have dedicated their lives to the study of their subjects. What is it like to be an elephant? Cynthia Moss, who has lived with the elephants of Amboseli National Park in Kenya for four decades, sums them up as “intelligent, social, emotional, personable, imitative, respectful of ancestors, playful, self-aware, compassionate.” It all sounds impressively human, but elephant societies are very different from our own. Female elephants and their young live separately from males, for example, so they have no conception of romantic love or marriage (though the females can be very interested in sex, enough to fake estrus in order to attract male attention).

Much published behavioral science, incidentally, is phrased in a neutral language that distances us from animals. Safina argues that we should use a common language of grief, joy, friendship, and empathy to describe the equivalent responses of both human and other animals. To this I would add the language of ceremony: What other word but “marriage” should be used to describe the ritual bonding, followed by lifelong commitment to their partners, of creatures like the albatross?

Sometimes it is the small things that best reveal shared life experience. When baby elephants are weaned they throw tantrums that rival those of the wildest two-year-old humans. One youngster became so upset with his mother that he screamed and trumpeted as he poked her with his tiny tusks. Finally, in frustration, he stuck his trunk into her anus, then turned around and kicked her. “You little horror!” thought Cynthia Moss as she watched the tantrum unfold.

Clans of female elephants, led by matriarchs, periodically associate in larger groups. As a result, elephants have excellent memories, and are able to recognize up to one thousand individuals. So strong is elephant empathy that they sometimes bury their dead, and will return repeatedly to the skeleton of a deceased matriarch to fondle her tusks and bones. Indeed, an elephant’s response to death has been called “probably the strangest thing about them.” When the Amboseli matriarch Eleanor was dying, the matriarch Grace approached her, her facial glands streaming with emotion, and tried to lift her to her feet. Grace stayed with the stricken Eleanor through the night of her death, and on the third day Eleanor’s family and closest friend Maya visited the corpse. A week after the death the family returned again to express what can only be called their grief. A researcher once played the recording of a deceased elephant’s voice to its family. The creatures went wild searching for their lost relative, and the dead elephant’s daughter called for days after.

Elephants have been known to extract spears from wounded friends, and to stay with infants born with disabilities. In 1990, the Amboseli female Echo gave birth to a baby who could not straighten his forelegs, and so could hardly nurse. For three days Echo and her eight-year-old daughter Enid stayed with him as he hobbled along on his wrists. On the third day he finally managed to straighten his forelegs and, despite several falls, he was soon walking well. As Safina says, “His family’s persistence—which in humans facing a similar situation we might call faith—had saved him.”

Most of us will never see a wild elephant, much less spend the time observing them that is required to understand them as individuals. But there are animals that share our lives, and whose societies, emotional depth, and intelligence are readily accessible. Dogs are often family to us. And it is astonishing how much of a dog’s behavior is pure wolf.

The Canidae—the family to which wolves and dogs belong—is a uniquely American production, originating and evolving over tens of millions of years in North America before spreading to other continents around five million years ago. The American origins of the wolf family did not save them from frontier violence. By the 1920s they had been all but exterminated from the contiguous forty-eight states of the US. Their reintroduction into Yellowstone National Park in January 1995 offered a unique opportunity to follow the fortunes of wolf families as they made their way in a new world. Yellowstone’s wolf research leader Doug Smith says that wolves do three things: “They travel, they kill, and they are social—very social.” But wolves are also astonishingly like us. They can be ruthless in their pursuit of power, to the extent that some will kill their sister’s cubs if it serves their ends. But they will also at times adopt the litters of rivals.

The best wolves are brilliant leaders that pursue lifelong strategies in order to lead their families to success. According to wolf watchers, the greatest wolf Yellowstone has ever known was Twenty-one (wolf researchers use numbers rather than names for individuals). He was big and brave, once taking on six attacking wolves and routing them all. He never lost a fight, but he was also magnanimous, for he never killed a vanquished enemy. And that made him as unusual among wolves as did his size and strength. He was born into the first litter of Yellowstone pups following the reintroduction of wolves in the park. Twenty-one’s big break came at age two and a half when he left his family and joined a pack whose alpha male had been shot just two days earlier. He adopted the dead wolf’s pups and helped to feed them.

A telling characteristic of Twenty-one was the way he loved to wrestle with the little ones and pretend to lose. The wolf expert Rick McIntyre said, “He’d just fall on his back with his paws in the air. And the triumphant-looking little one would be standing over him with his tail wagging.” “The ability to pretend,” McIntyre said, “shows that you understand how your actions are perceived by others. It indicates high intelligence.” That many humans recognize this in dogs, but have failed to see it in wolves, speaks strongly of the need for Safina’s book. For dogs are wolves that came to live with us.

