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A question about group selection

January 29th, 2016

I’m doing some work on evolutionary models of game theory and need to understand the debate about group selection. It seems pretty clear that the great majority of evolutionary biologists reject the idea of group selection, but I haven’t found an adequate (to me) explanation of why they do so. A crucial problem for me is that the literature seems, without exception as far as I can see, to conflate group selection with co-operation and altruism. But the problem of group selection arises in non-cooperative settings, provided they are not zero-sum.

To illustrate the problem I’m struggling with, suppose that two previously isolated species meet as a result of some change. In one species (peacocks), competition between males for mates takes the form of elaborate, and energetically costly, displays. In the other species (penguins) males compete by providing food to their mates. In all other respects (diet, predators and so on) the two are similar. It seems obvious to me that the penguins, with their more efficient social arrangements, are going to outbreed the peacocks and eventually drive them to extinction.

It seems to me there are only two possibilities here
(a) My reasoning is wrong, and we can’t judge which species, if either, will dominate; or
(b) Even though it involves one group being selected over another, this isn’t what is meant by group selection

I’d really appreciate some help on this. I’m happy to have thoughts from anyone, but I’d most like to hear from actual experts with contact details.

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  1. J-D
    January 29th, 2016 at 16:40 | #1

    I am not an expert, but I do have a couple of thoughts for you, one of which is that the example you have offered is not adequately elaborated.

    Why is it that males of the peacock species compete in the way they do? Why is it that males of the penguin species compete in the way they do? In what sense does the behaviour of feeding mates constitute ‘competition’ (for example, do males of the penguin species offer food to the mates of other males to induce mating)? Perhaps most importantly, how does an increase in the number of the penguin species have any effect on the survival of the peacock species?

    Another thought is that instead of worrying about what other people mean by ‘group selection’, you should try to elaborate in more detail what it is that you mean by the term. Sometimes when it seems that a term is problematic it can be helpful to try and restate the issue without using the term.

  2. John Street
    January 29th, 2016 at 16:42 | #2

    (Nothing like an expert) I have always thought of group selection as being exemplified by the hypothetical Man on the Titanic who, by allowing someone else to get in a lifeboat before him, improves the likelihood that his species will survive (although he will by his action have no chance of passing on his own genes).

  3. paul walter
    January 29th, 2016 at 16:53 | #3

    I can’t offer much, beyond asking whether we are clear on what constitues a species “need” as defined merely in the satisfaction of physical needs. Is “altruism” seperate from “group selection”. or a refined component that aims for survival based on the evidence of survival and as an extension, has then the capacity to seek and pursue cornucopia. Is cognition the game-changer?

    Yet instinct seems also to complicate cognition, is not a happy bedfellow, as the instinct for safety conflicts with what appears to be rational responses to developing situations. The social and cultural mechanisms that have allowed us to develop altruism as some thing basic to ensuring “vison”within a group contribute to the downfall of altruism and reversion to survival mode when fight or flight demands the destruction of icons for the survival of certain folk at the expense of others, then there is conflict. It is an imperfect, evolving system and priorities as to a group or groups with competing interests can conflict.

  4. GrueBleen
    January 29th, 2016 at 16:57 | #4

    Just a small question, ProfQ: how could penguins drive peacocks to extinction when they live in completely different environments and eat entirely different food ? Sorry, just couldn’t resist.

    But surely a key question of ‘group selection’ is: how can there be ‘group selection’ unless there is a group ? I mean a pride of lions might be considered a ‘group’, but what about the solitary tigers, leopards, cheetahs etc ? How would ‘group selection’ work with them ? The lions might practice ‘group selection’ in a fashion by a successful alpha challenger killing all the loser’s offspring so he can substitute his own – assuming that he won because of some level of ‘natural superiority’, then he’ll start to pass that along to the ‘group almost immediately.

    Besides, ‘natural selection’ isn’t the only way that ‘groups’ can acquire different characteristics: there is also ‘random gene drift’. How does that figure in your thinking ?

  5. paul walter
    January 29th, 2016 at 17:00 | #5

    As for the rest, JD makes an important point that briefly crossed my mind: that Peacocks mating ritual doesn’t deal with the problem of provisoning for the young, something covered by other means since there would be no Peacocks other wise. The Penguins appear to have incorporated provsioning into their mating rituals, by contrast and the various species thrive to this day also.

    Now, back to watching the rain. It hasnt rained for months here and it is such a thing of beauty as to defy the ignoring of its presence.

  6. Lachie A’Vard
    January 29th, 2016 at 17:16 | #6


    E. O. Wilson is probably your man. He gave an excellent lecture to the Long Now Foundation on Eusociality which may provide some insight. There is a podcast somewhere (try Youtube)


    His contact details are here: http://eowilsonfoundation.org/e-o-wilson/

    I am sure you are not particularly interested in the insights of a train driver!

  7. Kevin Cox
    January 29th, 2016 at 17:44 | #7

    In our work we are using Burgess’ Promise Theory as a theory of evolution and cooperation.

    In Promise Theory “autonomous entities” cooperate through the use of promises.

    We are using it to build a scalable personal identity.

    You can read some of what we are doing at http://www.welcomer.me/welcomer/blog/2016/1/12/a-description-of-identic . Identic is a member organization with any entity having an identity in Identic being eligible for membership.

    Unek.me is an application that provides an organization with a unique identity token for a person where every organization has a different token.

    Identic enables competing organizations to cooperate with respect to individual’s identities. This is a significant breakthrough in the development of IT systems as it provides a way of creating distributed cooperating systems. That is it combines identities through loose couplings to create a scalable identity system.

    We can use the approach to scale other things that prove problematic when we try to scale simply by making something larger. Money created for investments is a good candidate for the approach.

  8. QuentinR
    January 29th, 2016 at 18:38 | #8

    THOUGHTS: IMO, one group will survive at the expense of the other: the survivors will prove to be better at feeding, breeding and overcoming natural disasters, during the confrontation/interaction period, than the ones who die out.

    I don’t think your (JQ) assessment that the penguins’ social arrangements are more efficient counts much (if at all) towards their dominance over fowl. In a given space, how much can one species forage? In a given time, by how many can a group multiply? During floods/famines/fires/meteor strikes/occasional glut of predators, what percentage of each group survives?

    I realise that you say “In all other respects (diet, predators and so on) the two are similar.” But I don’t think you can predict (when two groups might first meet) which characteristic will be a defining one, so as to clearly predict which group might survive the confrontation period. Apart from the myriad of existing capabilities (which may assist or hinder the confrontation), how can you predict adaptability?

    Alternatively, if in all other respects they are identical, then perhaps they will interbreed and one or other of the mating rituals might continue or be replaced by something else. Who’s the winner then? – the group with the black eyes or the group with the blue eyes?

    So I favour your (a), perhaps because I don’t know what (b) [group selection] is, in academia.

  9. January 29th, 2016 at 19:59 | #9

    Well, I can point you towards some experts. Jerry Coyne wrote:

    “Group selection isn’t widely accepted by evolutionists for several reasons. First, it’s not an efficient way to select for traits, like altruistic behavior, that are supposed to be detrimental to the individual but good for the group. Groups divide to form other groups much less often than organisms reproduce to form other organisms, so group selection for altruism would be unlikely to override the tendency of each group to quickly lose its altruists through natural selection favoring cheaters. Further, little evidence exists that selection on groups has promoted the evolution of any trait. Finally, other, more plausible evolutionary forces, like direct selection on individuals for reciprocal support, could have made humans prosocial. These reasons explain why only a few biologists, like [David Sloan] Wilson and E. O. Wilson (no relation), advocate group selection as the evolutionary source of cooperation.”

