The Six Legacies of Edward O. Wilson

Note: An abbreviated version of this article is published in Nautilus Magazine.

Edward O. Wilson, who passed away at the age of 92 on December 26, 2021, is widely recognized as a giant of the Arts and Sciences. I include the Arts because Wilson regarded the creative dimension of science as an artistic endeavor, worked toward unifying the Arts and Sciences, and wrote beautifully for the general public, resulting in two Pulitzer prizes for nonfiction and one novel.

Wilson’s stature is so great, and reflections on his legacy upon his death are so numerous, that another reflection might seem unnecessary. The purpose of my reflection, however, is to make a novel point: Wilson left at least six legacies, which need to be combined to fully realize his vision. Combining the legacies of Edward O. Wilson requires first identifying them separately and then integrating them with each other.

The six legacies are:

1) His contributions to evolutionary biology.

2) His contributions to the conservation of biodiversity.

3) His contributions to a sociobiology that includes humans.

4) His contributions to the unification of knowledge.

5) His encouraging stance toward young scientists and other learners.

6) The new frontier that he was working on at the time of his death was ecosystems.  

My relationship with Edward O. Wilson

Before turning to these legacies and their integration, I will briefly recount my own relationship with Ed. I am 20 years younger so that he was already famous as a Harvard professor when I entered graduate school at Michigan State University in 1971. I first met him during the summer of that year. I was a student in an ecology course at the Marine Biological Laboratory in Woods Hole, Massachusetts. He was sitting in on the student project reports. After I reported my experiments on food size selection in zooplankton, Ed remarked “That’s new, isn’t it?” I was so proud to have impressed the great E.O. Wilson and contributed to the vast storehouse of scientific knowledge that I have remembered his comment ever since!

My graduate education was shaped in part by Ed’s influence on evolutionary biology, as I will elaborate below. My next personal interaction came near the end of my graduate career. I had constructed a mathematical model that provided support for the theory of group selection, which had been almost universally rejected by evolutionary biologists, as I will also elaborate below. Convinced of its importance, I wrote Ed asking if he would consider sponsoring it for the Proceedings of the National Academy of Sciences. Ed invited me to visit him at Harvard’s Museum of Comparative Zoology. As with my first encounter, I have a vivid memory of the visit, which began with a tour of his ant laboratory. Then he stood me in front of a blackboard, sat down in a chair, and said “you have 30 minutes until my next appointment.”

I talked like an auctioneer, filling the board with my equations. Ed was sufficiently intrigued to sponsor my article for PNAS after sending it out for review by two experts in theoretical biology. The article became my Ph.D. thesis, which is probably the shortest in the history of evolutionary science (four pages).

In the years that followed, I became one of the main advocates of group selection without directly crossing paths with Ed. I also took part in most of the other initiatives associated with Ed’s legacies without directly interacting with him. We were both involved in the formation of the Human Behavior and Evolution Society (HBES) and I hosted its third annual conference in 1993. On the theme of consilience, I started the first campus-wide program for teaching evolution across the curriculum and wrote one of the first book-length accounts of religion from an evolutionary perspective. It might seem strange that Ed and I shared so many interests without directly interacting, but just about everything associated with Ed’s legacies are in fact broad developments in the history of science involving many protagonists, a point to which I will return.

My next and by far most substantive interaction with Ed began at the 2006 annual conference of HBES. Ed was a plenary speaker and I was in the audience. Even though HBES members were in the avant-garde of studying human behavior from an evolutionary perspective, most of them were doctrinaire in their rejection of group selection. On his own, Ed had embraced group selection, converging on my own advocacy, and chose to break the news to the unsuspecting audience in his plenary. You could have heard a pin drop. Afterward, we found a corner of the lobby to talk alone.

“Did you like the grenade that I tossed in their midst?” Ed asked with a conspiratorial smile. On the spot, I suggested that we write a major article together, which became “Rethinking the Theoretical Foundation of Sociobiology”, published in the Quarterly Review of Biology in 2007. To reach a larger audience, we also wrote “Evolution for the Good of the Group”, which was published in the American Scientist in 2008. These were written by trading drafts and discussing them by email and phone. I still remember his voicemails, which sometimes went on for several minutes and were spoken in flawless extemporaneous prose.