The similarities between wolves and humans are arguably more extensive than those between humans and any other animal. Tough, flexible in social structure, capable of forming pair bonds and fitting into ever-shifting hierarchies, we were made for each other. And when we out-of-Africa apes met up with the arch-typical American canids a few tens of thousands of years ago, a bond was created that has endured ever since. Just who initiated the interspecies relationship is hotly debated. The traditional view is that humans domesticated dogs, but Safina makes a convincing case that the process was driven as much by the wolves as by the humans. The wolves that were better able to read human tendencies and reactions, and were less skittish of human contact, would have gotten access to more food scraps from human camps. And human clans willing to tolerate the wolves would have obtained valuable warnings of the presence of danger from other animals (and other humans). Eventually, Safina says, “we became like each other.” The partnership, however, has had some puzzling effects. The brains of dogs, as well as humans, have shrunk since we began living together, perhaps because we came to rely on each other rather than solely on our own wits.

Sperm whales have the largest brains on earth—around six times larger on average than our own—while bottlenosed dolphins have the largest brains relative to body size, with the exception of humans. Along with killer whales, these species have a place beside the elephants, dogs, and great apes in the animal intelligentsia. The Cultural Lives of Whales and Dolphins is a comprehensive academic work by researchers who have devoted their careers to studying sperm and killer whales. Ocean-going and deep diving, sperm whales are difficult to study, and researchers can as yet offer only a bare sketch of their societies. But it’s already clear that their social organization has remarkable parallels with that of elephants. Like elephants, sperm whale females and young often live in “clans” of up to thirty individuals, while adult males, except when mating, live separate lives.

Sperm whale clans possess distinctive “dialects” of sonar clicks. These are passed on by learning, and act as markers of clan identity. They are an important part of the whale’s communication system, which enables the creatures to synchronize their diving, feeding, and other activities. So social are sperm whales that females share the care of the young of their clan, for example by staying at the surface with a young whale while its mother dives for food. Clan members are so closely bonded that they spend extended periods at the surface, nuzzling one another or staying in close body contact. As with elephants, clans can gather in large congregations, so it seems reasonable to assume that sperm whales have the capacity to memorize large social networks.

Killer whales (otherwise known as orcas) have a very different social organization. Without doubt their most unusual characteristic is that all male killer whales are deeply involved with their mother. They never leave their mother’s clan, and despite their enormous size (growing to twice the weight of females), their fates remain deeply intertwined with those of their mothers. If their mothers should die, even fully adult males over thirty years old (they can live to over sixty) face an eight-fold increase in their risk of death. Just how and why the orphaned adult males die remains unclear.

Another striking feature of killer whales and near relatives is the extraordinary length of lactation. Short-finned pilot whales lactate for at least fifteen years after birth, even though puberty occurs at between eight and seventeen years. Sperm whales reach sexual maturity at nine to ten years of age, but traces of milk have been found in the stomachs of thirteen-year-olds. Killer whales and humans are unique in that they experience menopause (for the whales typically at around age forty). Because female killer whales can live up to eighty years, around a quarter of females in any group are postreproductive. Yet they remain sexually active. Grandmothers are evidently very important in killer whale societies, almost certainly because of the wisdom they have gathered over a lifetime.

An equally odd aspect of killer whale culture concerns food taboos and ways that whales observe them. In this they offer an extraordinary parallel with some human cultures. One clan of killer whales eats only a single species of salmon. Another kills only one species of seal. When members of a mammal-eating clan were captured for the aquarium trade in the 1970s, they starved themselves for seventy-eight days before eating the salmon being proffered, and then they ate the fish only after they had performed a strange ceremony. The two whales held gently onto either end of a dead salmon, and swam a single lap around their pool with it in their mouths, before dividing the fish between themselves and consuming it.

Killer whales are strongly xenophobic. Clans of salmon eaters never mix with mammal eaters, for example. Genetic studies show that clans with different food taboos don’t interbreed, leading to slightly different appearances and genetic makeup. Each clan has a distinctive dialect of vocalizations (perhaps we should call them languages), which facilitates coordination of their work, division of their labor, and care of one another.

At times, killer whales have developed special relationships with people. During the nineteenth and early twentieth centuries, at Twofold Bay south of Sydney, Australia, killer whales and humans set up a mutually profitable whaling enterprise. The killer whales would notify the whalers of the presence of humpback whales by performing a ritual in the waters of the bay fronting the whaler’s cottagers. The men would harpoon the humpbacks, and the killer whales would hold on to the harpoon ropes to tire the prey.

After a humpback was lanced and killed by the men, they observed the “law of the tongue.” The whalers would leave the humpback body for twenty-four hours so that the killers could feast on the lips and tongue. Remarkable proof of this partnership persists, in the form of the skeleton of “Old Tom”—a killer whale whose teeth were worn flat on one side while holding onto harpoon ropes—which can be seen in the killer whale museum in the town of Eden, Australia.