    And Steven Pinker wrote on “The False Allure of Group Selection” here:


    However, I don’t see your example as having anything to do with group selection. There is species A with one set of genes and there is species B with another set of genes. If we place both in an area where they compete for resources, then the individuals with the genes that best suit them to obtaining resources will leave more decendants than those with genes that aren’t as well suited for obtaining resources. And if species A starts off with an advantage in genes they will tend to crowd out species B and vice versa, but all the selection is occuring at the level of the individual. A member of species A that has genes that are more fit than average will crowd out members of its own species as well as species B.

    So I don’t see group selection as being involved here.

    I think some confusion about group selection comes from biologists (almost) always regarding evolution as occuring at the level of the replicator. That is genes. Meanwhile other people look around and see that obviously some groups have advantages other others and clearly succeed over others, and so think that group selection is is a thing biologists accept. But as far as the genes are concerened, the group is just part of the environment they find themselves in. So it doesn’t matter to a gene if it finds itself in a monkey that is part of a group that has learned to crack nuts with rocks or in a monkey that is part of a group that hasn’t learned that trick, it’s all just part of the environment to the gene. The environment is slightly different between the two groups, but it still is just part of the environment, along with plant toxins, parasites, weather, predators, and so on. To the gene it makes no difference. Its only metric of success is whether or not it replicates itself.

    So if one’s definition of evolution is a typical biologist one, “Change in gene allele frequency over time.” Then one is unlikely to think that group selection acts upon genes in a way that is different from the environment in general. But if one’s definition of evolution is different from this, then depending on the definition, group selection could definitely exist for that person. But that definition of evolution is not going to be shared by most biologists.

  10. James Wimberley
    January 29th, 2016 at 20:16 | #10

    “It seems pretty clear that the great majority of evolutionary biologists reject the idea of group selection…” I thought the CW was that group selection is theoretically possible but not demonstrated in the wild. (expertise disclaimer)

    You might look at bacteria, who exchange genetic material directly. Evolution can surely be a lot groupier there.

  11. Ikonoclast
    January 29th, 2016 at 21:07 | #11

    It’s probably worth looking at the literature for Multilevel Selection Theory (MLS).

    Maybe the online Stanford Encyclopedia of Philosophy on “Units and Levels of Selection” will provide a useful overview. It has a long bibliography too.


  12. Ernestine Gross
    January 29th, 2016 at 21:11 | #12

    It seems to me you are using a social-science notion of a ‘group’. But game theory (and its evolution – no pun intended) is mathematical in nature. Hence I would first look up how biologists apply the math notion of ‘a group’. Do the disagreeing biologists differ in their applications of the ‘a group’?

    If I remember correctly, the difference between cooperative and non-cooperative games is the existence or otherwise, respectively, of ‘binding commitments’ (enforcable behaviour).

    I don’t understand why you consider the behaviour (of a group of ?) penguins more ‘efficient’ than the behaviour of (the group?) peacocks?

  13. Ikonoclast
    January 29th, 2016 at 21:17 | #13

    Also, David Wilson Sloane claims “The Tide of Opinion on Group Selection has Turned”.


  14. Paul Boss
    January 29th, 2016 at 21:19 | #14

    I read this blog a lot, but have not found reason to comment based on my lack of knowledge of the usual topics ( but I have learnt a lot!). However, as a biologist I would argue that natural selection acts at the species level and you shouldn’t compare between species. That is, a mutation that improves the survival of subsequent progeny is only relative to the other members of that species. Perhaps in some cases that means that those offspring can better compete against another species, but the main comparison should be with genotypes of the same species. I’m happy to be corrected if my understanding is outdated!

  15. January 29th, 2016 at 21:58 | #15

    Surely humans are the obvious example? Our supremely high level of cooperation (and a few other attributes) has led to us dominating the planet.

  16. BilB
    January 29th, 2016 at 22:08 | #16

    I would love to hear you expand on that PaulB.

    I am only guessing but Ronald B brought out one of the key factors in and that is competition for resources. But the periods of greatest species diversity seems to be at times of abundance. So the primary driver is environment, secondary driver is volume of reproductions, third driver is evolution and change in food sources (resources), fourth driver is competition for resources, fifth driver might be serendipity, and the sixth driver though no in order must be to do with epigenetics.

    My expectation would be that primary evolution occurs at the individual level and manifests itself at the group level. This is evolution of profound change where life develops new body section or extra limbs and is visible at the zygote level. Secondary evolution is where the “model” changes slightly in response to its everyday environment, the territory of incremental change through selection of the most successful.

    I think that the confusion arises because we tend to look at the evolutionary end result, rather than at the process, and that process has many facets, many more than Darwin could have fully appreciated. It is only now with gene research we see how genes are turned on and off to adapt the common gene set for best advantage in the prevailing environment. And with epigenetics further refinement of how experience can be passed forward to offspring (please correct me Paul B if I have the wrong take on what epigenetics is about).

    So I think that group selection is a genuine process but one that is an aberration of times of rapidly changing environment. We might be seeing a lot of group selection in the near future, or perhaps in the present if we knew what to look for.

  17. paul walter
    January 29th, 2016 at 22:20 | #17

    Much more of the human cooperation shown in places like Iraq and Syria and we will return to a very minimalist form existence indeed, if that is the word.

    David Cox, to me, moved things to the next stage by introducing the concept of identity and from there it is much easier to get to consciousness, (free)will and responsibility and even bad faith as to the fulfilling of what we think of as ethical obligations, something humans have tried to alleviate through the court system at this early stage in evolution.

  18. James Wimberley
    January 30th, 2016 at 00:54 | #18

    @Paul Boss
    “as a biologist I would argue that natural selection acts at the species level and you shouldn’t compare between species ..” Ad hominem, Darwin’s masterwork is called “The Origin of Species”! Species do compete and succeed each other in the same or overlapping niches, as we see with rats introduced to Pacific islands etc and from the fossil record. The question seems to be whether this works by interactive natural selection operating on groups of each species, or simple replacement.

  19. ARJ
  20. ARJ
    January 30th, 2016 at 05:59 | #20

    I should have also mentioned this recent article by DS Wilson, which is brief and helpful for seeing points of controversy:


    Also, I am no expert. My academic focus was computational biology, but in the area of protein classification, not evolution.

  21. Chris Johnson
    January 30th, 2016 at 06:17 | #21

    Hello John

    In your hypothetical example, the real problem is whether a trait like mate-feeding is more successful – in the sense that individuals who possess it leave more descendants – than a competing trait, like display. That determines whether the trait becomes established and elaborated by further selection. Group selection just doesn’t help us to explain that process, because if traits don’t improve individual fitness they are unlikely to become established in groups of individuals, for the simple reason that individuals will be more successful (leave more descendants) if they do something else.

    Of course, it is unlikely that the fully-developed behaviours that you describe would directly compete within a species. The competition would be among precursor traits that might (or might not) lead to those complete behaviour strategies and relevant morphologies – depending on selection operating on individuals.

    A biologist would not regard the hypothetical example you give as a case of group selection. It’s species selection, which is a different matter.

  22. Julie Thomas
    January 30th, 2016 at 07:43 | #22

    I like the way Peter Turchin is approaching this problem; there are lots of well worth reading posts at his blog Cliodynamica and the comments are thought-provoking and I haven’t read it but his book “Ultrasociality” is on my list.

    This is a post from 2012 that seems relevant to the question.


    “For a century, the primary account of evolution has emphasized the gene’s role as architect: a gene creates a trait that either proves advantageous or not, and is thus selected for, changing a species for the better, or not. Thus, a genetic blueprint creates traits and drives evolution.