At the end of our “Rethinking” article, we summarized our argument for group selection as the theoretical foundation of sociobiology by stealing from Rabbi Hillel, who was reputedly asked to explain the meaning of the Torah while standing on one foot and replied “What is hateful to you, do not do to your neighbor. Everything else is commentary.” Our one-foot version of sociobiology was: “Selfishness beats altruism within groups. Altruistic groups beat selfish groups. Everything else is commentary.” This meme has become widely known and Ed repeated it all the way up to his final publications and interviews.

After this intense collaboration, Ed and I went our separate ways to continue pursuing our largely overlapping interests. The last time I saw him was at a conference at MIT, which was close enough to his home that he could attend without arduous travel. In the few minutes that we spoke together, he told me excitedly about ecosystems as the next big topic that he planned to synthesize. He retained his youthful spirit of exploration right up to the end.

I have one more story about Ed to tell before turning to his six legacies. In 2014, the evolutionary psychologist Barry X. Kuhle recorded a series of interviews with pioneers of HBES, including both Ed and myself. Ed must have relished the opportunity to talk at a professional level with someone as well informed as Barry because his interview lasted two hours. I was president of the newly founded Evolution Institute and Editor in Chief of its online magazine This View of Life (TVOL), which was named after the final passage of Darwin’s On the Origin of Species (“There is grandeur in this view of life…”). I was eager to feature a print version of Barry’s interview with Ed on TVOL, so I offered to transcribe it myself. There is something about transcribing a recording, word by word, that burns it into your memory more than merely listening to the recording or reading the transcription. This experience adds to my knowledge of Ed and his legacies, along with his published work and my personal relationship with him.

The Six Legacies

History—including the history of science--is a complex systemic process involving many actors and environmental (including cultural) contingencies. Attention often becomes focused on a few key people, such Albert Einstein, Sigmund Freud, and B.F. Skinner, which under-represents the contributions of many dozens of others. Iconic status is thrust upon a person as much as actively sought by the person. There seems to be a need to personify ideas as a form of simplification, among the general public and even, to a degree, among the experts.  

A few evolutionary biologists such as Ed Wilson, Richard Dawkins, and the late Stephen Jay Gould have achieved this iconic status. Yes, they made outsized contributions as individuals, but they also represent something larger than themselves. I think that Ed would agree. In his book Sociobiology: The New Synthesis, for example, he was relying upon the work of many hundreds of scientists to support his claim that there can be a single theory of social behavior informed by evolution.

The world “catalyst” also bears examination. In chemistry, a catalyst is a substance that increases the rate of a chemical reaction without being used up in the process. The way a catalytic molecule works is by holding other molecules in an orientation that binds them to each other and releases the catalytic molecule to repeat the operation. A person can play a catalytic role in cultural change in much the same way. As we will see, Ed was a catalyst par excellence. He made things happen that otherwise would have occurred much more slowly or not at all.

Against this background, calling Ed an “icon” and a “catalyst” honors the individual while also going beyond the individual to examine systemic trends in the history of science. It is in this spirit that I will review his six legacies.    

1) His contributions to evolutionary biology.

Here is how Ed described his contribution to evolutionary biology in his interview with Barry Khule:

We have to go back to the 1950’s. In the 1950’s, the molecular revolution had begun. It was clear that the golden age of modern biology was going to be molecular and would endure a long time. In fact, it did occupy the second half of the 20^th century and beyond. We felt here at Harvard immediately the pressure to start giving up positions to molecular biology. The Dean of the faculty and the President at that time were entirely in accord. We found—I say we, the organismic and evolutionary biologists here, comparative anatomists, comparative zoologists and so on–realized that we would not to be given much additional space anymore, that we probably would not get many if any new positions for a long time. They would be reserved to build up Harvard’s strength in molecular and cellular biology. What this did was have a tremendous impact on me personally because I realized…that those of us, my generation of what we came to call evolutionary biologists and organismic biologists, were not going to get anywhere by complaining by any means but we were going to have to—and we should be tremendously excited to plan this—develop an equivalent to molecular biology on our own.

