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David Sloan Wilson
David Sloan Wilson
is the SUNY Distinguished Professor of Biology and Anthropology at Binghamton University and Arne Næss Chair in Global Justice and the Environment at the University of Oslo

For anyone who thought that TVOL would pass down the Word, based on the authority of Science, think again. Science is a boiling cauldron of disagreement. The only thing distinguishing it from other boiling cauldrons is that it is supposed to result in progress rather than endless gridlock. Progress is the accumulation of knowledge, which starts as untested hypotheses and ends with something durable enough to be called fact. The earth is round. It is also very old. Continents drift. All of these claims were once hotly contested, but then science did its magic and they are now regarded as facts.

Science is not necessarily a gentlemanly sport, like tiddlywinks. It can be a contact sport, like boxing—but even boxing has rules and referees to make sure that the hitting is above the belt.

Science journalism reports on science. A science journalist must explain to nonscientists why a given topic is worth caring about, which often involves poetic license and omitting the minutia. Otherwise, science journalists are bound by the same code of ethics as scientists, such as respecting the authority of empirical evidence, avoiding ad hominem attacks, and so on.

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Against this background, welcome to the hard-hitting world of group selection and selfish genes. The controversy seems to persist forever. Is that because the topic is so very complicated, or because the data is so hard to collect? Or might it be the kind of futile gridlock that occurs when some of the participants stop functioning as scientists and start hitting below the belt?

In the most recent round, I came out swinging in an article titled “When Richard Dawkins is not an Evolutionist”. That might seem below the belt, but I chose the word “when” carefully. When does anyone function as an evolutionist, which also requires functioning as a scientist? In other words, what constitutes hitting above and below the belt? It’s not hitting below the belt to ask for a clarification of the rules!

Then I claimed that Dawkins was hitting below the belt on two subjects: religion as a human construction and selfish genes in relation to group selection. On religion as a human construction, there is a community of scientists who hit above the belt, they are making progress, and Dawkins is not among them. Readers who want to learn more should consult evolutionists such as Scott Atran, Jesse Bering, Pascal Boyer, Joseph Bulbulia, Joseph Henrich, Dominic Johnson, Richard Sosis, Harvey Whitehouse, and TVOL’s RELIGION editor Michael Blume, to name a few. ETVOL is proud to provide a journalistic forum for their work.

It would take a long time to document how Dawkins departs from factual reality on the subject of religion as a human construction, but selfish genes in relation to group selection is more straightforward: The concept of genes as replicators was initially regarded as a drop-dead argument against group selection, but it proved to be nothing of the sort. In a genetic model, genes are the replicators regardless of whether group selection does or doesn’t occur. Group selection occurs when genes (or memes in a cultural model) evolve in the total population, despite being selectively neutral or disadvantageous within groups (as a technical aside, a trait can also be favored by both within- and between-group selection). This can be regarded as a fact, as surely as a round earth, deep time, and drifting continents. It’s understandable that people might have been ignorant of the fact before (as with those other facts), but there is little excuse for an evolutionist today, since it was established decades ago. My claim is that Dawkins evades this fact and continues to treat selfish genes as an argument against group selection. Whether I am hitting above or below the belt depends upon the veracity of my claim.

In response to my article, the distinguished evolutionist Jerry Coyne came out swinging on his Why Evolution Is True blog. Because the mission of TVOL is to feature controversy, and not to present just one side, we immediately carried the story on our front page. Part of hitting above the belt involves acknowledging areas of agreement into addition to areas of disagreement. In my article, for example, I agreed with Dawkins on the lack of empirical evidence for supernatural agents that actively intervene in physical processes and the affairs of people, before disagreeing with him on the nature of religion as a human construction. I am now pleased to indicate my areas of agreement and disagreement with Coyne.