    This gene-centric view, as it is known, is the one you learnt in high school. It’s the one you hear or read of in almost every popular account of how genes create traits and drive evolution. It comes from Gregor Mendel and the work he did with peas in the 1860s. Since then, and especially over the past 50 years, this notion has assumed the weight, solidity, and rootedness of an immovable object.

    But a number of biologists argue that we need to replace this gene-centric view with one that more heavily emphasises the role of gene expression — that we need to see the gene less as an architect and more as a member of a collaborative remodelling and maintenance crew.”

  23. Ivor
    January 30th, 2016 at 09:12 | #23

    There is a strange undertone to all this.

    What is an example of one species being driven out by another unless there has been human intervention? Dingos did not drive out native species, but introduced cats and foxes did. But this is not pure group selection – it is artificial disruption.

    The main point can be reworded from species to races.

    …suppose that two previously isolated species meet as a result of some change. In one species (aboriginals), competition between males for mates takes the form of elaborate, and energetically costly, displays. In the other species (europeans) males compete by providing food to their mates.

    In all other respects (diet, predators and so on) the two are similar. It seems obvious to me that the (europeans) with their more efficient social arrangements, are going to outbreed the (aboriginals) and eventually drive them to extinction.

    Natural selection – the survival of the fittest – is a form of evolution primarily responding to the environment and food supplies not competition or conquering other species. If one species is more efficient at getting food and expands its population another may die out or evolve further.

    This is evolution NOT group selection.

  24. Ikonoclast
    January 30th, 2016 at 09:14 | #24

    Finally, a quick sense I get of the matter, notwithstanding David Wilson Sloane’s statement linked to above, is that group selection as an evolutionary theory does not hold water. Kin selection on the other hand is worth looking at. Quotes from Jerry Coyne Ph.D who is a Professor in the Department of Ecology and Evolution at the University of Chicago and a member of both the Committee on Genetics and the Committee on Evolutionary Biology.

    1. “Group selection is a fuzzy and nebulous concept that is far less coherent than is gene-level selection (see Pinker’s essay for an explanation).”

    2. “When group selection does work in theory, it can be shown to be mathematically equivalent to gene-level selection involving “inclusive fitness.” But the group-selection scenarios are far more unwieldy, and are often so complex that they can’t be modeled.”

    Then Coyne quotes other scientists in the field

    “No group selection model has ever been constructed where the same result cannot be found with kin selection theory”.

    “The group selection approach has proved to be less useful than the kin selection approach.”

    “The application of group selection theory has led to much confusion and time wasting.” It is, as the authors say, “easy to misapply, leading to incorrect statements about how natural selection operates,” it is “not distinct from kin selection”, and it “often leads to the confusing redefinition of terms and the use of confusing jargon.”


    Lots of good links in that article to academic papers.

  25. Ken Fabian
    January 30th, 2016 at 12:30 | #25

    In the case of humans I think there probably is group selection but it is based on learned and applied behaviours – memes rather than genes. It can be passed on to others but imprecisely and like mutations, that imprecision can be disruptive as well as constructive. The extent to which individuals can learn and apply new behaviours (or tools or technique) may get back to genetic adaptation – one sub population being less or more capable than another – but because memes can propagate rapidly this kind of evolution potentially outpaces the genetic sort. I suppose it’s easier to lose those memes too.

  26. sunshine
    January 30th, 2016 at 18:24 | #26

    Apparently when Homo Sapiens arrived in Europe we were able to beat the Neanderthals into extinction (apart from some interbreeding). This is despite their superior physiques and bigger brains ,and was not due to any technological advantage . How did the runts of the human family predominate ? One idea is that we were better co-operators. We lived in groups of up to 150 whereas the Neanderthals only lived in small family groups. Another advantage we had was that we had begun co-evolving with dogs and they had not, so our group included non-human members . Assuming they would want to ,a big group made of runty humans and dogs could easily crush a small group made of superior individuals.

  27. Barry
  28. James Wimberley
    January 31st, 2016 at 02:21 | #28

    Runty Cro-Magnons that drew pictures. A secret weapon somehow.

    Query: why didn’t they make pictures of Fido? Or did they? For that matter, why are there ten times as many photos of cats as dogs on Facebook?

  29. James Wimberley
    January 31st, 2016 at 04:10 | #29

    Why are we commenters helping out JQ here? It’s a good simple case of altruistic behaviour.
    1. We have been indoctrinated as children with the value of being helpful, especially when it doesn’t cost much.
    2. We are competing for the approval of an authority figure, JQ.
    3. We feel paternal towards our own ideas. Realistically, self -publication is not an effective way of propagating them. It’s a better bet to try to convince JQ, who has a higher soapbox.

    Individual explanations like these seem to do the job. I am inclined to agree with Pinker and Coyne that group selection is a textbook case of entities multiplied without necessity.

  30. James Wimberley
    January 31st, 2016 at 04:17 | #30

    BTW, “being helpful” to strangers is a survival trait in a group. Cohesive groups will likely do better than individualistic ones. (There’s a nice theory that the survival of the early Christian church had a lot to do with its support of widows and orphans, which ensured grateful young members.) This operates at the level of cultural patterns, not genes.

  31. James Wimberley
    January 31st, 2016 at 04:34 | #31

    To return to the initial challenge. Bower birds and the like have evolved by sexual selection in a forgiving environment with plenty of food and few predators. The issue here is long-run competition between species. Perhaps we can treat the bower birds as a branch which will eventually peter out or break. In the long run – millions of years – they will be replaced by species from duller and more flexible lineages, like corvids. In turn these may start another dead-end branch of exotic sexual selection.

  32. Ron E Joggles
    January 31st, 2016 at 07:05 | #32

    Climatic change is a major factor here. The current interglacial brought a replacement of larger with smaller species in a range of environments. The robust Neanderthal ice-age hunters were no match for fast and nimble gracile modern humans. The latter presumably evolved a higher level of cooperation, hunting in substantial groups, and consequently complex language, symbolism, and thus intelligence and technological development. A crude wooden lance used primarily for thrusting was no match for a lightweight multi-component spear that could be thrown with accuracy from a distance.

  33. Julie Thomas
    January 31st, 2016 at 07:30 | #33

    “I am inclined to agree with Pinker and Coyne that group selection is a textbook case of entities multiplied without necessity.”

    It’s just that some of us haven’t managed to see the ‘necessity’ yet; but this realization is coming.

    And about cats and dogs, Rudyard Kipling – the nasty old racist – wrote a very strange story about cats and dogs and other wild animals including ‘man’. I did love the Just So Stories and thought I ‘got’ them…. except for this one.

    “Originally all the tame animals were wild, but especially the Cat: he walked by himself and all places were alike to him. The Man was wild too until he met the Woman, who chose a Cave for them to live in, lit a fire in it and hung a horsehide over the opening. She cooked a meal of wild ingredients.

    Then, while the Man slept, she took the bladebone of a shoulder of mutton and made a Singing Magic. This attracted the Dog, and on the next two nights she similarly lured the Horse and the Cow to visit the cave. They agreed to provide services to the couple, the Dog in exchange for roast meat and the other two for hay that she had dried by the fire. Each time the Cat followed and eavesdropped, called them fools, and went off to tell no one. ”


  34. Ron E Joggles
    January 31st, 2016 at 08:03 | #34

    Ron E Joggles :
    The latter presumably evolved a higher level of cooperation, hunting in substantial groups, and consequently complex language, symbolism, and thus intelligence and technological development.

