David Sloan Wilson loses it again

March 9, 2012 • 6:27 am

For years, biologist David Sloan Wilson of the State University of New York at Binghamton has conducted a futile one-man crusade to convince the world of two things.  First, that group selection (or “trait-group” selection, to use his term) has been hugely important in evolution, especially in the evolution of altruism—and of human religion.  Wilson is also convinced that evolutionary biologists unfairly dismiss the tremendous importance of group selection. His second hobby-horse is his insistence that we can fix society only by incorporating evolutionary principles into social reform, especially (of course) Wilson’s own views on group selection (see my review in the New York Times of his new and bizarre book, The Neighborhood Project, which is about his use of evolution and group selection to fix his own dysfunctional city).

I’ve wavered between the view that Wilson is slightly off-kilter and self-promoting about this, and the view that he’s simply such an ardent believer in his own theories that he’s forcefully trying to argue for a neglected position.  After reading and reviewing his book, I decided that he’s a bit off balance, for in reality there’s simply not much biological evidence for group selection, and none for the evolution of altruism or human religiosity—which I doubt is even coded in our genes—via that process.  Still, Wilson persists.

His latest diatribe, “When Richard Dawkins is not an evolutionist,” has convinced me that Wilson is totally over the waterfall. It’s an attack on Richard Dawkins that appears on the website for which Wilson is the biology editor: “Evolution: This View of Life.”  It’s a strong piece, but also a piece infused with silliness.  Its thesis is simple:  Dawkins fails as an evolutionist in some areas.  First Wilson brings up, gratuitiously, Dawkins’s inability to recall the exact, full title of Darwin’s great book:

In a recent BBC radio interview, Richard Dawkins questioned the religiosity of Brits who consider themselves Christians but can’t name the first book of the New Testament. He was challenged to recall the full title of Darwin’s Origin of Species and failed, even uttering “Oh, God!” as he ransacked his memory.

Does this mean that Dawkins fails to qualify as an evolutionist? Of course not. But Dawkins might fail to qualify for other reasons. . . A person can easily qualify as an evolutionist on topic X but not topic Y. On this basis, I will state the bold hypothesis that Dawkins fails to qualify as an evolutionist on two topics for which he is well known: religion and selfish genes in relation to group selection.

Here are Wilson’s two accusations:

1. In The God Delusion, Dawkins did not discuss the origins and nature of religion as a human construction. Wilson:

In my review of The God Delusion published in Skeptic magazine, I criticized him at length for misrepresenting the nature of religion and ignoring the burgeoning literature on religion as a human construction from an evolutionary perspective. In his reply, Dawkins said that he didn’t need to base his critique on evolution any more than Assyrian woodwind instruments or the burrowing behavior of aardvarks, because he was only addressing question one and not question two. That’s bogus. Dawkins holds forth on question two all the time, and when he does he’s not functioning as an evolutionist–by his own account. Atheists can depart from factual reality in their own way, and so it is for Dawkins on the subject of religion as a human construction.

This is simply stupid.  As you can see from Dawkins’s earlier reply to this criticism, he notes that he devoted an entire chapter of The God Delusion (chapter 5) to this question. The fact that Dawkins isn’t obsessed with that question, which Wilson is, reflects the different purpose of The God Delusion: to show people that there was no evidence for God or the tenets of faith, and hence that we should abandon religion as a baseless superstition.  (Dan Dennett also dealt with the origins of religion in Breaking the Spell.)

Of course Dawkins realizes that religion is a human construction—what else could it be if it’s a delusion?—and he has indeed speculated about its origins.  But why should Dawkins be faulted for concentrating on the pressing problem—the damage caused by faith—rather than on the more arcane and academic question of why faith came to be? (By the way, that’s a question I think will be almost impossible to solve, for the origin of religion is lost in the mists of time. We can, however, study the origin of certain faiths like Mormonism. And I don’t think that belief in God has a genetic basis, or that religion is a group-selected evolutionary adaptation.)

Chastising Dawkins for the failure to be interested in the same things that obsess Wilson is simply another manifestation of Wilson’s hubris—a hubris amply on display in The Neighborhood Project.

2.  Dawkins’s argument against group selection, based on the concept of genes as “selfish replicators” as set forth in The Selfish Gene, is wrong.

Dawkins first achieved fame for his book The Selfish Gene (1976), which portrays genes as “replicators” that typically survive by forming individual-level “vehicles” but can also survive in other ways, such as at the expense of other genes within the same individual or by benefitting copies of themselves in other individuals. 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, but it soon proved to be nothing of the sort. In a genetic group selection model, the altruistic trait has a genetic basis, just like any other trait; the genes merely require a process of between-group selection to evolve when they are selectively neutral or disadvantageous within groups. When this happens, the genes for the altruistic trait are more fit than the genes for the selfish trait, all things considered, and therefore quality as selfish at the genetic level, as Dawkins defines selfish genes. Put another way, an argument against group selection framed in terms of selfish gene theory doesn’t depend upon the status of genes as replicators (which is always the case) but upon whether groups can qualify as vehicles of selection.

This is ancient history for just about everyone except Dawkins. He’s still claiming that the replicator concept counts as an argument against group selection, as if he can do so merely by decree. See these three posts on my Evolution for Everyone blog (I,II,III) for more.

This is sheer madness—an almost incoherent rant that completely misrepresents Dawkins’s views.  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.

That’s the classical argument against group selection (plus the observation that the reproduction of individuals far outstrips that of groups), and it has nothing to do with Wilson’s claim that “Dawkins and others. . .regarded the replicator concept as a drop-dead argument against group selection.”

Does Wilson even understand Dawkins’s argument here? It doesn’t seem so.  Replicators are replicators, and if they are to succeed then they have to increase the fitness of their “vehicle,” because that vehicle carries all the replicators.  It’s much easier to envision an individual as a successful vehicle than a group as a successful vehicle, although even in individuals replicators can subvert the “group” of genes within, too (segregation distortion, in which one form of a gene kills off the other one during the formation of sperm, is one way this can happen).

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.