First and foremost, Coyne and I agree that selfish genes do not constitute an argument against group selection. Here is Coyne in his own words:

Yes, genes are replicators, but no, Dawkins never claimed that their status as selfish replicators somehow rules out group selection. What he claimed, in The Selfish Gene and The Extended Phenotype, was that successful replicators must share the same vehicle if they are to be successful in the future. Usually that vehicle is the body of an individual organism, which is used by the replicators to propagate themselves. Dawkins’s argument against the efficacy of group selection was that this form of selection is usually unsuccessful because groups are vulnerable to subversion from within by those selfish replicators. That is, “cheating” replicators that are “good” for individuals but bad for the group as a whole will tend to propagate themselves. Yes, altruism may help groups propagate, but altruistic groups are susceptible to invasion by cheaters unless the “altruism” is based on kin selection or individual selection via reciprocity.

Coyne has made my work easy. Any competent historian of the subject knows that the “group selection is wrong because genes are replicators” argument was the received wisdom during the 1960’s and 70’s. Here is one of dozens of passages that could be quoted, this one from Richard Alexander in 1979:

In 1966 [G.C.] Williams published a book criticizing what he called “some current evolutionary thought” and chastised biologists for invoking selection uncritically at whatever level seemed convenient. Williams’ book was the first truly general argument that selection is hardly ever effective on anything but the heritable units of “genetic replicators” (Dawkins, 1978) contained in the genotypes of individuals.

I have discussed Dawkins’s role in the initial confusion elsewhere (here and here). For Coyne to claim that no one was ever confused on this point is historical revisionism, and that’s hitting below the belt.

Dawkins did acknowledge that group selection is a matter of vehicles, not replicators, in The Extended Phenotype, published in 1982. That’s why I was surprised when he took it back in 2007, in a response to an article written by Edward O. Wilson and myself in the American Scientist:

Genes Still Central: David Sloan Wilson’s lifelong quest to redefine “group selection” in such a way as to sow maximum confusion–and even to confuse the normally wise and sensible Edward O. Wilson into joining him–is of no more scientific interest than semantic double talk ever is. What goes beyond semantics, however, is his statement (it is safe to assume that E.O. Wilson is blameless) that “Both Williams and Dawkins eventually acknowledged their error [that the replicator concept provides an argument against group selection]…I cannot speak for George Williams but, as far as I am concerned, the statement is false: not a semantic confusion; not an exaggeration of a half-truth; not a distortion of a quarter truth; but a total, unmitigated, barefaced lie. Like many scientists, I am delighted to acknowledge occasions when I have changed my mind, but this is not one of them. D.S. Wilson should apologize. E.O. Wilson, being the gentleman that he is, probably will.

So, according to Dawkins, the status of genes as replicators is still central to the evaluation of group selection. That makes him a flat-earther on the subject and anyone refereeing the scientific process should call him out—including Jerry Coyne.

With someone of Dawkins’ stature saying whatever he pleases, it’s understandable that the rest of the world remains confused. Here is a passage from one of the people who commented on Coyne’s post:

Yes, it’s necessary to once again point out that individuals cooperating as a group is NOT evidence for group selection, in itself. This sort of behaviour has been shown, time and time again, to be the result of selection at the level of the individual’s genes.

The comments on Coyne’s post also demonstrate rampant confusion on the question of whether kin selection is a type of group selection or an alternative to group selection, another issue that was settled decades ago (go here). There is virtually no awareness among Coyne’s reading public of the concepts of equivalence and pluralism, which is the current frontier of debate (go here). I’m not to blame for this degree of illiteracy—go ask Coyne or Dawkins!

Coyne’s hard-hitting post in response to my article is a veritable flurry of below-the-belt punches, including:
• Questioning the mental competence of one’s opponent (e.g., “losing it again”, “totally over the waterfall”, “sheer madness”).
• Misrepresenting the number of scientists who accept group selection as a significant evolutionary force (e.g., “futile one man crusade”).
• Guilt by association (my association with the John Templeton Foundation, which evidently taints everything I have said or done; more about that in the future).
• Misrepresenting the amount of evidence for group selection, as in this passage:
The concept of genes as selfish replicators, which has held up perfectly well since The Selfish Gene was published in 1976, says nothing about the efficacy of group selection. Dawkins’s (and my) beef with group selection as a way to evolve traits that are bad for individuals but good for groups is that this form of selection is inefficient, subject to subversion within groups, and, especially, that there’s virtually no evidence that this form of selection has been important in nature).