    Of course, this occurred in Africa and was restricted to Africa until about 90K BP. African H.sapiens honed their skills in a highly competitive environment before exploring the rest of the planet.

  35. Ikonoclast
    January 31st, 2016 at 08:41 | #35

    @James Wimberley

    LOL, so true! We need to laugh at ourselves for this behaviour. At the same time, it is more pleasant for all concerned than “flaming”.

  36. Julie Thomas
    January 31st, 2016 at 09:25 | #36

    @James Wimberley

    “We have been indoctrinated as children with the value of being helpful, especially when it doesn’t cost much.”

    “From when they first begin to walk and talk and become truly cultural beings, young human children are naturally cooperative and helpful in many—though obviously not all—situations,” Tomasello said during one of two lectures about the origins of human cooperation. “And they do not get this from adults; it comes naturally.”


    Not much indoctrination is necessary to produce cooperative behaviour; children are more easily persuaded to help others than they are persuaded to be mean and greedy to others. It ‘feels’ better to be helpful and we realise this if we are able to experience, in fact or vicariously, these different ways of being.

    I think it takes a special sort of ‘indoctrination’ to produce the type of people who grow up to be libertarians and selfish greedy conservatives.

  37. Julie Thomas
    January 31st, 2016 at 09:28 | #37


    “We have been indoctrinated as children with the value of being helpful, especially when it doesn’t cost much.”

    But, “From when they first begin to walk and talk and become truly cultural beings, young human children are naturally cooperative and helpful in many—though obviously not all—situations,” Tomasello said during one of two lectures about the origins of human cooperation. “And they do not get this from adults; it comes naturally.”


    From my own experience and the growing amount of research it seems that not much indoctrination is necessary to produce cooperative behaviour; children are more easily persuaded to help others than they are persuaded to be mean and greedy to others. It ‘feels’ better to be helpful and we realise this if we are able to experience, in fact or vicariously, the different ways of being; scared and fearful of others or generous and willing to create a reciprocal relationship.

    I think it takes a special sort of ‘indoctrination’ to produce the type of people who grow up to be libertarians and selfish greedy conservatives.

  38. Ikonoclast
    January 31st, 2016 at 10:29 | #38

    @Julie Thomas

    Well, I see propensities and potentialities and I find them ambiguous. Children are both cooperative and competitive. I wouldn’t like to guess which is dominant before socialisation begins. Indeed, it might be hard to argue that the question of which is dominant makes any sense before socialisation begins. Among other things, there is no way to test it at the baby stage. Toddlers are already partly socialised and indeed significantly so. The fact that most behave relatively well according to Michael Tomasello’s studies may well be a tribute to what mothers do before wider society gets hold of their children. I do not see how he could support the broad-brush claim: “And they do not get this from adults; it comes naturally.” Anyone who has played a role in turning babies into toddlers knows this is an unsupportable assumption if made in these absolute “all of one thing, none of the other thing” terms.

  39. Julie Thomas
    January 31st, 2016 at 11:25 | #39


    “The fact that most behave relatively well according to Michael Tomasello’s studies may well be a tribute to what mothers do before wider society gets hold of their children. I do not see how he could support the broad-brush claim: “And they do not get this from adults; it comes naturally.” Anyone who has played a role in turning babies into toddlers knows this is an unsupportable assumption if made in these absolute “all of one thing, none of the other thing” terms.”

    Well I guess it was a bad paper if Tomasello didn’t explain how he supported his claim that it “didn’t come from the adults”. But, there are a lot of similar studies that come to the same conclusion about the propensity of infants to be sociable rather than selfish so I don’t think that it is guessing at all.

    And the idea that “anyone who has played a role …” is a bit sus. How many have you raised and how detailed was your observation? Were they your own children or are you able to compare your children with other children and the way they respond to their environment?

    Do you know about the idea of ‘temperamental differences’ in children – which is supposed to show that we are not blank slates – but it doesn’t. It shows that each slate is ready to be written on but each slate has individual characteristics that mean writing on it will require different writing implements if you want to create an adult.

    There are an awful lot of myths about ‘mothering’ and how good women are at it. Did you know that most first time mothers – unless they have had the experience of what is called ‘alloparenting’ – are unable to feed their infants without instruction from a nurse. The nipple needs to be fed or pushed into the babies mouth; it’s not sufficient for the infant to simply latch on and suck.

    Check out the idea of “alloparenting” and Sarah Blaffer Hrdy’s work in her book Mothers and Others.

    And you can be dubious about this idea too, but I think it is simply obviously ‘true’ :), that it was the ‘evolution’ of long lived women and allo-parenting that gave us the insights we needed to dominate the environment and the other animals.

  40. Ikonoclast
    January 31st, 2016 at 14:25 | #40

    @Julie Thomas

    I am not sure that sociable and selfish are psychological antonyms though they might be dictionary antonyms. I have seen sociable cooperative people and I have seen sociable selfish people. The latter are adept at seeming “nice”, seeming friendly, but are very good manipulators and exploiters of others and just about always have ulterior motives.

    But I will agree that socialising for cooperation is far more beneficial for society and the bulk of individuals. This would hold true I think whether or not we started out cooperation-biased, neutral or selfish-biased.

    Cooperation is clearly a work output multiplier. I have had occasion to compare painting a house exterior alone to painting a house exterior with a co-worker. These were two-story houses. Moving and setting up trestles and planks (just one example) becomes not half as difficult but about a quarter as difficult when you have two persons compared to one. In other words, if this ratio extended (which I don’t know if it does or doesn’t) then work output would increase as the square of cooperation. There are cases where work output increases infinitely by cooperation. Where human muscle power alone is used, there are loads one person can never move but six persons can move it quite effectively. In a more modern setting, 12 persons could run an assembly line with 12 stations requiring manual actions but any less than 12 could not run the assembly line at all so the increase in productivity from proper cooperation is again infinite.

    There in a nutshell, in my view, is why cooperation is vastly superior to competition, in economics anyway. Much of what people generally term competition is merely competition between firms. Within the firms, hundreds or even thousands of people cooperate to make things. Then the firms “compete” in what should be termed oligopoly or cartel competition which is far different from textbook economic competition which is a mythic construct. Textbook competition for the most part is nothing like what happens in the real world.

    If something approaching real competition occurs in our real economy, apart from some small business competition, it’s on the labour side where workers are pitted against each other and against the skilled unemployed to sell their labour. This competition has few benefits and many costs when it becomes exaggerated. It would say it is exaggerated whenever the unemployment rate is higher than frictional unemployment.

    The above is probably a “bloke” argument as well as a Marxian one.

  41. GrueBleen
    January 31st, 2016 at 15:34 | #41


    A good post, Ikono – your one of Jan 31st at 14:25pm above. I would have expected Jerry Coyne to be one of the very first experts considered in discussing ‘group selection’. P Z Myers (as expounded by Barry a post or two later) would also have been a candidate. Also Laurence Moran at the Sandwalk blog (just Google Sandwalk blog).

    But all of those possibilities would have ignored the fact that ProfQ is proposing an extraordinarily idiosyncratic definition of ‘group and ‘group selection’. Go back to ProfQ’s actual post and contemplate this (at the end):

    “It seems to me there are only two possibilities here
    (a) My reasoning is wrong, and we can’t judge which species, if either, will dominate; or
    (b) Even though it involves one group being selected over another, this isn’t what is meant by group selection”

    This is directly based on profQ’s discussion of penguins outcompeting peacocks and indicates that he is thinking of “the group” being essentially equivalent to “the species”. Most of the discussion, including the experts so far proposed, take “the group” to be intra-species.