I’m not sure why Wilson has produced such a misguided tirade, but I suppose it’s because he sees himself as someone crying in the wilderness—that his views have been neglected in favor of those of Dawkins.  And that’s indeed true: Wilson is pretty much seen by evolutionists as a scientific outlier, someone who’s forcefully and unreasonably pushing a theory that lacks evidence. When he comes up with hard biological evidence that group selection has been important in the evolution of altruism, religion, or other traits, then people will start listening to him. In the meantime, his constant harping on group selection, and his persecution complex, are growing tiresome.  When he claims that his second criticism of Dawkins “is ancient history for just about everyone except Dawkins,” he’s claiming support from others for his position, but that is support he doesn’t have.  Who are all the others (presumably everyone but Dawkins) who agree with Wilson? I’m not among them.

Wilson’s efforts, of course, are heavily funded by the Templeton Foundation, where he’s on the Board of Advisors.  It is, of course, typical of the Templeton Foundation that their advisors have received some form of Templeton funding; it’s the way they herd scientists into their posh stable. In fact, I recognize several scientists on the advisory board, and all of them that I know have received either a Templeton Prize or Templeton funding for their work.

As a side note, I recently did a podcast interview for the Evolution: This View of Life site.  Had I known that the biology part of the site was run by Wilson, and is used largely to promote his own views about religion and group selection, I would not have done it.

108 thoughts on “David Sloan Wilson loses it again

    1. Ouch! “What Jerry doesn’t seem to realize is that even the most severe critics of group selection nowadays, who know enough to publish in peer-reviewed journals, accept that group selection occurs.”

      Who then?

      1. When saying that “everyone agrees that group selection occurs” Wilson is actually quoting Andy Gardner. See: Kohn, M. 2008. The needs of the many. Nature 456, p.296

    2. John Maynard Smith would roll over in his grave if he where able to hear Wilson claiming him as an advocate of group selection. Smith spent his whole life proving that group selection was a fallacy.

  1. You tell us how busy you are then spoil us with this nice full analysis, for which thanks. I hope that you do not get too upset and then set really mean questions for that exam!

    1. Seeing as it is only March, I was puzzled about Jerry writing “a final exam”.

      Jerry, did you cram a grad course into a shortened semester? Or does the U of Chicago have a quarter system with an unusually long winter quarter?

  2. Wilson has never been deterred by the necessity for obtaining the same sorts of evidence for group selection as has been obtained routinely for genic or individual selection. He sees group selection everywhere, despite the reality that group formation and differential reproduction simply do not occur at the rates one would need to overwhelm individual selection (show me the data!). That is why group selection almost represents more of a hobby horse than a scientific hypothesis for him. Hence, for Wilson, group selection (in his formulation) has a quasi-religious feel to it, and is understandably and appropriately funded by Templeton! He seems to want to “support” the notion of group selection by fiat.

  3. “Wilson’s efforts, of course, are heavily funded by the Templeton Foundation, where he’s on the Board of Advisors.”

    I don’t think that what DSW has to say should be read through Templeton-flavored glasses, but it’s not something I control very well. Once disclosed that he’s a Templeton advisor, I’m deeply suspicious of anything he has to say about, well, anything. How limiting for me.

  4. Just to stir the pot a bit

    I would like to draw your attention to an article in the New Scientist magazine (1 October 2011, Re-educating superbugs could save lives, by Claire Wilson). The interesting part concerns the grouping together of Pseudomonas aeruginosa as a strategy for launching an all-out attack on our immune system. The thrust of the article concentrates on the cheating bacteria which subsequently develop and subvert the group strategy but none-the-less the development of this group attack in the first instance does demonstrate that such strategies can evolve and be highly successful. It should also be noted that the group attack involves a high degree of self sacrifice within the individual bacteria, 90% mortality to be exact. The fact that cheaters become a major factor in reducing the success rate of the group strategy does not, I would suggest, take away from the fact that the strategy has evolved and demonstrated to be successful.

    1. This is an interesting example, and something like this may have been involved in the evolution of multicellularity (our somatic cells “sacrifice” their futures for the benefit of the germ-line cells). Strassman and her colleagues have used stalk formation in “unicellular” slime molds to explore when we expect cooperation vs. cheating in a system much like this. I may be wrong, but I don’t think cooperative behavior like this (or pack hunting in unrelated carnivores to bring down a large prey) necessarily follows the group selection model proposed by Wilson.

    2. But the individuals in the group would all be closely related. So why would a group attack behavior be evidence for group selection rather than just kin selection.

    3. 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. In this particular example (from what you’ve described) it seems extremely likely that the cooperating bacteria would be highly closely related, if not direct clones of each other. Either scenario would strongly promote altruism on the basis of kin selection.

  5. Re the Selfish Gene, I wonder if Richard was familiar with this quotation:

    Genes build themselves into cells and cells into the gene hive called man in order to develop their potentialities, not man’s. The idea of man’s being able to develop was purely an anthropomorphic concept.

    — Brian W. Aldiss, “Gene Hive” (1958)

    (I suspect I might have mentioned this before… but do so again in the vain hope that Richard might comment in person. 😉 )


  6. Rather sad that so many people attempt to make names for themselves by maligning Richard. But, that seems to be the price of whatever degree of fame one has in an ignorant, tabloid world. The attacks are about the attackers, not Richard. Good that he has a thick skin, though. Or, maybe it’s simply a civility that puts theirs to shame.

  7. I know Professor Dawkins hates it when publishers re-title books, but he might agree that there should be two titles for “The Selfish Gene”

    “The Selfish Gene” for those who are open minded and don’t judge books solely by their covers, and “The Cooperative Gene” for those who’s biases cause them to.

    1. I’ve seen him comment (I don’t remember where, though) to the effect that he wishes he had added an “s” to the end of the title so that so many people wouldn’t have jumped to the conclusion that the book was about a gene for selfishness.

  8. Accepting that the arguments against group theory are correct due to subversion from within, is there a way that religion and culture can circumvent this principle in our advanced culture? Look at Islam, for example, where peremptory death awaits the apostate and paradise promised to the martyr. The population is enslaved as surely as any worker bee or ant and subversion from within is very effectively crushed. Those citizens who do not embrace the religious delusion will not prosper and breed as effectively as the happy copulating conformists. I realise that this aspect of religion would have to be a wrought within the eons of deep evolutionary time for it to be incorporated into our genetic structure but who can tell what path we took to get to where we are?