Once again, Coyne makes my work easy. Keep in mind that he is a fly geneticist who hasn’t published a single peer-reviewed article on group selection. Somehow, my own madness results in peer-reviewed publications in top journals, and not just me but dozens of my colleagues who straightforwardly demonstrate the evolution of genes and memes in multi-group populations that are selectively disadvantageous within the groups. For every colleague who dares to use the term “group selection” to describe this process, there are others who strip the term from their manuscripts to avoid harassment by people like Dawkins and Coyne, which only perpetuates the confusion, as Omar Eldakar and I describe in an article titled “Eight Criticisms Not to Make About Group Selection” published in Evolution.

For readers who are up for a challenge and want to learn more about the theoretical basis and empirical evidence for group selection from someone other than myself, I recommend Steven A. Frank’s “Natural Selection. III. Selection vs. Transmission and Levels of Selection (Journal of Evolutionary Biology, 2011). For Frank, it goes without saying that natural selection is a multilevel process and that the group level is often a significant evolutionary force.

Readers can decide whether I punch above or below the belt in my own articles. I expect to be refereed along with everyone else. But there is one move that surely qualifies as “not science” – the decision not to engage at all. Refusing to talk is one of the main reasons why other boiling cauldrons of disagreement fail to result in progress. If my evolutionist colleagues such as Coyne and Dawkins fail to show up for the match, then they’re not functioning as scientists for sure.


Join the discussion


  1. Synthetic says:

    Please, don’t.
    If you want this magazine to succeed don’t bring it down to the level of Coyne’s or Dawkins’ blogs. I understand your desire to set the record straight, and I’m entirely sympathetic to your point, but there is a reason I’m not reading Coyne’s or Dawkins’ blogs: I want to be informed, educated and not tho be the spectator of childish bickering between navel-grazers. I appreciate intellectual discussions and diverging opinions, but I’m not interested in personal attacks (and their responses) and bruised egos.

  2. Synthetic says:

    *navel gazers. Although, I sort of like the image of Coyne and Dawkins grazing on their respective navels.

  3. mikmik says:

    I am thankful for this article, David, and as a source of links to opposing opinions and research, is very enlightening for me. I rely on others to provide direction to relevant information, and I do not trust Coyne to provide it.
    You have a different atmosphere on your posts here, David, one that is professional and not personally motivated. It is important to challenge detractors when they are popular and sway, or contribute, to general opinion. I find ad hominum and denigration to be petulant and makes me very wary, but learning the actual details and reasons behind disputes is difficult in Jerry’s case (and others).
    It is important to expose these instances.
    I agree with Synthetic, succumbing to bickering is a no go, but you haven’t done that here. I find this to be an article that defends scientific integrity.

  4. Gerstle says:

    I’m going to go out on a limb here and suggest that David was thinking strategically when he ‘came out swinging’ at Dawkins on ETVOL, rather than on one of his blogs where (arguably) that kind of op-ed piece belongs. His charges against Dakwins would draw the latter’s defenders and add valuable online ‘hits’ to this emerging website.

    Unsurprisingly, Dawkins himself did not comment on David’s charges (which apparently means he’s not a scientist). But Jerry Coyne did, and so Jerry Coyne had to suffice. And now we get this follow-up essay on how uninformed Coyne is. David could not post this on his blog either, considering the majority of his submissions there from the last year already deal with that same topic.

    Anyway, we all enjoy a good fight, so I hope everyone keeps it up.  This behavior is not entirely conspicuous as opportunistic showmanship, and I’m sure this webzine will benefit from the spectacle. Here’s to the proliferation of your facebook likes, bookmarks, and tweets!