Ed then set about trying to modernize the biology of whole organisms, as part of a younger generation following the architects of the Modern Synthesis, which included names such as Ernst Mayr, Julian Huxley, and George Gaylord Simpson. This required finding and collaborating with people who had complementary expertise—especially the ability to build mathematical models of ecological and evolutionary processes. Names that Ed mentions as part of this younger generation include Robert MacArthur, Larry Slobodkin, and Richard Lewontin. These were some of the rock stars whose work I avidly read as a graduate student in the 1970s.

One of Ed’s most productive collaborations was with Robert MacArthur, an ecologist with mathematical training, leading to their landmark book The Theory of Island Biogeography, published by Princeton University Press in 1967 with Ed as the second author. What made the book so important was a theoretical framework that made sense of the great mass of natural history information on the distribution and abundance of species on islands—some of it collected by Ed for ant species around the world. The theory applied not only to actual islands but to all habitats that are island-like, such as mountains separated by valleys or patches of forest separated by deforested areas.  

While Ed played a prominent role in modernizing whole organism biology, he was by no means alone. Also during my time as a graduate student, a Nobel prize was awarded to Konrad Lorenz, Niko Tinbergen, and Carl von Frisch for pioneering the study of animal behavior and the geneticist Theodosius Dobzhansky titled an article for biology teachers “Nothing in biology makes sense except in the light of evolution”. Evolutionary theory was proving its explanatory scope and many people were taking part in the effort. What this meant to me as a graduate student was that I could choose any topic, begin asking intelligent questions based on evolutionary theory (often with the help of mathematical models), and then test my hypotheses on any appropriate organism. I didn’t need to become a taxonomic specialist and I could change topics at will. In short, I could become a polymath, based not on my personal attributes but on a theory that anyone can learn. This is the legacy of evolutionary biology, to which Ed made an outsized contribution.

2) His contributions to the conservation of biodiversity

As first and foremost a naturalist and ant taxonomic expert, Ed was passionate about the conservation of biological diversity and made room for it alongside his scientific career. His book Biophilia argued that we are genetically adapted to be surrounded by nature, with mental and physical health consequences if we are not. This bold conjecture has been largely supported by research. For example, hospital patients recover faster if their room has a window or is decorated with foliage and flowers.

Ed collaborated with Thomas Lovejoy, who coincidentally passed away just a day earlier at the age of 80, to preserve the biodiversity of the Amazon. According to a remembrance in the New Yorker magazine, it was they who coined the term biological diversity, which became shortened to biodiversity. They even drew upon the theory of Island Biogeography by studying the effect of the size of forest reserves on species loss.

With his gift for marketing whole disciplines and initiatives, Ed coined the term “Half Earth” for the goal of preserving half of the earth for nature and the other half for humankind—not in separation, but in a way that is interdigitated, so that humans can live within nature and nature can flow along corridors. Anyone who values nature should want to continue this legacy but doing so requires changing the minds and hearts of people, along with their cultural practices, in the real world.

3) His contributions to a sociobiology that includes humans

Ed’s 1975 book, Sociobiology: The New Synthesis, was in the same mold as Darwin’s “there is grandeur in his view of life” and Dobzhansky’s “nothing in biology makes sense except in the light of evolution”. Ed’s claim was that evolutionary theory provides a single conceptual toolkit for studying the social behaviors of all creatures great and small. Thanks to Ed’s gift for identifying whole fields of inquiry and writing for non-specialists, Sociobiology combined the authority of an academic tome with the look and feel of a coffee table book, complete with over 200 illustrations by the artist Sarah Landry. Thanks to his stature and gift for promotion, its publication was noted on the front page of the New York Times.

It was the last chapter on human social behavior that landed Ed in trouble and a systemic view of the history of science is needed to understand why. For all its explanatory scope, the study of evolution was restricted to genetic evolution for most of the 20^th century, as if the only way that offspring can resemble their parents is by sharing the same genes. This is patently false when stated directly since it ignores the cultural transmission of traits entirely, but it essentially describes what became known as the modern synthesis and was consolidated by the molecular biology revolution described by Ed in his interview with Barry Kuhle.