    1. Well yeah.  In principle it’s possible that a group who adheres to a delusional belief that has the affect of out breeding other cultures, could out compete other groups, and over time other cultural practises within their belief system could have an affect on their genes as well.  “In Principle”.   But the question is, how often do such things happen in nature?

      1. That is a tricky question because if djihadists know that they are going to die, are Pseudomonas aeruginosa aware that they are going to die even if 90% of its army will die when they’ll try to take the control…

        Is altruism a proper word when it come to non-human species, c’est à dire tout le reste except us…

    2. Much of religious behavior is surely not acquired through genetic evolution. You can become a Muslim even when your parents are Christians.

      1. This is perhaps taking the proposition a little too literally. There obviously would not be a gene for Islam and another for Christianity. It may be as loose as a predisposition for believing what one has been told without question. This may simply derive from a tendency for believing that snakes should be avoided without necessarily obtaining firsthand proof, for example.

    3. exotic behaviors occur for the benefit of groups; consider corporations, sports team, military etc.

      This does not make them evolutionary, beyond the possibility that a general concept of cooperation (common in many animals and probably originally kin based) exists. Humans can however see value in complex cooperation without it being an evolutionary design.

  9. I only found “This view of life” a week or 2 ago, and the first thing I read was the dim article which is the subject here. I have not much liked D S Wilson since I tried to read his quite self-satisfied book, and I like him less now. I’ve unbookmarked “This view of life”

  10. I was planning to visit the website, since Jerry is podcasting there. But after this article, I’d rather not. Un-Dawkins are very common, and so far I’ve never enjoyed any.

    True, it is sad that so many people try to be famous by maligning the truly famous ( as with Einstein, Hawkings, Dawkins).

  11. Empirical evidence for group selection seemed to have been observed by a danish team who studied the population of atlantic cod in the St-Lawrence gulf. What is even more surprising is that it would have take only a 3 decades to operate the changes.

    Canadian Journal of Fisheries and Aquatic Sciences, 2009, 66(10): 1719-1733, 10.1139/F09-132

    “Decreasing temporal trends in probabilistic maturation reaction norm (PMRN) midpoints, symptomatic of earlier maturation despite environmentally induced variation in growth, have been observed in many exploited fish stocks. Here, we studied the growth and maturation trends of female and male Icelandic cod (Gadus morhua) by estimating PMRN midpoints for cohorts 1964–1999 and found evidence that a shift towards maturation at smaller sizes and younger ages has occurred independently of changes in growth, condition, and temperature. Weighting the data with regional survey abundance estimates to account for spatial heterogeneity in maturity status and sampling intensity did not qualitatively affect the temporal trends. Length-at-age also decreased through the study period, which, through simulations, could be attributed to the energetic costs of earlier maturity at maturing age groups but not at younger ages. These findings support the hypothesis that such changes in maturation schedules are not caused by environmental factors alone but could also reflect a genetic change, potentially in response to intensive fishing.”


    (click on english at the right top if it appears in french)

    1. This is not evidence for group selection. It is evidence of plain old everyday Darwinian selection. Go check up on the what group selection is. Wikipedia is only a few key clicks away.

    2. Sorry, but unless I’m badly misunderstanding this, there’s NOTHING in this abstract that points to group selection.

      1. I’m very very far form being an expert in Evolution but why the change in question isn’t due to a group selection if a whole population of cod belonging to the same region is able to modify its growth rate within 30 years?

        I’m also surprised to learn in Wiki that “cultural evolution” isn’t something that is widely accepted? Evolution officially can only work on a biological level?

        1. Because the largest and fastest growing individuals are those that are selected out of the population by commercial fishing. Plus evolution always happens at the population level.

          1. Oh, and those individuals that mature more quickly and start reproducing at a younger age are more likely to leave behind offspring before they are harvested.

            1. If it is obvious that those who are selected by the commercial fishing have less chances to survive, how could a specie change in only 30 years its age of maturation?

              If it wasn’t the norm to mature at the younger age they now mature, how could a few individuals that would be able to do it could transmit their “ability” to an entire population within 30 years?

              “…midpoints for cohorts 1964–1999 and found evidence that a shift towards maturation at smaller sizes and younger ages has occurred independently of changes in growth, condition, and temperature.”

              “These findings support the hypothesis that such changes in maturation schedules are not caused by environmental factors alone but could also reflect a genetic change, potentially in response to intensive fishing.”

              1. JF: Please invest in a basic evolution textbook. This is not advanced material.

              2. The basic I know about evolution is that it happens on a long run, not on a 30 years run…

                I also read that it might happen suddenly, because of some sudden changes in the environment. But the idea that an organism can develop antidote in reaction to the sudden changes seems to be very suspicious among the darwinian interpretation. From what I read, it has to be latent accidentally in the organism, it cannot be an organized response to what is felt as a threat… Yes , no?

              3. If I read the abstract right, these people haven’t even shown rapid evolution, much less group selection. They only *speculate* about a genetic basis for the changes.

                Without that, we are only talking about phenotypic plasticity, possibly as a reaction to reduced competition by larger cods, changes in resource availability or the ever-popular climate changes.

              4. They indicate that the changes of temperature and that the size of cods is not a factor in the younger maturation of the species…

        2. Darwinian evolution is an extremely robust theory of biological changes over time based on replication and selection of genetic material. Cultural evolution is a much less robust and fuzzy theory of change in culture over time. They are completely separate.

          Group selection is a concept within biological evolution. It is not well accepted as an significant contributor to biological evolution.

          Please note: the word “evolution” does not require capitalization unless it appears at the beginning of a sentence. Like “gravity”, it is not the name of a church or faith group.

          1. So according to darwinian evolution, the homo sapiens didn’t really evolve since he appeared, despite the radical changes that he was able to create?

            1. This is a trolling question. It expresses a level of ignorance so profound that a serious response is not required. If you were really interested in the subject you would go read basic text on the subject. In fact, you might consider the book that this web site is named after.