  5. David Sloan Wilson says:

    Thanks to everyone for their comments so far. Here are a few thoughts in return.

    1) I emphatically do NOT want the topics that concern me most to “take over” the magazine. There are eleven section editors and each has their own portfolio of interests. We want to develop a culture for the magazine in which each editor articulates his or her vision of evolution without any form of censure. If there are disagreements, then that is fodder for the magazine. All legitimate controversies are welcome and none should be excluded.

    2) We are discovering that the magazine is an evolving entity in its own right. Issues bubble up, and the beauty of the magazine is that we’re able to respond to them quickly with original content. For me, two issues that have risen to the fore are the nature of science and the nature of science journalism. It’s easy to wax poetic about science—I certainly do—but what distinguishes it from other forms of discourse and why is it supposed to result in more progress, if in fact it does? How does science journalism differ from science or other forms of journalism. What liberties can be taken by science journalists in the name of interesting a non-scientific audience, and what is off-limits to science journalists, no less than to scientists? What authority does a professional evolutionist have when writing for the general public on a topic that is not their speciality? These are deep questions and I look forward to featuring them as part of my contribution to the content of the magazine, as one of multiple visions of “this view of life”.

    3) There is tremendous insight in viewing science as like a sport, with rules and the need for refereeing, just like any other sport. You can’t find a single sports match on TV that doesn’t have referees watching every move and calling fouls during each and every game. Without rules and referees, sports matches would quickly devolve into chaos as the urge to win erodes the rules with no counterforce to maintain them. There seems to be an implicit assumption that scientists are so well behaved that they don’t need referees, or that the refereeing process that takes place at the professional level (e.g., peer reviewed journals) are sufficient to keep them in line. On the contrary, it seems to me that scientific discourse is badly in need of refereeing and popular science discourse even more so. In the absence of refereeing, scientists stop functioning like scientists and they are frankly not worth listening to. Policing is required and it’s not churlish to throw down the flag when there’s an infraction. I look forward to developing this theme and clarifying the science and science journalism rulebooks on the virtual pages of ETVOL smile


  6. Notanevolutionist says:

    Immortal Coil – “There we saw that selfishness is to be expected in any entity that deserves the title of a basic unit of natural selection. We saw that some people regard the species as the unit of natural selection, others the population or group within the species, and yet others the individual. I said that I preferred to think of the gene as the fundamental unit of natural selection, and therefore the fundamental unit of self-interest. What I have now done is to define the gene in such a way that I cannot really help being right!”………

    “In sexually reproducing species, the individual is too large and too temporary a genetic unit to qualify as a significant unit of natural selection.(3) The group of individuals is an even larger unit. Genetically speaking, individuals and groups are like clouds in the sky or dust-storms in the desert. They are temporary aggregations or federations. They are not stable through evolutionary time. Populations may last a long while, but they are constantly blending with other populations and so losing their identity. They are also subject to evolutionary change from within. A population is not a discrete enough entity to be a unit of natural selection, not stable and unitary enough to be ‘selected’ in preference to another population.”

    Is this not good enough evidence?

  7. Notanevolutionist says:

    “If anything is hitting below the belt here, this is. Actually, it’s not even hitting below the belt, it’s just immaterial.”

    Which one is it!? Why even state the first sentence if it is irrelevant?

    “Writers do not control what their audience chooses to read, how well they understand it, or how effectively they reproduce it”

    Bogus. Science journalism and peer reviewed publication is a communication system that is suppose to hold authors accountable for they say so that readers are not misguided by falsehoods. Are you arguing that Coyne and Dawkins don’t influence readers’ decisions?

    “I lean toward Coyne and Dawkins (though apparently not your interpretation of Dawkins), but I also believe there is an important and under-appreciated role for multi-level selection.”

    You lean toward what?—-Do you mean you lean toward errors and falsehoods? I hope not.