With the exception of our species, killer whales are earth’s most capable predators. When they evolved ten million years ago, half of earth’s whales, seals, and dugong species became extinct. Because they specialize in a particular food type and are so intelligent, killer whales continue to have a huge impact on their prey. As a result of global warming, killer whales have appeared in Arctic waters. Horrified Inuit describe them as voracious and wasteful killers that have reduced populations of some Arctic mammals by a third.

Safina comes to an unfamiliar but empirically based conclusion: prior to the domestication of plants and the invention of writing, the differences between human societies and those of elephants, dogs, killer whales, and dolphins was a matter of degree, not kind. Why, he asks, has it taken us so long to understand this? Are our egos “threatened by the thought that other animals think and feel? Is it because acknowledging the mind of another makes it harder to abuse them?”

The discovery of nonhuman societies composed of highly intelligent, social, empathetic individuals possessing sophisticated communication systems will force us to reformulate many questions. We have long asked whether we are alone in the universe. But clearly we are not alone on earth. The evolution of intelligence, of empathy and complex societies, is surely more likely than we have hitherto considered. And what is it, exactly, that sets our species apart? We clearly are different, but in light of Beyond Words we need to reevaluate how, and why.

Beyond Words will have a deep impact on many readers, for it elevates our relationships with animals to a higher plane. When your dog looks at you adoringly, even though he or she cannot say it, you can be as sure that love is being expressed as you can when hearing any human declaration of eternal devotion. Most of us already knew that, but have withheld ourselves from a full surrender to its implications. Along with Darwin’s Origin and Richard Dawkins’s Selfish Gene, Beyond Words marks a major milestone in our evolving understanding of our place in nature. Indeed it has the potential to change our relationship with the natural world.

Tuesday, September 29, 2015

2039. After 60 Million Years of Extreme Living, Seabirds Are Crashing

By Jeremy Hance, The Guardian, September 22, 2015
Arctic tern (Sterna paradisaea)
Every day for sixty million years, seabirds have performed mind-boggling acts of derring-do: circumnavigating the globe without rest, diving more than 200 meters in treacherous seas for a bite of lunch, braving the most unpredictable weather on the planet as if it were just another Tuesday and finding their way home in waters with few, if any, landmarks. 

But now seabirds, like so many other species, may have met their match.

Conservationists have long known that many seabird populations are in decline, but a recent paper in PLOS ONE finds the situation worse than anticipated. According to the researchers, seabird abundance has dropped 69.7% in just 60 years – representing the deaths of some 230 million animals.

“I was very surprised with the result, it was considerably greater than I’d expected,” said Edd Hammill, co-author of the paper, with Utah State University. “What we should take away from this is that something is serious amiss in the oceans.”Ben Lascelles, a Senior Marine Officer with Birdlife International, who was not involved in the study, said he found the research alarming because the decline appeared practically indiscriminate, hitting a “large number of species across a number of families.” 

Seabirds, which include any bird that depends largely on the marine environment, comprise nearly 350 species worldwide – an astonishing variety of extreme-loving birds. For example, the indefatigable wandering albatross, which sports the largest wingspan on the planet; the child-sized Emperor penguin, the only bird that breeds during the Antarctic winter; and the tiny storm petrel that practically capers on the water as it feeds – they are named for St. Peter after all.

But, given that seabirds inhabit both the open ocean and the shoreline, this eclectic mix of birds faces a litany of threats: overfishing, drowning in fishing lines or nets, plastic pollution, invasive species like rats in nesting areas, oil and gas development and toxic pollution moving up the food chain. And as if these weren’t enough, the double-whammy of climate change and ocean acidification threatens to flood nesting sites and disrupt food sources.

“Seabirds are particularly good indicators of the health of marine ecosystems,” explained lead author, Michelle Paleczny with the University of British Columbia and the Sea Around Us Project. “When we see this magnitude of seabird decline, we can see there is something wrong with marine ecosystems. It gives us an idea of the overall impact we’re having.”

But with such a large number of species across such a wide variety of environment one is left asking: how did the scientists count so many birds?

First, the team of researchers scoured all the population data on seabirds available. They found demographic data on 3,213 populations. But they couldn’t use all of theses counts, since conservationists had surveyed many of these far-flung populations just once or twice – not enough to show a real trend.

The team eventually selected 513 populations that had been counted at least five times. In all, these populations represented about 19 percent of the world’s seabirds.

Still, Hammil said he believes the team’s findings “are an accurate representation of what is happening worldwide.”
He added, “although we did not include every population, all seabird families were included, and we included populations from every major coastline in the world.” 

Paleczny also said that when the researchers looked at the differences between monitored and unmonitored populations, they saw “no evidence that the monitored populations are declining more.”