              1. JF: I did not say you were not worthy, I said the question was a trolling question and not worthy of response. It remains unworthy of a response. You are not interested enough to learn the very basics. And I am not interested in playing a juvenile troll game.

              2. GB, I think I understand evolution more as it is understood by most of the common mass. What is now confusing me is the more I read about evolution, the more I see different interpretations on how how it happened. You’ll understand that it is not just because the host of this blog doesn’t think that selection group is not something relevant that I’ll buy it.

                When you read about group selection, you can see that it is an avenue that is highly considered and that the theory of evolution is a road that is made from several little avenues (excuse my french).

                So my question maybe sound naive but it is not a trap, I’d really like to know if accordingly to the theory of evolution, I am more evolved or not than the first homo sapiens who appeared, even if he couldn’t sing, dance and read like I do?

              3. So Ant, if I understand well the link you paste, accordingly to Dawkins and the theory of evolution, there is no significant difference between the first homo sapiens and me.

                I really just want to understand. Is it possible to have a yes or no? If not, explain why please.

                If it is obvious, pardon my ignorance (and my french).

              4. JF: Your understanding may be exactly the same as the average among the common mass. That only puts you in company of a large group of ignorant people. It remains impossible to seriously discuss the subject because evolution, like gravity, are not matters that are determined by vote.

                Your question remains, as before, a trolling question. I strongly suggest you purchase a decent book on evolution and learn a bit. Our host’s book is an excellent one, not technically burdensome, and easily accessible to the lay reader. There are many others that you might choose from. The Ancestors’ Tale by Richard Dawkins is also an excellent read. Go pick up one of these books. Read it. Not only will it demonstrate that you are taking your own questions seriously (which I currently doubt) but it will also provide you with material for some really interesting questions that are worth the effort to articulate.

              5. First GB, I don’t question the truth of evolution. Secondly, I don’t really have the time to read books about evolution. Probably because it is not a passion for me. So, yes, in a sense, you can say I’m not serious enough to learn about it.

                But on the other end, thanks to evolution(?), I am just a few clicks and blogs away when I have questions about evolution. That is how a troll evolutionist (?) gave me an interesting trolling answer to my trolling question about that fuzzy cultural evolution thing:

                “There is no doubt that it is often important to remind overly-enthusiastic orthodox Darwinians of the importance of culture. For example, it seems that the increased incidence of lactose-tolerance among human populations has arisen as a consequence of a cultural innovation—namely dairy farming. The relatively recent appearance of this genetically-controlled adaptation demonstrates that human physiological nature is something that continues to change, and it also demonstrates the causal impact of culture on genes (Richerson and Boyd 2005, 191–92). Such examples by themselves show the rashness of any view that claims either that human nature has remained fixed since the Stone Age, or that genes are somehow in the evolutionary driving seat. Yet none of this shows that we can develop a general, informative theory of cultural evolution.”


              6. You have plenty of time. Just take a pause from posting inanities here. Bingo… time for reading.

                What you are seem to be lacking is honest willingness to learn. That’s the signature of a troll.

            2. POI: Unlike “evolution”, “Darwinian” is capitalised. As are generic names (but not specific names, an error the popular press is uncommonly fond of); and binomials are usually italicised; thus, Homo sapiens.

              You’re welcome.


              PS. Btw, “specie” (your comment above) means money in the form of coins; the singular of “species” is “species”. Almost the inverse of French; “en espèces” & “espèce”. English, he is so funny!

    3. This is not group selection. The genes in organisms that produce large size and slow maturity in a population are too costly to be maintained in an organism under population stress. Reproductive fitness will be greater in those organism whose genes produce smaller size which allows them to mature and reproduce quicker. The reduced nutritional requirements of the smaller size can also bring a fitness advantage as well.

      1. But that is always the case. Improvements are always welcome, no?

        The data tells us that even if the size of the cod doesn’t change, the maturation changes.

        Again, how can a species can adapt its sexual maturation within 30 years because of an external pressure accordingly to a darwinism that would exclude group selection?

        1. Cod that mature sexually at a younger age start getting their chances to reproduce earlier. Having a chance to reproduce earlier means more chances to reproduce. More chances to reproduce means more likely offspring. More likely offspring – themselves more likely to reproduce earlier – will mean more cod reproducing earlier.

          There’s that pressure among the cod each generation. _Every_ cod runs the risks of getting killed before reproducing, or reproducing much, and in this case, that pressure’s enough to offset the contrary pressures, perhaps to grow larger and have a better chance to keep reproducing over a longer period.

          Evolution by natural selection just takes selective pressures and variation among heritable traits that are relevant to them. Give it one generation and there’s a little evolution. Up the pressure, get more evolution. Give it more generations under that pressure, get more evolution. Start with more heritable variation, get evolution that can go further quickly.

          It takes longer when it’s a complex adaptation, when the pressure is mild or erratic, or when generations are long. None of those apply here.

          Even if they did, it’s not the sort of adaptation that’s bad for individual’s reproductive chances but great for the group. Each cod’s on its own here, with genes or without them for becoming an ancestor in its environment.

  12. It’s true, the Templeton Foundation is “oxymoronish” and supports contradictory viewpoints. Nevertheless, it is a lighthouse that warns us of nearby danger while at the same time blinds us like approaching headlights in our eyes. I am always skeptical of everything with a Templeton label.

  13. Dawkins’s earlier reply to Wilson’s criticism contains this wonderful zinger of a phrase, worth requoting:

    As for group selection (either as normally understood or in the idiosyncratic sense of Wilson’s private re-definition, about which he has been obsessing for thirty years)…

    That man can write!

  14. 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.

    That’s what altruism is, selection based on reciprocity, both socially and for survival of the individual. Groups are not as vulnerable as you guys say, because obviously, we wouldn’t have societies based on compassion and empathy that co-operation and trust is based on, the recognizing it in others.
    Just read a study that people find altruism and compassion more attractive, all things being equal.
    Altruism only has meaning on a group level. If it was so vulnerable to ruthlessness, then we wouldn’t have it, or complex societies. Sheesh, even bonobos exhibit altruist behavior, and it is most strongly manifest in high level individuals.