  8. evolutionist1 says:

    I think your tone in this post is quite respectable and level-headed, which is more than I can say for your first post or indeed for Coyne’s reply, which was bordering on apoplectic.

    It’s disappointing to me that you seem to base your belief (that Dawkins makes the genes-as-units-of-selection-ergo-no-group-selection argument) on his extremely brief reply (“genes still central”) to your article in New Scientist. In your original post, you argued: “A major objective of The Selfish Gene was to argue against a theory known as group selection, whereby traits such as altruism evolve ‘for the good of the group’, despite being selectively disadvantageous within groups. Dawkins and others at the time regarded the replicator concept as a drop-dead argument against group selection…” 
    However, in your current post, you recognize that “Dawkins did acknowledge that group selection is a matter of vehicles, not replicators, in The Extended Phenotype, published in 1982.”  I’m glad you finally recognize that neither Selfish Gene nor Extended Phenotype made this argument. If the 2007 Dawkins reply seems to make an about-face in this regard (and I admit it does appear this way) then it is an unfortunate error on his behalf, but one has to admit that his peer-reviewed publications do not take this position.

    Lastly, I don’t see what point there is in you, the eminent David Sloan Wilson, picking a fight with one of the commenters on Coyne’s blog. If anything is hitting below the belt here, this is. Actually, it’s not even hitting below the belt, it’s just immaterial. Writers do not control what their audience chooses to read, how well they understand it, or how effectively they reproduce it. I could just as easily point to an uninformed commenter on your own post (“when dawkins is not…”) who mistakenly believes that Dawkins makes the genes-as-units-ergo-no-group-selection argument in the Immortal Coils chapter of Selfish Gene. Regardless of whether Dawkins ever believed this, or believed it in 2007, or believes it now, it’s not in this chapter. Is it your fault, Dr. Wilson, that this commentor made this error? Nope.

    but all in all, I’m glad you followed up with a post such as this. Kudos to you. I lean toward Coyne and Dawkins (though apparently not your interpretation of Dawkins), but I also believe there is an important and under-appreciated role for multi-level selection.

    Best of luck with the website! may it inspire critical thought

  9. evolutionst1 says:

    RE: notanevolutionst, 12:25
    first quotation: describes that the units of selection must be selfish. this has no bearing on group selection.
    second quotation: describes that groups are not stable enough to be “units” of natural selection. Again, no bearing on group selection.
    I believe there are two questions that you are conflating:
    1) whether groups can be “selected” in relation to other groups in the same way that “units” are “selected”
    2) whether it is possible for between-group competition to cause directional change in gene frequencies.
    Dawkins disagrees with 1 but does not close the door on 2.  Thus the “tendency” for selection at lower levels to undermine that at higher levels, about which Dawkins and everyone else is clear.  Yet Wilson and others seem to imply that Dawkins makes the claim: 1 is false, therefore 2 cannot be true.  This is a misrepresentation of Dawkins, in my opinion.

    RE: notanevolutionst, 12:55
    why state the first sentence? I wrote in conversational tone. My mistake; I’ll more carefully edit my comments in the future. Thanks for holding me to task. The answer is I don’t know, it’s subjective. my belief, however, is that it’s immaterial.
    Am I arguing that Coyne and Dawkins don’t influence readers’ decisions? No I’m not. This is not a question of influence, it is a question of control. We all influence each other. My point is that an ill-informed comment on Coyne’s website is no more Coyne’s fault for mis-educating his readership than it is Wilson’s fault for failing to be more persuasive such that Coyne’s readers are drawn to the light of truth. There is always influence, but communication systems are not as simple as you portray them. For example, research on confirmation bias suggests readers go out in search of writing and evidence that supports their pre-conceived beliefs. In this sense, writers have very little control over the content of their readers’ beliefs. this is murky ground, thus my belief that to hold coyne accountable for the comments on his page tells us little of relevance to this debate.
    I lean toward belief in the gene as the unit of selection, as well as the plausibility of group and multi-level selection. when you take a closer look at all of these debates, you realize that the space between you and your enemies is narrower than you would have guessed.