    After all, if we’re going to go to the species level, why not go all the way up to the Clade. Why not consider, for example, that our very, very remote ancestors – the original synapsids – were actually Earth’s dominant major (ie largish scale) life form, and that the sauropsids (otherwise known as ‘dinosaurs”) were subordinate to we synapsids. Hooray. Then came “The Great Dying” – the very, very fatal Permian-Triassic extinction of about 252 million years ago. And guess what – those god-fearing sauropsids displaced our ancestors as the dominant macro clade.

    But then … well, about 65 million years ago we had the Cretaceous–Paleogene extinction, and all those very large sauropsids, and some fairly small ones too, went the way of our long gone synapsid ancestors. But a bunch of fairly small, generally furry, mostly nocturnal scavengers and vegetarians – synapsids all of them – survived and thrived. Ain’t that just miraculous ? 65ma we simply out-competed them at the clade (group) level.

    And then, round about 5ma (5 million years ago) the evolutionary path that ends (so far, anyway) with us began. So obviously, if you aren’t too fussy about timescales, it is clear that ‘group selection’ worked to replace the dinosaurs by human beings.

    Ain’t that the clear and obvious truth ?

  42. harleymc
    January 31st, 2016 at 20:50 | #42

    John while Darwin socialised with a lot of economists the two sciences of evolutionary biology and economics share no common basis.

    (b) Even though it involves one group being selected over another, this isn’t what is meant by group selection.

    A group in evolutionary biology is an association (an in-group) with no more or no less genetic similarity to each other than the out-group. In other words it is not a kin group. So whether the in group or the out group somehow gets a higher number of off spring, doesn’t change the frequency of genes.

    Kin selection is a diferent model and is a strong basis for the evolution of both cooperation and altruism.

  43. January 31st, 2016 at 21:56 | #43

    Despite not being an expert, I will have a crack at explaining why most biologists reject group selection, which I didn’t really do in my previous comment.

    To almost all biologists, biological evolution refers to the selection of different gene varieties. So if there is a group of monkeys that learns to cooperate together better, and as a result this the group expands and outcompetes other monkey groups, then that group of cooperative monkeys would certainly have succeeded at survival and expanded at the cost of less cooperative monkey groups, but no group selection has taken place in this example. This is because the monkies’ genes haven’t changed as a result of learning to cooperate better. No more than the genes of a human child change when they learn to cooperate from watching Sesame Street.

    Okay, you might say, so that’s not group selection according to biologists because there is no selection of gene varieties going on. The monkeys were just using learned behaviour that presumably any monkey group could learn under the right circumstances. But, you may say, what if a monkey has a mutation that makes it more cooperative? And this genetic change helps it and its offspring survive and it spreads through the monkey group making the group more cooperative as a whole and then it outcompetes other monkey groups. Since there is a genetic change, you might think this is definitely an example of group selection.

    But it isn’t. You see, the group wasn’t responsible for that mutant gene spreading. It spread because it helped the individuals who had it to be successful and leave more offspring. It is just an example of natural selection which operates at the level of the individual.

    For group selection to occur, there would have to be a genetic change that is selected for that benefits the group and not individuals. And as far as we can tell, that doesn’t happen. Or at least no clear, undisputed examples have been presented so far.

    One important reason we don’t see group selection occurring is that any genetic change that advantages the group but disadvantages an individual means that the individual with that genetic change is less likely to survive and pass it on. And if the genetic change does make the individual more likely to survive and reproduce, then it is not group selection but plain old natural selection that is occurring.

    So, John, I would say we definitely have a case of (b) here. That is: Even though it involves one group being selected over another, this isn’t what is meant by group selection.

  44. Fabrizio
    February 1st, 2016 at 01:20 | #44

    In the situation you describe, both the penguin strategy and the peacock strategy seem evolutionary stable. A peacock adopting the penguin strategy would not be able to invade the incumbent peacock population because he would be unable to attract mates. Symmetrically, a penguin adopting the peacock strategy would be unable to invade for the same reasons. What you appear to be describing is a situation with multiple equilibria where one equilibrium is more efficient than the other. In this case, it is plausible that, in the very long run, the more efficient equilibrium may prevail. (I believe I heard Ken Binmore defending this approach in a seminar, you may want to look at his recent stuff on sexual selection).

    As far as I understand, when people criticize group selection, they cast doubts on outcomes that are incompatible with equilibria of the underlying game. This is not the case you are describing.

  45. James Wimberley
    February 1st, 2016 at 07:31 | #45

    @Julie Thomas
    You are probably right that there is a genetic basis to human cooperation. Small children are both possessive and sharing at different times. SFIK other animals do not share food and other resources like toys at all. Apes groom each other, which is a social gift exchange but not the same. I amend “indoctrination” to “reinforcement”.

  46. James Wimberley
    February 1st, 2016 at 07:39 | #46

    “In the long run, the more efficient equilibrium may prevail”. This fits a point I tried to make earlier using the branching tree metaphor. In support, look at the pronking of springboks. There are many species of the antelope family on the African savannahs, but IIRC only one goes in for this nonfunctional and costly display. Sexual selection is Nature’s evolutionary joker, but it’s not the commonest card.

  47. James Wimberley
    February 1st, 2016 at 07:50 | #47

    Stephen Jay Gould pointed out that survival through mass extinctions is down to luck. The features that allowed one species or family to get through Snowball Earth or the Mexican asteroid or the Deccan Traps were by definition irrelevant to their adaptive fitness in normal times. You can call this competition between species if you want, but it’s very far from Darwin’s scheme of insensible change.

  48. GrueBleen
    February 1st, 2016 at 08:14 | #48

    @James Wimberley

    No, actually James, I called it competition between clades, not species, if you read what I wrote. That was just a bit of “exaggeration for dramatic effect” though, so don’t give it any attention.

    And it may have escaped normal observation – I’m not familiar with everything that S J Gould ever wrote – but it was precisely those features that allowed our synapsid ancestors to coexist with rampant dinosaurs, namely small (less food needed), furry (kept warm in planetary winter), nocturnal (can handle extended darkness), scavengers and largely vegetarian (can get by on almost anything) that greatly contributed to their post Cretaceous–Paleogene survival.

    But yes, I guess it was just luck that our ancestors had had to adapt to survive in that way in a dinosaur dominated world that allowed for their survival. So I guess Gould’s reputation is intact.

    However, tell me what animal species survived ‘snowball Earth’ (if in fact it ever happened), and just what ‘luck’ enabled them to do it ?

  49. J-D
    February 1st, 2016 at 09:48 | #49

    The original example describes interaction between two different kinds of organism (‘peacocks’ and ‘penguins’) resulting in one kind becoming extinct — but it isn’t clear how.

    One way that interaction between two different kinds of organism might result in one of them becoming extinct would be if there was a by-product of the metabolism of type P organisms that was lethally toxic to type Q organisms.

    This is not pure speculation: for example, free atmospheric oxygen is a by-product of the metabolism of photosynthetic organisms but is lethally toxic to the kind of organisms that biologists refer to as ‘obligate anaerobes’.

    It seems to be generally agreed by scientists who study the topic that the concentration of free atmospheric oxygen in early geological history was negligible, and that it rose to present levels principally because of the evolution and multiplication of photosynthetic organisms. Many biologists also think that this resulted in the extinction of many obligate anaerobes. (Obligate anaerobes still exist, but not in environments exposed to the atmosphere.) Biologists don’t treat this idea as an example of ‘group selection’. Biologists who reject ‘group selection’ don’t mean ‘it never happens that one group of organisms drives another to extinction’; that it sometimes happens that one group of organisms drives another to extinction is an accepted fact of biology, but it’s not what biologists (pro or anti) mean by ‘group selection’.