    Individuals achieve status by being altruistic, and they attract mates. Pretty strong arguments that seem to trip people up, but this is based on a faulty assumption, that at the group level, ruthlessness is more powerful. No it isn’t, ruthlessness is a threat to everyone’s individual and genetic survival.

    1. You are mistaking “selfish genes” with “genes for selfishness”.

      For instance, “Altruism only has meaning on a group level. If it was so vulnerable to ruthlessness, then we wouldn’t have it, or complex societies.” How do you know it’s not the selfish genes that are promoting altruism?

      1. Yes, it is the gene promoting, or coding for, altruism, and it only flourishes when individual fitness and chance for reproduction is increased. That is my point, that it takes groups for this to happen, and since a co-operative group increases the chance for each individual to reproduce, the altruism gene, or replicator, is promoting its own success.
        There is absolutely no doubt that every adaptation can be seen as a purely selfish ‘success’ of a gene, that is never in dispute.

        The group security is the selecting factor. Individual success is the selecting factor.

        I am saying that co-operation and altruism enhances both group and individual fitness, whereas individuals acting without concern for group success and only their own, are seen as poisonous, a threat, to the group, and excluded

        Nowadays, we put people in jail. This isn’t the best answer much of the time, but the overall intent is the protection of society.

        When ruthless individuals succeed, they succeed by tricking the group into thinking the rogue is actually not trying to undermine other members of the group, or the group as a whole.

        But, yes, it is easier to test empirically when , or formalize, as gene competition, but the obvious fact of the matter is that selection is done at the group level. I find it hard to believe anyone would deny this. Evolution works by adapting to the environment, and groups have been around long before humans have as an environmental condition, but it hasn’t been possible to form large groups, and larger groups with shared purposes without the altruism replicator, the altruism replicator allowing groups to form.
        Wait, what did I just say!

  15. I find it irritating that DS Wilson seems to take group selection (the kind of multilevel-selection he happens to call thus) as necessarily benign. At least that was the impression I got from a talk of his available as an internet video. He asked the audience what they associate with group selection and individual selection and got all positive associations for group and negative ones for individual selection.

    Not even Wynne-Edwards, whose concept was wrong (my opinion) or naive (Wilson’s wording), thought of it like that. On the contrary Wynne-Edwards thought that group selection could ‘sentence’ individuals to death in the midst of plenty. Oh, and Hamilton by the way thought of group selection as mainly bad (war, genocide and all that).

    This false good vs. bad dilemma seems to bring all the ideological dispute into the issue, which would have otherwise probably long been settled.

  16. I’ve never found DSW’s points about the group-adaptive value of religion to be an effective critique of the “Gnu Atheists.”
    Even if one concede’s to DSW that religion is in some way the product of group-level selective processes I don’t see how you have reason to dislike religion any less than before.

    A group-selection view of religion at most suggests a change in phrasing from

    “We should liberate individuals from selfish memes”
    To “We should liberate individuals from tyrannical group-selfish adaptations”

    If religions are the product of a product of group selection, they are adaptive for competing groups or societies, but can be every bit as deleterious from the point of view of the individual as a selfish meme.

    Competition between societies is clearly important in human history and even the human present day. But isn’t competition between groups of the sort that some flourish at the expense of others something worth actively discouraging in this era of suicide bombings and nuclear weaponry?

    Yet, there is no natural (evolutionary) process for which seeing all of humanity as a community is adaptive. There is no Darwinian process that will get us there. Individual selection won’t. Selfish gene-ism won’t. Selfish meme-ism won’t. And DSW’s hobby horse group-selection won’t (since it depends competition between groups). Dawkins is right when he says that morality is about opposing Darwinian processes, whether these are at the individual or higher level of selection.

  17. This thread, the other recent one here, and the one over at PZ’s have convinced me once and for all that the comment sections of the blogosphere (including the websiteosphere) are exactly the wrong place to try discussing the group-selection issue. There are simply too many misunderstandings and misconstruals in the way (I readily ackowledge that some of them might be mine).

    1. I’d welcome your comments over at mousetrap. There are tidbits on what Darwin, Wynne-Edwards, Wright, and others said that is relevant to the history of that controversy. I’m trying to develop a differentiated picture of the issue and to steer clear of the purely semantic misunderstandings.

    2. But wouldn’t be an evolutionary goal to make the misunderstandings more understandable to dumb people like me, or like you, if you agree that some of them might be yours?

      In other words, is evolution strictly a biological process?

      I’d really like to know the answer…

      1. @JF Fortier
        1st question: As I understand it (onw misunderstandings possible), one party uses the term group-selection as “That what has been rejected in the late 1960s” (a scenario in which cheats can indeed undermine the system). The other party uses the same term for multilevel selection applied to a particular tier of the natural hierarchy (a scenario in which cheats cannot undermine the system, because of particular dynamics of group establishment and resolution that lead to a statistical swamping of within group benefits by between group differences). As long as the semantics are that sclerotic, this sort of debate will not go away, and that’s why most evolutionary biologists now avoid the term group-selection and speak of multilevel selection instead.

        2nd question: As long as we are not talking about cultural evolution, it is a strictly biological process.

        1. Mr. Fortier, please teach yourself some evolutionary biology before you come over hear and put more than a dozen comments on this thread.

          1. I can’t exactly remember how I got here but it had to do with one of your colleague who was using morality as an argument against theist evolution. I found that it was a weak argument and explained why, and a lot of comments were made around the answers. I got enthusiast about your blog because you can’t usually have these kind of civil debates on these topics.

            But I now realize where I am (I didn’t who you were) and that I am too novice to really follow what is going on when it comes to strict biological evolution. So I’ll temper my enthusiasm and learn more about it, which I did since the last days btw.

            So thank you for that and sorry for the overspreading.

            But I must say to my surprise that I didn’t expect cultural evolution to be a controversial issue…

            Bon dimanche,


  18. 1. Please use your Google+ account, Jerry! I know that’s kind of a long shot, but I figured I’d try.

    2. “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.”