“The trends for many seabird species have clearly been downwards for a number of years, and this paper provides further evidence of this,” Lascelles said.

Still, Paleczny and Hammil’s research arguably paints an even more alarming picture of the state of the world’s seabirds. For example, according to them, the tern family has fallen by 85%, frigatebirds by 81%, petrels and shearwaters by 79%, and albatrosses by 69%. 

Such dismal findings point to one of the study’s patterns: open ocean birds – such as albatrosses, frigatebirds, petrels and shearwaters – are generally faring worse than birds that stick near the coasts.

“[Open-ocean] seabirds are hit especially hard due to their large geographic ranges. Because these species travel so far, there is a greater chance they will encounter threats,” said Hammill who noted that coastal birds “in some cases” are doing better because of improved management of breeding areas and improved fishing gear.

But even when threats were minimised, Lascelles noted that recovery requires diligence and patience.

2038. Darwinian Natural Section: Individual vs. Group Selection

By Steve Mirsky, Scientific American, December 18, 2008

Want to start a brawl at an evolution conference? Just bring up the concept of group selection: the idea that one mixed bag of individuals can be “selected” as a group over other heterogeneous groups from the same species. Biologists who would not hesitate to form a group themselves to combat creationism or intelligent design might suddenly start a pie fight to defend the principle that “it’s every man for himself.”

Yet Charles Darwin himself argued for group selection. He postulated that moral men might not do any better than immoral men but that tribes of moral men would certainly “have an immense advantage” over fractious bands of pirates. By the 1960s, however, selection at the group level was on the outs. Influential theorist George Williams acknowledged that although group selection might be possible, in real life “group-related adaptations do not, in fact, exist.”

Richard Dawkins of the University of Cambridge, whose writings have reached millions, maintains that selection might not even reach such a high level of biological organization as the individual organism. Instead, he claims, selection operates on genes—the individual is the embodiment of the selection of thousands of selfish genes, each trying to perpetuate itself.

In the past few decades, however, group selection has made a quiet comeback among evolutionary theorists. E. O. Wilson of Harvard University and David Sloan Wilson (no relation) of Binghamton University are trying to give group selection full-fledged respectability. They are rebranding it as multilevel selection theory: selection constantly takes place on multiple levels simultaneously. And how do you figure the sum of those selections in any real-world circumstance? “We simply have to examine situations on a case-by-case basis,” Sloan Wilson says.

But the Wilsons did offer some guidelines in the December 2007 issue of Quarterly Review of Biology. “Adaptation at any level,” they write, “requires a process of natural selection at the same level, and tends to be undermined by natural selection at lower levels.”

Experiments with actual groups illustrate the point. Pseudomonas fluorescens bacteria quickly suck all the dissolved oxygen out of a liquid habitat, leaving a thin habitable layer near the surface. But some bacteria spontaneously develop a beneficial mutation. These group-saving individuals secrete a polymer that enables bunches of individuals to form floating mats. As a mat, all the bacteria survive, even though most of them expend no metabolic energy producing the polymer. But if the freeloaders get greedy and reproduce too many of their kind, the mat sinks and everybody dies, altruists and freeloaders alike. Among these bacteria, then, groups that maintain enough altruists to float outcompete groups with fewer altruists than that minimum number. The former groups survive, grow and split up into daughter groups. Thus, altruistic individuals can prosper, despite the disadvantage of expending precious resources to produce the polymer.

Perhaps the biggest change that group selection brings to evolutionary theory is its implication for so-called kin selection. What looks like group selection, some theorists argue, can actually be understood as genetic relatedness. Evolutionist J.B.S. Haldane pithily explained kin selection: “I would lay down my life for two brothers or eight cousins.” In this view, altruistic bacteria in the Pseudomonas mats are saving close relatives, thereby ensuring the survival of most of the genes they themselves also carry.

Turning that argument on its head, the Wilsons assert that kin selection is a special case of group selection. “The importance of kinship,” they note, “is that it increases genetic variation among groups.” The individuals within any one group are much more like one another and much less like the individuals in any other group. And that diversity between groups presents clearer choices for group selection. Kinship thus accentuates the importance of selection at the group level as compared with individual selection within the group.

The Wilsons think evolutionists must embrace multilevel selection to do fruitful research in sociobiology—“the study of social behavior from a biological perspective.” When doing so, other investigators can keep in mind the Wilsons’ handy rule of thumb: “Selfishness beats altruism within groups. Altruistic groups beat selfish groups.”

Saturday, September 26, 2015

2036. The Spandrels Of San Marco Revisited: An Interview With Richard C. Lewontin

By David Sloan Wilson, The Evolution Institute, March 29, 2015
Richard Lewontin

One of the most widely cited and discussed articles in evolutionary biology is “The Spandrels of San Marco and the Panglossian Paradigm: A Critique of the Adaptationist Programme”, which was written by Harvard biologists Stephen Jay Gould and Richard C. Lewontin and published in the Proceedings of the Royal Society of London in 1979. Their critique of their own field of evolutionary biology spilled out of the Ivory Tower onto the pages of general intellectual forums such as the New York Review of Books.