What became of the study of cultural evolution? It was ceded to other disciplines in the human social sciences and humanities. Each discipline developed into a sophisticated body of knowledge, but not in reference and sometimes in perceived opposition to evolutionary theory. And all of those disciplines did not remotely become integrated with each other. Instead, they became an archipelago of knowledge with little communication among the islands. The lack of consilience for human-related knowledge stands in stark contrast with the consilience of biological knowledge, at least when it comes to genetic evolution.

Darwin’s theory is often said to have earned a bad reputation for itself in the human-related disciplines by providing a moral justification for inequality (Social Darwinism). The real history of Darwinism in relation to human affairs is more complex and interesting. Socialists such as Peter Kropotkin and progressive thinkers such as William James and John Dewey were inspired by Darwin along with “nature red and truth in claw” types. The bottom line is that any powerful tool can also be used as a weapon and Darwin’s theory is no different than any other theory in this regard.¹

Returning to the reception to Sociobiology, when critics accused Ed of genetic determinism, they were absolutely right. The entire field of evolutionary biology was gene-centric and Ed was no exception. Yet, critics from the human social sciences and humanities had no synthesis of their own.

Only after the publication of Sociobiology did evolutionary thinkers begin to take cultural evolution seriously. Ed was among them with books such as On Human Nature, Genes, Mind, and Culture (with Charles J. Lumsden), Promethean Fire (also with Lumsden), and The Social Conquest of Earth. Other major thinkers included Richard Dawkins and his concept of memes, Luigi Luca Cavalli-Sforza and Marcus Feldman (Cultural Transmission and Evolution), and Robert Boyd and Peter Richerson (Culture and the Evolutionary Process, Not By Genes Alone). The importance of symbolic thought began to occupy center stage with books such as The Symbolic Species by Terrence Deacon and Evolution in Four Dimensions by Eva Jablonka and Marion Lamb.

Today, Darwinian evolution is widely defined as any process that combines the three ingredients of variation, selection, and replication, no matter what the mechanism of replication. This definition is true to Darwin’s thought (since he knew nothing about genes) and can accommodate a plurality of inheritance mechanisms such as epigenetics (based on changes in gene expression rather than gene frequency), forms of social learning found in many species, and forms of symbolic thought that are distinctively human. While human cultural inheritance mechanisms evolved by genetic evolution, that doesn’t make them subordinate, as if genes hold cultures on a leash (one of Ed’s metaphors). On the contrary, as the faster evolutionary process, cultural evolution often takes the lead in adapting humans to their environments, with genetic evolution playing a following role (gene-culture co-evolution).

Part of the maturation of human cultural evolutionary theory is the recognition of group selection as an exceptionally strong force in human evolution—something else that Ed got right. According to Harvard evolutionary anthropologist Richard Wrangham in his book The Goodness Paradox, naked aggression is over 100 times more frequent in a chimpanzee community than in small-scale human communities. This is due largely to social control mechanisms in human communities that suppress bullying and other forms of disruptive self-serving behaviors so that cooperation becomes the primary social strategy (this is called a major evolutionary transition). Nearly everything distinctive about our species is a form of cooperation, including our ability to maintain an inventory of symbols with shared meaning that is transmitted across generations. Our capacity for symbolic thought became a full-blown inheritance system that operates alongside genetic inheritance (dual inheritance theory). Cultural evolution is a multilevel process, no less than genetic evolution, and the increasing scale of cooperation over the course of human history can be seen as a process of multilevel cultural evolution.

While the critique of genetic determinism was accurate for Sociobiology and evolutionary biology as a whole in 1975, this is no longer the case for the modern study of humans from an evolutionary perspective—which brings us to Ed’s next legacy.

4) His contributions to the unification of knowledge.

Something that can be said about Ed’s books is that they are all visionary—imagining whole new fields of inquiry—but vary in the degree to which Ed has made progress carrying out the vision. He made the most progress for ants and other social insects, of course, and Sociobiology reflected a thorough reading of the literature on animal social behaviors. A book such as Consilience, however, is long on vision and short on execution.