    I think as an addendum you’d have to add if the altruistic tendencies of a gene increased mating success via sexual selection. 🙂

  19. [oops, I accidentally posted this on one of the old related blog posts. please ignore/delete the other one, as suits you]


    I have followed this selfish-gene vs. group selection debate for years, and in blog comments and forums have always defended the selfish-gene position. However, since the 2010 Nowak et al Nature paper, something has nagged at me — that buried in their opaque mathematics in the appendix of that paper and their poor choice to frame their ideas in terms of “group selection”, they have a valid point. It may be a point well known and understood to biologists, so the debate is about interpretation and terminology… or it may be a somewhat novel point (although from a mathematical perspective it should be almost as obvious as Hamilton’s rule).

    Here is what I *think* he might be getting at: suppose you have a trait with reduces the reproductive success of a individual *relative* to the “group”, but which enhances the overall reproductive success of the group as a whole more than enough to offset the personal “sacrifice”. The “group” need not be related in any way and might be defined merely by proximity (local neighborhood). Such a trait would certainly be favored by natural selection, simply by considering the net effect (to increase reproductive success of the carrier).

    To clarify with a concrete example: suppose I have a behavioral trait which gives my immediate neighbors, or my “group” a 0.1% reproductive advantage over me, but also gives my group (including me) a 0.2% advantage over neighboring groups. I still get a 0.1% net advantage as compared to the global population. My progeny who carry the trait become less frequent among the local population but more frequent globally. This trend will continue towards some evolutionarily stable distribution with the global population.

    Just to further show how this kind of trait can lead to apparently altruistic behavior, consider that the .1% or .2% advantages are statical averages. And further suppose that the .1% “sacrifice” is virtually guaranteed with only a little variation, but suppose that the .2% payoff for the “group” is the result of a 100% payoff which only happens .002% of the time. Such a behavior would easily be seen as “self sacrificing” as the connection to the payoff would not be obvious.

    A trait like this is encompassed by selfish-gene theory which ultimately only cares about the net effect. But such a trait would result in behaviors which appear altruistic with respect to the local “group”. A trait like this would actually be a special case of “win-win” cooperative behavior — might call it “win-WIN” cooperation to emphasize the lop-sidedness of who benefits.

    If this is the what Nowak, Wilson, et al, are trying to get at, they are not doing a particularly good job of explaining their position. It certainly does not support “group selection” in the sense of groups as units of selection. (It does not have much if anything to do with Wilson social/political view either as far as I can tell)

    However, in discussions of the evolution of altruism (or seeming altruism) I have read (such as here, in Dawkins’ books, other blogs, forums, etc.), this kind of asymmetric, indirect “win-WIN” (maybe I should call it “lose-WIN”) effect never seems to come up.

    1. @s.k.graham

      I’m not an expert so keep that in mind.
      I think you are describing everything correctly. Those who are in favor of group selection/multi-level selection (MLS2) agree that traits are determined at the genetic level. DSW is also not opposed to looking at the gene level to do the accounting. The question is where is the selecting happening? I believe his position is that since the trait would be net deleterious if there weren’t between group competition, the selecting is happening at the group level.

      1. This suggests that the whole controversy is one of definition of terms and nothing of substance. In which case I think Wilson & Nowak and their allies are fabricating controversy by changing definitions. “Group selection”, as I understand it, implies that groups are identifiable units which reproduce more or less as units. But what I have outlined does not require delineated “groups” at all — I merely need locally varying gene frequencies.

        Even with identifiable “groups”, the success of the trait depends on individual success, and the fact that non-carriers out-compete carriers within each groups has a major effect on how the trait spreads through the population as a whole, and that is certainly a gene or individual level selection effect.

        1. Let me recommend this talk by Stuart West

          I did get something wrong though. DSW is defending multi level selection 1 (MLS1) not MLS2.

          I don’t think it is just semantics that is being argued. Jerry’s post is about why he thinks group selection is wrong and it isn’t just for semantic reasons. He’s explicitly accepting the concept of multi level selection when he says things like, “Yes, altruism may help groups propagate” and “even in individuals replicators can subvert the “group” of genes within”. The latter refers to genes that “trick” the cell machinery into replicating them even though they weren’t the gene that was selected at the individual (phenotype) level.

          The question is about whether or not selection at the group level can persist and be important in evolution as a whole. In other words, it is an empirical question.

          It may be helpful to go back and re-read Jerry’s post with this in mind.

          PS If I’ve got it all wrong I hope someone more knowledgeable will correct me.

          1. “I did get something wrong though. DSW is defending multi level selection 1 (MLS1) not MLS2.”

            I once tried to pin down DS Wilson on the MLS1 MLS2 distinction at his own blog (Evolution for Everyone) and got away with the impression that he does not think it is a very important distinction to draw in the first place. Maybe that’s why he thinks Wynne-Edwards needs to be rehabilitated. Only if you do draw the distinction, you can see that Wynne-Edwards tried to apply an MLS2 scenarion where actually an MLS1 would have been needed.

    2. “My progeny who carry the trait become less frequent among the local population but more frequent globally. This trend will continue towards some evolutionarily stable distribution with the global population.”

      Yes the trend will continue to the stable distribution of zero. The trait becoming less and less frequent in the local population means it will eventually disappear. Why do you think it would stabilize before this, if your local competitors always outcompete you? Your group may do so well even to the point of eliminating all the other groups, but how would that help you?

      1. “The trait becoming less and less frequent in the local population means it will eventually disappear.”

        Do the math. If the local population grows by, say 2% (a factor of 1.02) and my progeny grow by a *relative* factor of .99, then my progeny grow by an absolute factor of 1.01. Once the trait is widespread, then there are frequency dependent effects that depend on local variations in the frequency of the trait. If all neighboring local sub-populations have equal numbers of the trait, then the local or “group” advantage is no longer conferred and the trait will locally diminish. But “groups” or sub-populations near the geographic border of the expansion of the trait will still have an advantage over the groups further out where the trait does not yet exist. Furthermore, due to unrelated, effectively random effects, you will never have perfectly equal local frequencies of the trait from one group or local sub-population to the next, and the groups or local sub-populations with more members carrying the trait will expand (thus benefiting the local altruists, though not as much as the local non-altruists benefit at any given time).