Gould died in 2002 but his coauthor is still active. Richard C. Lewontin is a population geneticist by training and pioneered the method of gel electrophoresis among many other accomplishments. His academic books include The Genetic Basis of Evolutionary Change (1974), which I eagerly read as a graduate student. His biological books for the general public include The Triple Helix: Gene, Organism, and Environment (2002) and Human Diversity (part of the Scientific American Library Series; 1982). In his role as social critic and theorist, his books include Biology Under the Influence: Dialectical Essays on Ecology, Agriculture, and Health (with Richard Levins; 2007), It Ain’t Necessarily So: The Dream of the Human Genome and Other Illusions (2001), Biology as Ideology: The Doctrine of DNA (1993), and Not in Our Genes: Biology Ideology, and Human Nature (with Leon J. Kamin and Steven Rose; 1984). Finally, Lewontin has served as a mentor for many PhD and postdoctoral students, in philosophy in addition to biology, including my longstanding philosophical collaborator Elliott Sober.

I talked by phone with Lewontin on March 2 2015. In his mid-eighties, he is still scientifically active and could recall his collaboration with Gould in detail. Our conversation is highly relevant to the “Just so story” critique that is frequently leveled against Evolutionary Psychology.

DSW: I’m so happy to be talking with you and thanks for making the time.
RL: I don’t know if anything I can say to you at this stage has any usefulness.
DSW: I think that is overly humble on your part.

RL: You can’t be overly humble.

DSW: That’s true. Humility is a religious virtue but it is also a secular virtue and a scientific virtue. I couldn’t agree more. I am interested among other things in social history. To get started, you are one of the preeminent evolutionary biologists of our time—many achievements. In 1979 you felt the need to write this article with Stephen Jay Gould that became a classic.

RL: Sorry—which article is that?

DSW (laughs): The Spandrels of San Marco.

RL: I thought it was but I didn’t remember the exact date.

DSW: I wonder if you could tell me—what were the circumstances that moved you and Steve Gould to write this article?

RL: Sure, I can give it to you in detail. I was invited by, I think it was the Royal Society, to come and give a lecture. For one reason or another that I can’t remember, I couldn’t go. So I asked if it would be alright if I asked Steve Gould if he would go in my place–Steve and I were teaching evolution together—and they said sure. So Steve went and he gave a talk from the standpoint of what interested him at the time, which was the notion that some traits arise simply as a structural byproduct of selection on other traits, and he chose to call them spandrels. I did make a contribution to the written version of that article, but most of it was Steve’s.

DSW: Ok!

RL: Now I should warn you about my prejudices. Steve and I taught evolution together for years and in a sense we struggled in class constantly because Steve, in my view, was preoccupied with the desire to be considered a very original and great evolutionary theorist. So he would exaggerate and even caricature certain features, which are true but not the way you want to present them. For example, punctuated equilibrium, one of his favorites. He would go to the blackboard and show a trait rising gradually and then becoming completely flat for a while with no change at all, and then rising quickly and then completely flat, etc. which is a kind of caricature of the fact that there is variability in the evolution of traits, sometimes faster and sometimes slower, but which he made into punctuated equilibrium literally. Then I would have to get up in class and say “Don’t take this caricature too seriously. It really looks like this…” and I would make some more gradual variable rates. Steve and I had that kind of struggle constantly. He would fasten on a particular interesting aspect of the evolutionary process and then make it into a kind of rigid, almost vacuous rule, because—now I have to say that this is my view—I have no demonstration of it—that Steve was really preoccupied by becoming a famous evolutionist.

DSW: So he was trying to grab center stage at every opportunity.

RL: Yeah, I think so.

DSW: Is the Spandrels paper like that?

RL: Well, I made a lesser contribution than he did. Most of the Spandrels paper was written by Steve. There is a section in there, which one can easily pick out, where I discuss the various factors and forces of evolution…

DSW: Yes, I can see that division.

RL: That paper never would have been written by us as a joint paper if I hadn’t asked Steve to go to the Royal Society and give a talk in my place.

DSW: Fascinating! To what extent was this paper motivated, either for you or Steve, by Sociobiology?

RL: Well, I don’t know to what extent it was motivated. Sociobiology was certainly contextually relevant. I think we would have written exactly the same paper if Sociobiology had never come into existence. Looking back, that’s hard to say, but I think the idea was to avoid selective caricatures, the making up of selective stories just because you felt you had to. That is, as you are aware, a very common phenomenon in writing about and teaching evolution. For example, why is blood red? The fact that hemoglobin happens to have that absorption spectrum in the visible is not sufficient for some people, who have to show that it’s a good thing that blood is red because it scares off predators who come and scratch you and stuff like that. So we had a lot of that sort of thing to deal with—what we called “Just so stories”.