  50. February 1st, 2016 at 10:42 | #50

    One aspect of the theory says that male peacock expenditure on plumage provides a way for the female peacocks to judge effectively which males are the healthiest, thus resulting in healthier offspring; and if this is true then the competition between better-fed and healthier young penguins versus better-genetically-selected and healthier young peacocks is not clearly going to favour penguins.

    Certainly in the human and sociological arenas we see the signalling strategy “Only a very powerful person could survive the utterly idiotic proposal I am about to impose, which proves I am more powerful than you and you should adopt the deferential position” in the workplace every day, and often working very well for the individuals concerned.

  51. Ikonoclast
    February 1st, 2016 at 11:45 | #51

    I haven’t yet seen a clear definition of what “group selection” is supposed to be. I haven’t seen clear definitions of replicators, interactors, “vehicles” and groups. Nor have I seen a clear statement of which of the above is the “real unit of selection”. Until an advocate of group selection can clearly define all of the above (and/or rule some out as meaningless or redundant) then the discussion cannot even begin.

    Herding animals are an interesting case. Is the gene for herding (assuming there is one) sensibly regarded as an individual possession or a group possession? I have no idea how to answer that.

    One researcher has written;

    “Try this new line of thought for a moment: what if we dropped the idea of group selection and instead discussed interaction selection? This would be a study of how the interactions between two or more individuals, between two or more groups, or between two or more species resulted in differential fitnesses of organisms or genes. In discussing interaction selection, we would transcend strict essentialist concepts such as individual selection and group selection, and reach a more pure form of analysis in which the process is emphasized rather than the unit of selection.” – Rebecca Beroukhim

    This contains the germ of a very interesting idea I think. From the point of view of complex systems theory it seems to make a great deal of sense.

  52. GrueBleen
    February 1st, 2016 at 13:37 | #52


    Frustrating, isn’t it. Re your first paragraph, I seem to remember saying very similar things in the thread on the ASIO guy supposedly “ringing around to politicians” re IS and Islam. What is this “normal process” I asked, and how many pollies did he ring, and what did he say to each of them. I never did get an answer, and you’re not going to get one either.

    And you’re absolutely right: without at least some answers, there’s nothing to discuss.

    Never mind, mate, I put it down to bio-quantum entanglement, myself: whatever happens to a dominant individual also somehow affects the group (if only we could know what group is).

  53. sunshine
    February 1st, 2016 at 14:23 | #53

    @James Wimberley
    ‘ other animals do not share food and other resources like ‘ [humans do] .

    Tho we are certainly very good sharers ,I think it would be hard to define sharing so that there are there are no examples of it in the non-human animal world. The urge to do so is more interesting to me than the definition itself. Rick Dawkins says there is almost no basic kind of human societal structure that cant be found in the ant world. On that a-bit-off-topic line of thought ,although we currently direct massive resources our way, wouldnt the champion groups on long time scales have to be the insects ? There are so many species we cant even count them quicker than they mutate into new species .Our knowledge will never completely envelope them.

    Not having done any supplementary reading, I dont think I have a good handle on the group v’s individual selection debate . As Ronald Brack points out above -it seems that by virtue of the definition of terms ,genetic group change will always be traceable back to genetic change of an individual . However there does seem to be lots of recent evidence showing that expression of genes can be affected by environment and environmental effects on parents can be passed several generations down. There is a continuous complex two way interaction between genes and the world. Is there no way group action could change genes ? Is that what Prof Q is looking for? And wouldnt people then just say ,as Ronald pointed out, ‘well that group action must just be the result of genes anyway’ .As so often happens I suspect the answer may lie in better defining the question .

  54. Jim Birch
    February 1st, 2016 at 14:55 | #54

    I can’t understand the debate either. It seems a case of science advancing with the death of each scientist.

    I think the problem might be conceptual tension, as in ‘By discarding inclusive fitness on the basis of its limitations, they [Nowak et al] create a conceptual tension which, we argue, is unnecessary, and potentially dangerous for evolutionary biology.” From Ferriere & Michod Nature 471, E6–E8 (24 March 2011)

    BTW, your example is incorrect. Penguins outcompeting (outcooperating) peacocks is standard evolution. Group selection would be where one group of peacocks have a gene that makes them more penguin-like, grow smaller tails, spend less energy in intra-group competition and collectively prosper relative to the peacock muster across the valley. The problem is working out what would cause the cooperation gene to propagate from an individual mutation into the group if the benefit is applied to all individuals in the group. It may be a smart idea but that isn’t enough. This is a kind of magical thinking which is present in popular thinking about evolution and I think this might be part of the issue with group selection.

    As I understand it, what Nowak did was show that group selection is actually mathematically possible although under a more limited limited set of circumstances than the magical version. In the process he also take a swipe at the inclusive fitness criteria – which has been a bit of a cornerstone of the evolution theory for a few decades – and attempts to replace it with something more like a population model than a simple criteria that applies to the individual.

    There’s some analysis of the arguments here: http://www.christopherxjjensen.com/2010/10/13/robert-trivers-and-colleagues-on-nowak-tarnita-and-wilsons-the-evolution-of-eusociality/

  55. February 1st, 2016 at 16:57 | #55

    Again, I am no expert, but I’ll have a go at defining some terms:

    According to wikipedia, evolution is: Change in the heritable traits of biological populations over successive generations.

    While the terminology is different, that’s very similar to the one I gave earlier – change in gene allele frequency over time.

    And again, according to wikipedia, group selection is: A proposed mechanism of evolution in which natural selection is imagined to act at the level of the group, instead of at the more conventional level of the individual.

    I would expand on that by saying that for group selection to have occurred, the genetic change selected for would have to benefit the group and disadvantage the individual that has it. If that is not the case and the genetic change benefits the individual, then it is not group selection, but selection at the individual level instead.

    It is easy for people who are not familiar with the subject to think that any selection that benefits the group is group selection, but that is not the case. Individual birds benefit from hanging out with other birds in a flock because more eyes mean predators are more likely to be spotted. So the gene that encourages flocking behaviour is not group selection. I know this may sound odd because if birds didn’t flock there wouldn’t be a group at all, but that’s not what group selection refers to.

    I’ll mention that one behaviour that has been proposed as an example of group selection, the warning cry a bird gives when it spots a predator such as a stalking cat, may not be for the benefit of the other birds. One of the multiple possible reasons, several of which may apply at once, is it is actually a message to the cat telling it, “I’ve spotted you and I’m already escaping. Go for one of the slower birds, not me!” So even for examples that have been put forwards as being group selection alternative, competing explanations always apply. So far we have no clear examples of group selection occurring outside of special examples such as social insects where all the individuals in a colony are closely genetically related.

    Harleymc gave a definition for group above, which I think is the one used for discussing population genetics, but biologists also use the term group more informally to mean, “This bunch of thing I am talking about,” and it might be easier for us to just continue using it that way ourselves.

  56. Ikonoclast
    February 1st, 2016 at 17:11 | #56

    @Ronald Brak

    And as harleymc explained, in-group (not kin-group) success or failure “doesn’t change the frequency of genes” so there is no selection pressure:

    “A group in evolutionary biology is an association (an in-group) with no more or no less genetic similarity to each other than the out-group. In other words it is not a kin group. So whether the in group or the out group somehow gets a higher number of off spring, doesn’t change the frequency of genes.” – harleymc.

    By that definition of “the group” there can no such thing as biological natural selection stemming from group success.

  57. February 1st, 2016 at 18:00 | #57

    Ikonoclast, yes, that’s right.