I do not intend this observation as a criticism. Ed had only 24 hours in a day, like the rest of us, and his visionary gaze is worthwhile even if the execution is left to others. In Consilience, the vision is “a conviction, far deeper than a mere working proposition, that the world is orderly and can be explained by a small number of natural laws (p4)”. While this vision stretches back to antiquity and includes knowledge of the physical world in addition to the living world, there is something about evolutionary theory that fulfills the vision for the living world in an extraordinary way. Here is how Ed describes his first encounter with evolutionary theory in the opening pages of Consilience. He’s an 18-year old kid newly arrived at the University of Alabama, with a passion for identifying plants and animals using field guides.

Then I discovered evolution. Suddenly—that is not too strong a word—I saw the world in a wholly new way. This epiphany I owed to my mentor Ralph Chermock, an intense, chain-smoking young assistant professor newly arrived in the provinces with a Ph.D. in entomology from Cornell University. After listening to me natter for a while about my lofty goal of classifying all the ants of Alabama, he handed me a copy of Ernst Mayr’s 1942 Systematics and the Origin of Species. Read it, he said, if you want to become a real biologist.

The thin volume in the plain blue cover was one of the New Synthesis works, uniting the nineteenth-century Darwinian theory of evolution and modern genetics. By giving a theoretical structure to natural history, it vastly expanded the Linnaean enterprise. A tumbler fell somewhere in my mind, and a door opened to a new world. I was enthralled, couldn’t stop thinking about the implications evolution has for classification and for the rest of biology. And for philosophy. And for just about everything. Static pattern slid into fluid process…A new enthusiasm surged through me. The animals and plants I loved so dearly reentered the stage as lead players in a grand drama. Natural history was validated as real science.

Coincidentally, Ernst Mayr’s Animal Species and Evolution was one of the first evolution books that I read as an undergraduate student. While it was not thin (811 pp!), I was similarly enthralled. Compare Ed’s epiphany with passages from Charles Darwin, such as “I can remember the very spot on the road…” and “he who understands the baboon would do more toward metaphysics than Locke”, which was scribbled in his notebook in 1838. There is something about the simplicity and generality of evolutionary theory that starts working at the very beginning, for Darwin as the originator and Ed Wilson as an unschooled kid. Now recall what I said about being a graduate student in the 1970s—that I could become a polymath, based not on my personal attributes but on a theory that anyone can learn. What this means is that by the 1970s, what Darwin and Ed glimpsed from the start was now proving itself for the length and breadth of the biological sciences. Every time an evolutionary biologist decides to switch to a new topic and/or organism--which happens all the time—consilience is being demonstrated in action.

The prospect that human-related knowledge can become unified in this way is both old and new. It was how Darwin thought and he originated group selection theory as much to explain human morality as “for the good of the group” traits in nonhuman species. But you can’t make sense of humanity without acknowledging its groupish nature and the importance of culturally transmitted symbolic meaning systems. As Emile Durkheim wisely put it: “Social life, then, in every aspect and throughout its history, is only possible thanks to a vast body of symbolism.” Only now are we in a position to synthesize human-related knowledge in the same way as biological knowledge, thanks to an expanded definition of Darwinism as any variation/selection/replication process. Ed’s vision in Consilience is right on and its fulfillment is now in progress.

5) His encouraging stance toward young scientists and other learners.

No remembrance of Ed would be complete without noting the way that he encouraged people to become scientists, to follow their hearts, and to cultivate a reverence for nature. Visit #eowilson on Twitter and you’ll find quotes such as these offered by those whose lives he touched.

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“Adults . . . are prone to undervalue the mental growth that occurs during daydreaming and aimless wandering.” -- The late great Edward O. Wilson

“Nature first, then theory. Love the organisms for themselves first, then strain for general explanations, and with good fortunes discoveries will follow.”

“You are capable of more than you know. Choose a goal that seems right for you and strive to be the best, however hard the path. Aim high. Behave honorably. Prepare to be alone at times, and to endure failure. Persist! The world needs all you can give.”

“Nature holds the key to our aesthetic, intellectual, cognitive and even spiritual satisfaction.

"There can be no purpose more enspiriting than to begin the age of restoration, reweaving the wondrous diversity of life that still surrounds us.”

“The evolutionary epic is the best myth we will ever have.”

“You teach me, I forget. You show me, I remember. You involve me, I understand.”