DSW: So forget about Sociobiology—basically, you saw na├»ve adaptationism being a problem in general in the field of evolutionary biology?

RL: Exactly. It was really not Sociobiology itself, but a tendency to try to find, in every instance, some selective advantage for things. We were teaching the students—and Steve was not as keen on that as I was—that there are a whole variety of forces that give rise to observed traits and they are not all directly selected for.

DSW: This interests me very much. I’m interested to know that was the primary motivation for the article, not Sociobiology.

RL: Yeah.

DSW: What’s the right way to do it? When I talk about the adaptationist program, I say that an adaptationist hypothesis is often the best way to start because it doesn’t require much information to know what an organism should be like to be well adapted to its environment. So it’s a good starting point, although certainly not the end point. Then the inquiry can go in a direction where you decide—as with the color of blood, by the way, an example that I use myself—that this is probably not an adaptation. You can arrive at the truth of the matter. But thinking in adaptationist terms is part of the process. So what’s the way to do it right and what’s the role of adaptationist thinking in an appropriate procedure?

RL: Well (laughs), you’re asking me what the right way to do it is. I think the right way is to start with the sentence: “We do not have any hard evidence of the forces leading to the following evolutionary change.” There has to be a prelude to the discussion of evolutionary change to make it clear that although the theory of natural selection is very important and happens lots, there are other forces, or other mechanisms, that lead to change and we are not obliged by being Darwinians and being evolutionists to invent adaptive explanations for all changes. I think that’s where you have to start. Then, as either a philosopher or biologist, ask in a particular case what is the direct evidence, besides the desire that we want to find something, that a particular story is true or not true. Most of the time we’re going to have to say that this happened in the Eocene or the Paleocene and we haven’t the foggiest notion of why it happened. I think the admission of necessary ignorance of historically remote things is the first rule of intellectual honesty in evolution.

DSW: Good. Thank you for saying that so clearly. At the same time, sometimes the past can be inferred with amazing certainty. All the historical sciences are like that, right?

RL: Right. And so, I think that the right general strategy for explanation, writing, and teaching is to begin with some really clear cut cases where we have in our very hands the evidence for a particular causal pathway–a greater reproduction and survivorship of one form versus another–and then move from that to living cases where we’re not quite so sure because we can’t actually count the number of offspring of each type, and so on, to somewhat hazier cases, and then go back to extinct organisms and evolutionary past and say, we could make up a good story, but we don’t know how to show that it’s really true.

DSW: Right. There is the humility again that began our conversation. Although I have to say that it was the lack of humility, in part, that caused the Spandrels article to become so widely read. Steve Gould the showman put that on center stage and it became a three ring circus for a long time, as you know. The adaptation wars and all that. I don’t know whether that is good or bad.

RL: All we can say is that sometimes the stories are right and sometimes they’re wrong. One way to deal with this is to take a case where we can in fact—because it’s present and we have the organisms, and we really test what’s going on. Take such a case and put it aside and don’t even mention the result of our determined discovery about it, and make up three stories, all of which sound perfectly biologically realistic and reasonable, and then give the right answer and say now, aside from the necessary observations and experiments—forget the real answer, why should we choose one of these versus another?
DSW: Uh Huh. Interesting.

RL: What we have to decide is whether we’re going to put behind us certain motivations, one of which is the general motivation to struggle against religious anti-evolutionary views, and at the other extreme to be as individuals successful as evolutionary biologists by giving an explanation of something interesting even when we don’t have the observations. I was raised not as an evolutionist but as a population geneticist.

DSW: Right.

RL: That’s a big difference.

DSW: Why is that a big difference? Let’s clarify that for me. I tend to see it as a small difference. What’s the difference between being a population geneticist and an evolutionist?

RL: A population geneticist by theoretical training has certain parameters of population change. That’s become broadened by the realization that there are between population changes and so on, but within a population we’re talking about changes in gene frequency and we have a catalog of the causes: selection, inbreeding, chance, mutation, and so on. Our job as population geneticists is to do the necessary observations of the various things that give us estimates of the strength of those different forces. Now, historically one of the most interesting—now I want to talk a little about the sociology of our science—Theodosius Dobzhansky, my professor and then greatest living evolutionary biologist…

DSW: Mr. “Nothing in biology makes sense except in the light of evolution…”
RL: Yeah, right. He was a very bad field observer. Theodosius Dobzhansky never, in his entire life, nor any of his students, me included—I would go out in the field with him, actually–ever saw a Drosophila pseudoobscura in its natural habitat.
DSW (laughs): Yeah, OK!