  10. Tim Tyler says:

    D. S. W. – it seems as though you had to edit the “Dawkins eventually acknowledged their error” quote – by inserting material in square brackets.  Why not simply interpret Dawkins’ denial as a denial that he ever made this error in the first place?

  11. Tim Tyler says:

    If Dawkins “acknowledged his error” regarding group selection as claimed – where did he do that?  I for one don’t recall him doing that so far.  He doesn’t seem to recall doing that either.  Did it ever happen?  Or is this a fictional event.

  12. Synthetic says:

    Let me qualify what I meant by bickering between navel gazers, as I think that it is relevant to your initial question on the nature of science journalism.
    My impression is that some of the most prominent blogs on evolutionary biology (Dawkins, Coyne, Meyers, …) have essentially become self referential and that their main content is coverage of the reactions to other blog posts, if not deliberate attacks. In this context, science becomes secondary, only a smoke screen used try to confer legitimacy to a post, instead of being the central topic of analysis (in contrast, take a look at the phylogenetically community, where blog posts lead to scientific insights).
    In this sense, I don’t think that it’s fruitful to respond directly to Jerry’s post. Rather challenge him to rise up to your level. Invite him to write a response in ETVOL, asking him to adhere to the rules, rather than letting him play on his own field.

  13. newt rality says:

    The group-profile is the sole evolvable entity. There need not even be any between-group selection for altruism to evolve(although, there probably is and it probably helps!). All that is needed for sincere kindness or altruism or blind cooperation is the individual member, as a species of unit, being ultimately dependent in the medium and long-term on the group, for both the individual’s and the group’s survival and the need for the group as a whole to police it’s individual members.. for the sake of the group’s survival and it’s individual members.
    It makes no sense to reduce the notion of the organic multi-scale complexity of of the whole life-cycle to the the herded individuals and their herded genes. The life-cycle is not shaped by Ayn Rand’s and John Nash’s artificial selection.
    The notion of natural selection being able to discriminate between a more versus a less memberistic tendency in a group-dependent species in an intra-species transparent environment(helped by gossip-language, etc.)and with the less attractive, less popular individualism ‘bred for’ irreversible group-failures and sexual competition etc.,is unproblematic.
    Rock on Dave. Best wishes from the UK. Forgive the lack of supporting evidence.. this is mainly a conceptual-positioning grin

  14. Gerstle says:

    “The notion of natural selection being able to discriminate between a more versus a less memberistic tendency in a group-dependent species in an intra-species transparent environment (helped by gossip-language, etc.) and with the less attractive, less popular individualism ‘bred for’ irreversible group-failures and sexual competition etc., is unproblematic” ~ Newt Rality

    …  This sentence is about as impenetrable as any I’ve ever read, though that’s incredibly typical in arguments about units of selection. Your post, like David’s, goes in fifty different directions at once. If there is going to be ‘constructive disagreement,’ there has to be some attention to clarity and focus regarding (1) what you’re arguing and (2) not exhausting your audience. Any scientific journalist would tell you guys that. Not that you’re listening.

  15. David Sloan Wilson says:

    Thanks to everyone for their comments, which partially inspired my newest article titled “Why is everyone still so muddled about selfish genes?” Let’s move the conversation over there—and hit above the belt smile


  16. newt rality says:

    Gerstle, my long-winded sentence, that you quoted, was probably worth three smaller ones but it’s unfortunate that you found it impenetrable. My point is that the selection of a sincerely ‘memberistic’ tendency is unproblematic so long as there is adequate within-group transparency and member-accountability for a species whose members are ultimately mutually-dependent. Selfishness (not the more socially and sexually attractive ‘individuality’) would only then be possible where the group was dysfunctional. No wonder(?) that we thrive where there is transparency and accountability.. and don’t, where there isn’t.

    And, to David, thanks, see you ‘over there’.