        Once established, in order for the trait to disappear completely would require almost perfectly uniform geographic distribution of the trait, so that no local population could gain any advantage — and the distribution would have to remain uniform until the trait was gone completely. Seems highly unlikely that you would not get clumps here and there of higher concentrations of the trait than in other places. Think of it this way: if the trait ever did reach the point of almost disappearing, you are going to have one or a few local pockets of the trait, and those pockets will then have an advantage and begin expanding again (in absolute numbers).

        Whether this results in an evolutionarily stable solution, per se, or some kind of oscillating solution, or even chaotic oscillations, will depend on the exact local-variation-of-frequency-dependent effects of the trait.

        But traits along the lines of what I have described could certainly maintain themselves within a population.

        1. Very interesting – what you are describing does sound reasonable. But can this work if the local population is not composed of close relatives? That is, does the mechanism require that two members of the local population are genetically closer (on average) than locals are to members of the distant population? I’m thinking about how what you describe would get started. Let’s say I have a mutation of the sort you described, producing a trait that is a slight disadvantage to me relative to other locals but would confer a group advantage to the local population IF SOME PROPORTION OF LOCALS HAVE THE SAME TRAIT. I’m stumbling over that IF. If local versus distant populations are not differentiated genetically, then the fact that I have the trait doesn’t mean that other members of my local population will. I am no more likely to be a part of an advantaged group than not, if MY having the trait is not somehow predictive of the rest of my local population having the trait. So it seems that all I would have is my local disadvantage selecting against the trait. If, on the other hand, I am genetically closer to members of the local group than members of the distant group, then it seems like we are back in kin selection territory. Am I missing something here? I guess you would not need the genetic differential if one lone individual with the new trait could benefit the entire group by themselves. That seems implausible, though.

          1. [in loose and simple terms] The benefit to the local group would be in terms of some competitive advantage over neighboring groups, and would depend on how many “altruists” the local group had in comparison to the neighboring groups.

            The reproductive advantage (for the selfish gene) would of course be further enhanced if their is a kin relation, so the trait-carrier is that much more likely to be benefiting others who carry the same trait.[**]

            Whether a group advantage could be conferred by a single individual within the group just depends on the nature of the “altruistic” behavior trait. Suppose I spend more time building a defensive structure (a wall or something) or “on lookout” against invasion by neighboring groups. Both I and my group benefit, but my “selfish” group-mates benefit more than I do. Of course there can be many such behaviors where the group advantage will fail to compensate for my personal disadvantage. But there are also plenty of behaviors where the benefit to me will more than offset. As humans, with foresight, we can easily identify such situations, but there is no reason why the “blind watchmaker” won’t stumble onto them from time to time.

            Bear in mind also that like any mutation, advantage/disadvantages conferred can be quite tiny, even imperceptible, but will nonetheless be magnified exponentially over hundreds/thousands/millions of generations. It would only be the accumulation of many such tiny changes in behavior that that you arrive at behaviors that appear noticeably altruistic.

            [**] I should add that the extent to which Wilson & Nowak & others want to downplay the importance of kin-selection, I think they are pretty much dead wrong.

  20. As a non-scientist and merely a science geek I try to hear as many points of view as possible but I try to not be to judgmental. After all with my limited knowledge of science my opinion should be taken with skepticism. But the one observation that I notice from Wilson is that he makes extraordinary claims yet does not provide enough evidence to even satisfy me. I notice with other scientist such as Dawkins and Coyne they do not make claims unless they can present us with supporting evidence.

  21. Originally posted by Jerry Coyne:
    “(By the way, that’s a question I think will be almost impossible to solve, for the origin of religion is lost in the mists of time. We can, however, study the origin of certain faiths like Mormonism. And I don’t think that belief in God has a genetic basis, or that religion is a group-selected evolutionary adaptation.)”

    Hm… I’m not so sure. I’m studying to be a Cultural Anthropologist, and one of my three main interests is the origins and perniciousness of Ideological Fanaticism.

    Part of that is attempting a robust hypothesis of the origins of religion (while granting that a fully realized theory may not be possible).

    My idea (not yet really “tested”, as far as an anthropological idea can be tested, so I’m not willing to call it a hypothesis, yet… and I admit that it’s not really an original idea) is that religion started with the soul, and that idea started with the recognition of death.

    At some point in our evolutionary history, we finally became cognizant enough to recognize when somebody died. This produced two reactions: grief at the loss of someone we had learned to love, and fear that it will one day happen to us (this didn’t all happen at once, of course… we had to become cognizant of the idea of attachment [what we now call “love”] and fear, first).

    So how did we deal with this? Over time, we came up with the idea of the soul and an afterlife… “death” wasn’t a “goodbye forever” thing. It was more of a “see you later” thing.

    At the same time, we were becoming more and more aware of our natural surroundings, so much so that we could think about them. Of course we attempted to explain the natural world and all the natural happenings, from the existence of life to earthquakes and volcanoes and tornadoes and other natural disasters.

    We already had souls and the afterlife, so, taking into account our early attempts to understand the natural world, it’s really just a small step to deities and, finally, religion. And all you need is a basic understanding of memes to see how it might have spreaded so quickly.

    In this sense, I actually agree with Sam Harris when he calls religion “failed science”. If you see science, as I do, as the tool we use to answer questions about the nature of reality, then religion absolutely is a failed science, at least if my idea (please note that I’m not even willing to call it a hypothesis, yet) is right.

    1. It’s a plausible enough idea for an origin of religious thinking. (You may want to take a look at some of the offerings from Boyer, Atran, and Dennett on this score too.) The problems come when you try to penetrate those mists of time to seek to confirm the hypothesis or disconfirm rivals.

      What predictions would your “souls-first” hypothesis make that would differ from those of, say, a hyperactive agency detection device hypothesis, in which early people attribute agency to natural phenomena across the board? What would disconfirm it? What trace of those early days remains now to tell the difference? Religions like Mormonism come up in a vastly different cultural milieu than that of, say, hunter-gatherer bands developing sentences for communication 200,000 years ago? (Yeah, that’s a lot of wild guesses so much as describing them. There’s the point in that.) Even the development of ideological fanaticism has to be much more recent.