RL: We didn’t know where they laid their eggs. We couldn’t have counted the number of eggs of different genotypes. How did we study Drosophila in the wild? We went out into the desert, into Death Valley, we moved into a little oasis, we went first to the grocery store, and bought rotten bananas. We mushed up the bananas with yeast till they fermented a bit, we dumped that into the paper containers, put it out in the field and the flies came to us.

DSW: Right! No naturalistic context whatsoever.

RL: None…at…all. And to this day we do not know anything about the actual habitat of Drosophila pseudoobscura, although by the way, interestingly enough, in more recent years, Tim Prout actually succeeded in trapping pseudoobscura in orange groves, so we don’t even know how much they hang out with cultivated fruit.

DSW: Right.

RL: Now let me go one step further because we cannot understand the development of evolutionary biology if we don’t understand questions of the sociology of academic life. If I wanted to study evolutionary forces acting on some genetic polymorphism in Drosophila, I would go and look for some species of Drosophila where I could actually look at, perturb, and work with the actual breeding sites and egg laying sites and pick up larvae in nature and so on. And in fact there is such a group of Drosophila. They the cactophilic ones. There is a group [of scientists] from Texas and other places that studies the cactophilic Drosophila in an ecologically sensible way of going to the rot pockets and perturbing them, getting larvae out of them and so on. That group never acquired the prestige associated with the Dobzhansky school because—I don’t know why. They were doing what one has to do. That’s why, for example, I try to convince students who are entering evolutionary biology not to study animals at all but to study plants. Plants stay in one place. You can manipulate them. You can move them. Plants are much better than animals for studying things in nature. Yet, plant evolutionary biology is not, for sociological reasons that I don’t understand—I could make up stories—has never had the prestige that animal work has had when it comes to population genetics.

DSW: Right. I think that [there was an] all consuming interest in physical mechanisms as opposed to a more fully rounded approach. I place a lot of emphasis on the classic paper by Niko Tinbergen, “The Methods and Aims of Ethology”, in which he says that you have to ask four questions: Function, History, Mechanism, Development. Are you familiar with that paper?

RL: No, I’m not. Send me a reference to it.

DSW: It’s such a succinct summary of what a fully rounded approach needs to be. Dick, I’d like to spend a little bit of time on Sociobiology and also Evolutionary Psychology, because even though that didn’t motivate the Spandrels paper, it still motivated you to be a critic and Steve too. I wonder if you could bring us back to that point and what you saw as problematic about Sociobiology and then Evolutionary Psychology.

RL: This is what I have been talking about for the last five minutes. This is a branch of academic life that consists entirely, as far as I can see, of making up what would seem to be plausible stories. I would say that’s not what we are in business to do. I don’t know what else to say. Look, when I look at Sociobiology, the book or some of the other books he [E.O. Wilson] has written, it drives me mad. For example, if you read—I’ll take an extremely nasty example because it’s so clear—it is written that aggression is a part of human nature. It says that in the book. It lists features of human nature and aggression is one of them. So then I have said to Ed and others of his school, what do you do about people who have spent almost their entire lives in jail because they refuse to be conscripted into the army? What do you think the answer is? That is their form of aggression.

DSW: Well, OK, that’s facile.

RL: I don’t know what you can do about it. If everything can be said to be a form of aggression, even the refusal to be physically aggressive, what kind of science is that?

DSW: Would it be more acceptable to say that aggression is part of the repertoire of human behavior? That leaves it open to be part of the repertoire that we don’t always use.

RL: Before I will allow you to make even that statement, I will insist that you write down how I know whether any particular phenotypic manifestation is or is not included in your definition of aggression.

DSW: That’s fair.

RL: I don’t know what those people would say. But if you are willing to make a clear enough—and most important, and this is perhaps the fundamental contradiction, potentially there has to exist a group of cases of non-aggression. Because if everything by definition can be shown to be aggression then it ceases to be a useful concept in our scientific discussions.

DSW: The problem of explaining everything, and therefore nothing, recurs again and again. What were some of the political implications of Sociobiology that worried you? Misuses of biology, or misuses of evolutionary reasoning–back then and are they still with us today?

RL: My main complaint is not the list of specific manifestations but the underlying claim that there exists a human nature, which then the claimant must give examples of, and so each claimant gives examples that are convenient for his or her pet theory. I think the worst thing we can do in science is to create concepts where what is included or not included within the concept is not delimited to begin with. It allows us to claim anything. That’s my problem with Sociobiology. It’s too loose.

DSW: That brings us to the topic of cultural evolution, which is something that I study a lot and I think that you have thought of a lot. One of the exciting things that I think has taken place is the idea that the study of evolution became too gene-centric over the course of the 20th century. Evolution requires heredity, not genes, and there are other mechanisms of heredity. Culture really is an evolutionary process. That is in part why we are so open-ended, why there’s not a human nature in terms of a fixed human nature because we’re so adaptable. I wonder how much you have thought about cultural evolution in that way and if you have any comment to make upon what studying culture as a genuine evolutionary process with its own inheritance mechanism, including symbolic thought—the evolution of meaning systems—what that does to change the picture of evolution.