“Humanity is part of nature, a species that evolved among other species. The more closely we identify ourselves with the rest of life, the more quickly we will be able to discover the sources of human sensibility and acquire the knowledge on which an enduring ethic, a sense of preferred direction, can be built.”

Passages such as these spell the difference between science and a science-based worldview. By itself, science merely tells us what is. A worldview provides a sense of values and motivates action. A science-based worldview does this based on reverence of the natural world rather than a supernatural agency. Ed is remembered at least as much for the science-based worldview that he offered as his scientific discoveries.

6) Ecosystems as Ed’s final frontier

Ed’s next book was to be titled “Ecosystems and the Harmony of Nature”. I don’t know if it will be published posthumously but we can get a glimpse of what he had in mind from its title, a brief article on the E.O. Wilson Biodiversity Foundation website,² and a short lecture on YouTube.³

In the article, Ed is quoted as saying: “We know that ecosystems, which are really what we are trying to protect--not just single species but ensembles of species that have come together and have reached the ability—sometimes over thousands or even in some places millions of years—have formed ecosystems that equilibrate. And we don’t really know how equilibration comes about.” Ed also encourages young people to join “the coming development of a new biological science, one of the next big things, which is ecosystem studies.”

I must confess that I am puzzled by these statements since the study of whole ecosystems dates back to the beginning of the 20^th century and has become increasingly integrated with evolutionary ecology over the last 50 years. It turns out that multilevel selection theory is essential for understanding the nature of ecosystems, no less than single species societies. I will be fascinated to know if Ed has converged upon this conclusion.

To explain what I mean, a critical distinction needs to be made between two meanings of the term “complex adaptive system (CAS)”: A complex system that is adaptive as a system (CAS1), and a complex system composed of agents following their respective adaptive strategies (CAS2). A human society in the grip of civil war is an example of CAS2. It can be understood in terms of the conflicting interests of the warring factions, but it does not function well at the level of the whole society (CAS1) and no one would expect it to.

Many single-species societies in nature are like my human civil war example. Members of social groups are largely in conflict with each other and at most cooperate in specific contexts. We need look no further than chimpanzee communities for an example, where naked aggression is over 100 times more frequent than in small-scale human communities and the main context for community-wide cooperation is aggression against neighboring communities. Social strife in chimpanzee communities is stable—there is no reason to expect it to change, given the selection pressures that are operating—but that doesn’t make them harmonious or desirable from a human perspective.

Many multispecies ecosystems are also like this. For example, if you want to understand the nature of beaver ecosystems, ask the question “what’s in it for the beavers?” They are modifying the environment for their own benefit, flooding it to protect themselves from predators and eating the most palatable plants. Consequences for biodiversity and ecosystem processes such as nutrient cycling are collateral effects of beavers pursuing their interests. There is no reason to expect the whole ecosystem to be functionally organized and harmonious, any more than a chimpanzee community or a human society in the grip of civil war.

This is a hard lesson to learn about nature. We want it to be harmonious. Religious cosmologies often portray nature as harmonious (e.g., the Garden of Eden) except when disturbed by humans. The early study of ecosystems often treated them axiomatically as harmonious.  But Darwin’s theory of evolution tells a different story. It tells us that functional organization for any given system, at any given scale, requires a process of selection at that scale. That is the only way to achieve the status of CAS1 rather than merely CAS2, where functionally organized agents impose suffering on each other in the course of pursuing their respective adaptive strategies. That statement goes for human society, single-species animal societies, and multispecies ecosystems.  

Are there examples of whole ecosystems that have evolved into superorganisms? Yes! Microbiomes are an example. Every multicellular organism is not only a collection of mostly identical genes but also an ecosystem composed of trillions of microbes comprising thousands of species. When the host organisms differentially survive and reproduce, this is due in part to variation in their microbiomes along with variation in their genes. Thanks to selection at this level, microbiomes have evolved to be largely mutualistic with their hosts. There is also potential for selection among microbes within each host, however, leading to the evolution of pathogenic strains. It all depends on the level of selection.