    But I’m not aware of people of people using that definition when discussing group selection. If we strictly define a group as being a bunch of organisms with gene frequencies that are identical to other bunches of organisms, then group selection is impossible. Along with any other form of selection. In other words, if we strictly follow the definition, we freeze evolution.

    ORGANISM 1: Hey! My offspring has a novel mutation!

    ORGANISM 2: No it doesn’t, because if it did, that would make our group’s gene allele frequency different from other groups, and by definition, that is impossible.

    ORGANISM 1: Oh, you’re right. Its DNA just popped back into place via the power of definition.

    That definition of group is a bit of jargon that is useful in the right circumstances, for example discussing population genetics where they also sometimes declare organism populations to be infinite and mating to be completely random to make their equations simpler. But it’s not useful when discussing group selection because it defines away the possibility of group selection or any change in allele frequency at all, and we know from experience that changes in gene allele frequency can and do occur.

    So I would suggest we not use that definition for group, or at least not apply it too strictly, because it contradicts what we have learned from both observation and experiment, which is gene allele frequencies can change over time. (They just don’t appear to change due to group selection.)

  58. dedalus
    February 1st, 2016 at 20:28 | #58

    Evolution is both adaptive to and exploitative of environments. There are two processes going on here. One is biological, the other cultural.

    “Adaptive” is suitable for looking at passive changes in the biology of large groups or species over very long time spans. “Exploitative” suits the description of behaviour within groups or species reacting more proactively to an environment in the short term. Exploitative evolution is mainly only observable in higher primates – specifically humans. To say a certain species of plant “exploits” an environment is wrong. (This is often said of introduced species – but in fact an introduced species will thrive not through exploitation but simply through being suited to a previously addapted environment.)

    Evolution in most species, then, is a mainly adaptive process taking very long times to work through. Human evolution by contrast also involves an exploitative process leading to relatively rapid changes, mainly cultural. For example the height and longevity of humans has markedy increased over the past 50 of so years, but this is not the result of biological adaptataion, but rather of cultural exploitation, namely the marketing of medical and nutritional advances. It is this mainly exploitative nature of humans that has led to the destruction or farming of other species.

    In the norms of long term evolution, we can’t really predict how the human species will “evolve”. We can only speculate on cultural-type advances (science, politics etc).

  59. Ikonoclast
    February 2nd, 2016 at 05:59 | #59

    @Ronald Brak

    I don’t follow your reasoning at all. The in-group definition given by harleymc does not define away individual selection. It’s just that individual selection under that definition would be precisely that, individual selection.

    What is your suggested definition of group for the purposes of testing a group selection hypothesis?

  60. J-D
    February 2nd, 2016 at 10:24 | #60

    Suppose biologists studying some particular kind of organism observe an increase over time in the frequency of a particular gene (let’s call it P, which could stand either for ‘peacock’ or for ‘penguin’) in the population’s gene pool.

    They might hypothesise something roughly like this:

    Hypothesis 1: Gene P has effects which result, directly or indirectly, in individuals with gene P producing more offspring that survive to reproductive age (than do other individuals without gene P).

    (If something like Hypothesis 1 were true, the natural expectation would be for the frequency of gene P to increase; and if biologists were thinking along those lines, they could go on to investigate, in more specific detail, what the effects of gene P are and how they might have that kind of result.)

    Something like that could be an ordinary kind of evolutionary explanation that would be unlikely to be referred to as ‘group selection’.

    But suppose there’s another hypothesis:

    Hypothesis 2: Gene P has effects which result, directly or indirectly, in groups that include some individuals with gene P producing more offspring (that survive to reproductive age) than do groups with no individuals with gene P.

    Biologists who favour the idea of ‘group selection’ are interested in finding cases where something like hypothesis 2 is true.

    Biologists who reject the idea of ‘group selection’ tend to argue along the lines that it’s fairly predictable that hypothesis 2 will be true in cases where hypothesis 1 is true (at least often), that in cases where hypothesis 2 is true it’s only as a by-product of hypothesis 1 being true, and that hypothesis 2 gives us no extra information and adds no value.

    I think it would be interesting if there were cases where something like hypothesis 2 is true but something like hypothesis 1 is not true. But are there? Biologists who reject the concept of ‘group selection’ seem to doubt it.

  61. Ikonoclast
    February 2nd, 2016 at 11:10 | #61


    In hypothesis 2, I think you have described a kin-selection variant. Nobody, so far as I can see in this thread, has described any form of group selection which goes beyond kin-selection and which makes any kind of logical sense. The idea of group selection, given what selection is, seems to a non sequitur. I’ve seen nothing so far here or elsewhere to convince me that group selection makes any sense with strict respect to genes as the replicators.

    It seems to me (though as a non-expert I really don’t know) that for any group selection theory to work, the replicator, formally understood, would have to be above the level of the gene. We would have to conflate the replicator with the interactor at some level i.e. with the individual or the group itself. Does this idea work biologically or mathematically? I really don’t know but I doubt it. I think we are all stumbling around here without a clue.

  62. February 2nd, 2016 at 12:05 | #62

    Ikonoclast, let’s say we have two groups of insects with identical genetic similarity. That is gene allele frequencies are the same. One group is on island A and the other group is on island B. On both islands insect numbers are controlled with DDT. On island A a chance mutation results in resistance to DDT and spreads through the population. On island B this does not happen.

    So now we have two populations that differ in gene allele frequency. But if a group has “no more or no less genetic similarity to each other than the out-group”, then we have a problem because we do have a group that differs in genetic similarity. So either the population on island A is no longer a group and we have to call it something else or the situation I have described is imposible and so never happens:

    REALITY: Luke, allele frequencies in group A differ from the out-group.

    DEFINITION: No. No! That’s not true. That’s impossible!

    REALITY: Search your polymerase chain reaction results. You know it to be true.

    DEFINITION: No! No! If that’s true, then you can’t call it a group anymore!

    REALITY: Now you’re just being silly. Join me and together we will document the changes in allele frequency that are occurring before our very eyes.

    DEFINITION: I will never join you. [Jumps down conveniently located chasm.]

    So I don’t find that definition of group very useful. Not when discussing selection since if there is no change in allele frequency then no selection has taken place. So I propose the following definition of the word group: The word group means “the bunch of things that I am talking about.”

    So what do you think of the population on island A that has developed pesticide resistance, Ikonoclast? Do you:

    (a) Think it is impossible for a change in allele frequency to have occurred since the definition of group you quoted states that a group has “no more or no less genetic similarity to each other than the out-group”. That is, no genetic change has actually occurred.

    Or do you:

    (b) Think that a change in allele frequency has occurred and therefore we can’t refer to the population on island A as a group since the definition states a group has “no more or no less genetic similarity to each other than the out-group”. So we have to call it something else now.

    Or do you:

    (c) Think that there has been a change in allele frequency on island A and we can still refer to it as a group. That is, you think the following statement is acceptable, “The group on island A is genetically different from the group on island B.”

  63. Ikonoclast
    February 2nd, 2016 at 12:49 | #63

    @Ronald Brak

    The answer is “c”. You can talk about groups in any way you like (as in dividing by island of residence as you do).

    However, if on island A, a chance mutation happens, in an individual called let us say “Alexander Beetle”, then it can only “spread through the population” by reproduction. All of the later resistant individuals on island A will be descendants of “Alexander Beetle”. It’s a clear case of selection of individuals. Which I assume you realise.

    So you have “proved” the trivial assertion that we can define “groups” any way we like and that this adds nothing to the discussion of group selection as a concept. What is your point?