RL: I think that the evolution of this thing that’s in our cranium, however it happened, has changed all the rules for the history of the species, for its biology, for everything about it. I mean, rational thought and the kind of communication we have with human language, as opposed to the stereotypical communication of other animals, has really made a fantastic change in the conditions of life and the rates of reproduction of individual types and so on. I would say human evolution is in that sense unique because of the possibility of: a) the details of communication; b) the notion of historical memory; well, everything about human thought. I really do think that if we want to understand evolution, the first species we should keep out of our consideration is Homo sapiens. I’m sorry, but that’s the way it is for me.

DSW: Well, but that’s genetic evolution. What if the principles of cultural evolution, although different in some respects–because another inheritance mechanism needs to be functionally an inheritance mechanism but need not resemble other inheritance mechanisms in detail–what if the basic way of going about genetic evolutionary reasoning also could be employed for cultural evolutionary reasoning? Wouldn’t that put evolution back in the game for the study of human cultural evolution?

RL: Well, let me ask you a question. Why do you use cultural evolution instead of cultural history? Why evolution instead of history? Can you avoid—let me put it another way—can you generally avoid the false similarities, the made up structures that we are criticizing, if we continue to use the word evolution when what we really mean is historical change?

DSW: I’ll actually take you on, on that issue. I’m not sure how much time we’ll have during this conversation. What the mechanisms of human cultural evolution do is adapt human populations to their environments, often with a fine degree of sophistication. When you just say cultural history you’re being agnostic about adaptation. Cultures adapt to their environments. [Without a history of adaptation] they could no more survive and reproduce in their environments than a genetically evolved species. Just to say history leaves out adaptation, don’t you think?

RL: But the problem, here, is that it’s a form of adaptation that hasn’t been studied enough in animals and plants, which is that each change in the species changes what we call the environment, so there is a co-evolution of organism and environment. Historical change in our species has been increasingly the consequence of the organism itself. We’re inventing it all. As the brain grew into what we now have, it became the chief mechanism by which organisms constructed their environment. Look, let me interject here. I think it is extremely important to go to a fundamental issue, which is organisms create their own environments. All organisms make their niches. The whole notion of ecological niche is a very bad notion. There are no niches without organisms. This notion that there is a hole in the world that the organism evolves to fill. The organism by its evolution changes the conditions of its life and changes what surrounds it. Organisms are always creating their own hole in the world, their own niche.

DSW: You pioneered the concept of niche construction, which has become a hot topic.

RL: I think that one mustn’t see niche construction as a special issue. There are niches and then there is reconstruction of the niches. My claim is a very strong one and I could be wrong: there is no niche without an organism.

DSW: I’ll accept that provisionally, but there is such a thing as a purely physical environment…

RL: A physical world, oh yeah, but let me tell you one of my favorite seminars that I ever heard. I can’t trace it any more, unfortunately. A guy came to Chicago and gave a talk in which he showed motion picture photographs of all kinds of organisms, plants and animals, using what are called Schlieren optics, which are sensitive to differences in optical density. What he showed was every organism of which he took moving pictures—both plants and animals—have around them a layer of warm moist air–even trees have it—which is being produced by the organisms themselves. So every organism, at least every terrestrial organism (I don’t know about aquatic ones) is by its metabolism producing a layer of warm moist air with certain gases in it that are its immediate environment.

DSW: I think that’s probably even more so for aquatic organisms. You make your point very nicely. Each organism is manufacturing its own local environment.

RL: Exactly. That’s the wind chill factor. That’s why it gets colder when the wind blows.
DSW: Right. But that makes it a complicated evolutionary story. It’s still an evolutionary story, and when you just say history you’re leaving all that out. History seems to me too broad. Sure everything is history but we’d like to say something more specific. If there is a process of adaptation going on, even if it’s one of rapid niche construction and coevolution, that’s still a more specific set of ideas than just plain history, which really does encompass everything and therefore nothing. Don’t we want to use some of those more specific ideas about adaptation and coevolution and niche construction? That’s more than just history!

RL: Oh no, I’m with you! If I could convince people to use that notion of niche, not as a fixed thing, but as something that is manufactured by the organism, I would be very very happy. But when I talk to biologists about it, they’re always surprised.

DSW: It is still a new idea, in part of course because it’s a complex idea. Complexity is complex, it’s hard to study. We’re always trying to keep things simple, even when we should be embracing complexity in some sense.

DSW: What a pleasure, Dick! Thank you so much for this conversation. Have a great day.

RL: You too.