      It’s very interesting – it’s hopelessly interesting, and neither you nor I will likely quit speculating. But it’s an answer for which we’re likely to have to settle for vague and uncertain answers.

      1. Unfortunately, I have no way to respond to any of your questions, which is why I’m not even willing to consider it a hypothesis. It’s really just a reasoning along the lines “it makes sense”. I think perhaps the only effective way of really studying it would be to study elephant who appear to mourn for their dead and, of course, our closest evolutionary cousin the Chimpanzee, and see what happens.

        It seems to me that death would be the first natural event we would become aware of, because it’s such a pervasive event (it literally happens all the time and one never needs to go very far to find it, even in the animal kingdom) and it’s such a *personal* event. With perhaps a few very rare exceptions, *somebody* will be impacted by death in some capacity.

        I can see those primitive humans inventing a primitive version of the soul and the afterlife (it would start off extremely simple, I imagine) to help deal with the recognition of death. If death is the first natural event we become aware of (and by “aware of”, I obviously mean we gain the ability to start thinking about it… we have become evolved enough that it actually affects us in some long-term capacity), then I would think that soul would be the first real supernatural invention.

        As for Ideological fanaticism, I actually don’t think it started with religion. I think it started with tribal loyalty and perhaps, by extension, xenophobia. I’ve watched video of baboon societies and sometimes, within the internal fighting and the close-knit groups, I see what I think could tentatively be classified as a sort of primitive fanaticism in the form of tribal loyalty. I see it with chimpanzees, too. Of course, I’d have to do some long-term field observations of both groups, as watching videos doesn’t tell me much of anything. Hopefully I will get the chance to do that…

        1. Well, saying that “things make sense” is not, as you realize, any way understand what really happened.

          I take an apophatic approach to the origin of religion in human society: if I don’t know anything about it, I shut up about it.

          1. Of course. I never attempted to claim otherwise.

            And I’m not claiming any kind of studied knowledge on the subject, as I’ve only *just* started on my Cultural Anthropology degree, and the origins of religion/fanaticism is not where I’m starting.

            I want to start off with an ethnographic documentary about atheism in the United States. I’m in my Junior year of my Bachelor’s now, so I’m starting with a basic study. My Master’s Thesis will be the written version, and I hope to do actually make the film as my PhD project.

            Then I want to do a study on a phenomenon within music culture: P2P, torrenting, bootlegs (unofficial live recordings, studio outtakes/sessions, etc); the idea that music should be free (I know plenty of people who, politically, are staunch *conservative* Libertarians… but when you ask them about music, they become the biggest socialists you will ever meet). Why are they popular? Why are they so wide-spread? Can they be used by the music industry as money-making tools?

            After that is when I want to study ideological fanaticism, attempt to come up with a hypothesis about its roots, and try to understand it’s perniciousness… if there’s a biological evolutionary reason for fanaticism, or if it’s strictly a cultural phenomenon.

            Now I may not do these three things in that order, but all three are my interests and are why I’m studying to be an anthropologist in the first place.

            1. Clarification:

              I’m in my junior year of college as a transfer student to Florida Atlantic University (now in my second semester). I started on my anthropology degree in my junior year because, unfortunately, it took me that long to realize that this is what I wanted to do.

              1. Good luck to you, Nathan! You certainly have no lack of enthusiasm, ideas, or goals!

                Regarding primate research–have you read any Robert Sapolsky?

  22. I think it’s just name-dropping in an effort to draw attention to the writer. Memorably lampooned by Spike Milligan in his war memoirs, the first book of which was titled Adolf Hitler: My part in his Downfall.

    It’s a bit sad when pseudo-atheists or pseudo-evolutionists join the Goddists in misappropriating Richard Dawkins’ name, though.

  23. @ s.k.graham

    Thanks for the clarification. To make sure I understand, is this an appropriate way to re-phrase what you are saying?: A trait that is beneficial for (that is, increases the reproductive success of) an individual might be *even more* beneficial for local con-specifics that do NOT have the trait. So say the trait is sleeping less when night-time ambush is a serious survival threat. If I get the trait I get the benefit of less chance of ambush and the cost of losing sleep, but the former might outweigh the latter. Others in my group get the benefit without the cost, so it’s more of a boon to them than to me. The trait benefits the organisms that hold it, so there is a standard selection pressure promoting the spread/maintenance of the trait. The fact that it is even better for other members of the local group than the individuals that have it is why we can say it is a RELATIVE disadvantage in the local context, but natural selection doesn’t stop working just because there are free loaders. If that is the argument, I’m on board.

    1. That’s a pretty good example of the kind of trait I am talking about.

      It is just a particular case of win-win cooperation, that happens to be asymmetric.

  24. Since when is kin not a group?
    Since when did any mutation not get ‘kin selection’ to begin with?
    Since when did altruism get designated to particular gene sequence, where you either have it, or don’t?

    Since when did an individual have to have any benefit to a mutation that improved its fitness? What about old age? You are not more fit when you are ancient, yet the group protects you, for obvious reasons.
    In fact, intelligence substitutes for physical fitness, why wouldn’t altruism substitute for usury?

    We don’t even know where altruism cam from, it probably is coincidental to self awareness, but I can see it existing before that.

    Posted March 9, 2012 at 5:04 pm

    As a non-scientist and merely a science geek I try to hear as many points of view as possible but I try to not be to judgmental. After all with my limited knowledge of science my opinion should be taken with skepticism.

    No, as a blogger, poster, expert, naive knave, commenter – all views should be taken with scepticism.
    Your view, you should be taken as anecdotal evidence based on an unknown methodology that may involve all sorts of selection bias.
    Personally, I try to evaluate based on logic, but any viewpoint that assumes knowledge of motivation and/or employs ad hominum and pronouncement of truth is immediately suspect, and reason to be very sceptical.

  25. I believe that you are being unfair to Professor Wilson. I have read some of is books and listened to several of his online lectures, and I believe that he is largely correct in his perspective on Evolution. His views on “Group Selection” cannot be summarily dismissed.

    Wilson’s views may not be completely correct, but they are certainly not completely wrong.

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