Nowadays, whole forests are being imagined as mutualistic networks, with trees connected into a network by mycorrhizal fungi. Is such a thing possible? Yes, but only if selection has operated at the scale of whole forests with sufficient strength to counteract selection at lower scales. Otherwise, forests become merely CAS2 systems, composed of species that interact at cross purposes, rather than CAS1 systems.

Above all, it is important to avoid confusing “harmony” with “equilibrium”. Ecologists have started to use the word “regime” to describe stable assemblages of species. This is a well-chosen word because it evokes what we already know about human political regimes. All political regimes have a degree of stability, or we wouldn’t call them regimes, but they span the range from despotic (benefitting a few elites at the expense of everyone else) to inclusive (sharing their benefits with all citizens). Some of the worst regimes are also depressingly the most stable. Using the language of complex systems theory, there are multiple local stable equilibria and positive change requires escaping the gravitational pull of one local equilibrium to enter another local equilibrium. This requires active management and will not necessarily happen by itself. The management of ecosystems must itself be a human cultural evolutionary process informed by multilevel selection theory.

Combining the legacies

In this remembrance of Ed Wilson, I have tried to honor the person while also placing him in the context of broad trends in the history of science. Without mentioning Ed, we can say that Darwin’s theory of evolution has an amazing explanatory scope, that this scope was largely restricted to the study of genetic evolution for most of the 20^th century, but now is rapidly expanding to include all aspects of humanity in addition to the rest of life. As I put it in my own book This View of Life: Completing the Darwinian Revolution, Dobzhansky’s statement “nothing in biology makes sense except in the light of evolution” can be extended to include everything associated with the words “human”, “culture”, and “policy”.

Without mentioning Ed, we can also say that evolutionary theory is capable of functioning as a worldview in addition to a body of scientific knowledge. Science only tells us what is, whereas a worldview inspires us psychologically and moves us to action. Creating a worldview informed entirely by science, as opposed to supernatural belief, is part of the enlightenment project that led to humanism as a philosophical worldview and social movement. While humanists accept Darwin’s theory as a matter of course, the recent developments that I have recounted have not been incorporated into the humanist movement for the most part. Thus, humanism and what it stands for is due for a renaissance, along with a renaissance of basic scientific knowledge.

Some simple calculations will help to put Ed’s career into historical perspective. Starting from when he received his Ph.D. in 1955 to his death in 2021, his career lasted for 66 years. If we mark the beginning of evolutionary science with the publication of Darwin’s On the Origin of Species in 1859, then Ed was present for 40% of the history of evolutionary thought. If we mark the beginning of the scientific revolution with the publication of Copernicus’s On the Revolution of the Heavenly Spheres in 1543, then Ed was present for 14% of the scientific revolution. As 20 years Ed’s junior, my numbers work out to 28% and 10% respectively.

These numbers remind us that evolutionary science and the scientific revolution are still works in progress. If science in general and evolutionary science, in particular, have revolutionized the way we see and therefore act upon the world, then we can look forward to further improvements in the near future. This leads to a form of hope and optimism, even in the darkest of times, that is part of Ed’s legacy.

For me, the next frontier is not just ecosystems but becoming wise stewards of evolution in all its forms. Variation/selection/replication processes are taking place all around us at different time scales, including genetic evolution, cultural evolution, and intra-generational personal evolution. Without wise stewardship, these evolutionary processes result merely in CAS2—complex systems composed of agents following their respective adaptive strategies, often inflicting harm on each other and on the entire system over the long term. Work is required to transform CAS2 into CAS1—systems that are adaptive as whole systems. This work will be required for all forms of positive change—individual, cultural, and ecosystemic. The ability to see this clearly and to act upon it has only become available during the last few decades and is currently shared by only a tiny fraction of those who need to know about it. Catalysis is needed, so that positive evolution can take place in a matter of years rather than decades or not at all. The best way to honor Ed’s combined legacies is to join in this catalysis.

References:

[1] For more, see the TVOL special edition titled “Truth and Reconciliation for Social Darwinism”.

[2] https://eowilsonfoundation.org/inspiring-a-new-generation-to-fight-for-biodiversity/

[3] https://thefestivalofdiscovery.com/session/watch-now-e-o-wilson-ecosystems-and-the-harmony-of-nature/