  64. February 2nd, 2016 at 13:51 | #64

    Ikonoclast, my point is that if we are going to discuss selection, then we have to accept at least the possibility that groups can differ in allele frequency which is something that can’t happen if we define a group as, “…an association (an in-group) with no more or no less genetic similarity to each other than the out-group. In other words it is not a kin group. So whether the in group or the out group somehow gets a higher number of off spring, doesn’t change the frequency of genes.”

    Since you chose (c) you appear to agree with me.

  65. J-D
    February 2nd, 2016 at 13:52 | #65


    No …

    … or at least I don’t think that’s the clearest way of putting it.

    As far as I know, biologists generally accept that kin selection happens. A generic description of kin selection using the same kind of wording and structure that I used for the other generic descriptions I gave would be something roughly like this:

    Gene P has effects which result, directly or indirectly, in individuals closely related to the P-bearing individual producing more offspring (that survive to reproductive age).

    In a situation where that is true, both hypothesis 1 and hypothesis 2 would be true; kin selection is one example (but not the only example) of what I described earlier, hypothesis 2 being true only as a by-product of hypothesis 1 being true, so that hypothesis 2 gives no extra information and adds no value (and, in fact, kin selection was an example I had in mind, and not the only one–I left out the examples I was thinking of for simplicity).

    By contrast, the idea of kin selection does add something; it is more specific than hypothesis 1 by itself, although still a fairly general concept.

  66. Peter T
    February 2nd, 2016 at 14:25 | #66

    Kin selection ids favoured because it is micro-founded – it rests on the established mechanism of transmission of traits – the genotype.

    Kin selection: selects for actions which are disadvantageous to individuals but overall advantageous to the genotype. The eusocial insects (and a few others) do this by ensuring a very high degree of genotypic similarity in the group – typically by concentrating reproduction in one or a very few individuals (the queen in bees or ants and the naked mole-rat). So the male mole-rat that flings itself at the invading snake is trading its future sons and daughters for a quantity of nieces and nephews large enough to outweigh the loss.

    The challenge is to find a way that actions disadvantageous to the individual can persist in the absence of genotypical similarity. Humans clearly achieve this (just search the news for “suicide attacks”). How? My guess is the work-around involves neotony, the central role of culture in brain development and the large-ish minimum group size needed to sustain a culture (roughly 1000).

  67. jungney
    February 2nd, 2016 at 19:15 | #67

    You do know, JQ, that Mary Midgely has described the misapprehension of Darwin in these terms:

    “Thus the picture of natural selection which was shaped around Darwin’s main proposals used only two sombre colours – black for death and white for survival. Nature appeared in it only as an obsessive accountant, spectacles on nose and ledger in hand, testing every action in those terms and destroying the failures. Any trait still appearing in the world was deemed to have passed her audit in some distinct, discoverable way which constituted its evolutionary function.”

    She places the interpretation of Darwin within the distorting ideological landscape of liberalism.

    As to the errors within the group selection dialogue it is not necessary to say more than that the attribution of causality to one component of the whole, the gene, is such a pit of exhausted debate that there are only bones at the bottom. Peering in from the top of the pit are the poets, artists, writers, dancers, musicians, dreamers, philosophers and believers who constitute the rest of the whole of which the gene is but a mere switch.

  68. J-D
    February 3rd, 2016 at 08:30 | #68

    What I want to know is this: has all this discussion been of any use to John Quiggin?

  69. Ikonoclast
    February 3rd, 2016 at 09:29 | #69


    I was wondering that. I suspect not as we are all amateurs bumbling around with unscientific ideas on this topic. I was idly wondering if this post was a sociological experiment. You know, as in; post something a bit “left-field” and see how silly the replies get. I’ve noticed a strong tendency in myself and in others, for the need to have an opinion on everything, even on things we don’t know anything about.

    Clearly it has to do with discomfit about uncertainty. We humans seem to dislike uncertainty intensely. Uncertainty roughly aligns with insecurity. Indeed, we dislike uncertainty so much we manufacture false certainty by taking positions without evidence. We seem to be wired this way to some extent. It makes humans such fertile ground for dogma, denialisms and fundamentalisms. By contrast, it takes discipline to admit lack of knowledge, go looking for empirical evidence and then form opinions only after that process if such opinions are justified. It takes wisdom and resignation to realise there are many things we can never get certain knowledge about and so we must always live with considerable uncertainty.

  70. John Quiggin
    February 3rd, 2016 at 10:07 | #70

    I’ve got a couple of benefits from reading the comments here and at Crooked Timber. One is that there have been lots of interesting points raised, in a generally civil fashion. The other is to determine that the idea of group selection is both controversial and not very well defined. So, while I was thinking of referring to an analogy with group selection in some work I’m doing, I’ve decided this would create more confusion than enlightenment.

    So, thanks everyone for talking about it, and feel free to keep going.

  71. Jim Birch
    February 3rd, 2016 at 13:01 | #71

    Evolution theory is a bit like economics, the actions and impacts are statistical ensembles but everyone wants to distil out coherent actors, intents and narratives.

  72. ZM
    February 3rd, 2016 at 18:58 | #72

    There was an article about Australian Fairy Penguins taking over the habitat of New Zealand Penguins 400 years ago in The Guardian today.

    A scientist said these two groups of Penguins were difficult to distinguish apart from their behaviour, Australian Fairy Penguins:

    “come ashore in big rafts of 100 or 200…. The Australian ones can breed twice a year, while the native New Zealand one can only breed once, Waters said. Digital audio analysis also revealed they had different calls, Waters said. “It’s a bit like the accent of saying ‘fush and chups’ rather than “feesh and cheeps’,” he said.
    Many of New Zealand’s animal species, birds in particular, have suffered at the hands of people. The really exciting thing about these findings is that they show how quickly nature can respond to human impacts,” said Jonathan Waters from the University of Otago, who led the study.”


  73. February 4th, 2016 at 16:58 | #73

    I have mentioned why with the example given (b) applies. That is, it is not what is meant by group selection. But I will also make a comment or two on (a) which is whether or not we can judge if one species will dominate. Note that my comments don’t concern group selection as it does not apply in the given example.

    The “penguins” appear to have a better system than the “peacocks” (or “peafowl” if you prefer) because penguin males contribute resources to egg production and chick raising and so it appears that species is able to devote much more resources to reproduction than the peacocks where males contribute nothing but gametes.

    But, a colony of 50 female penguins, 49 contributing males, and one male who spends all his time commenting on the internet, has the same amount of resources to contribute to reproduction as a colony of 99 females and one sufficiently fertile male. So maybe the peahens stumbled onto a good thing by insisting the peacocks have ridiculous tails, as it must result in a lot of males getting killed and so reduce the numbers of the mostly useless male portion of their species.

    However, the ridiculous tails probably don’t kill enough males for an efficient male female ratio to result. But while the surviving male peacocks would be strutting around the place gobbling up resources that could have gone to females and therefore reproduction, they would presumably also be gobbling up resources that could have gone to penguins. So male peacocks could be a pox on both penguins and peahens. While peahens have to waste resources producing males in the first place, once they leave the nest they might be an equal burden to both species. (While not relevant for the example, real peafowl chicks can fly within days of hatching and with help from their mother start finding food for themselves almost immediately, and so take less effort to raise than the chicks of many other bird species.)

    If we assume all else is equal between the “penguins” and the “peacocks” then it looks to me that the “penguins” still seem to have an advantage, but not nearly as much as one might think at first glance. And since all else is rarely equal in real life, other factors would come into play. For example if “peacocks” can reproduce faster than “penguins” under good conditions, the place might become swamped with “peacocks” while the “penguins” only have a foothold.

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