When I was in graduate school at Duke University in the early 1980s I remember a young professor visiting from Michigan State who gave a talk about group selection. The professor was of course David Sloan Wilson, because at the time he was the only academic willing to stick his neck out for group selection. I thought his ideas were eminently sensible and was surprised to learn that none of the other graduate students agreed with me. Group selection was not particularly close to my research interests then (I was working on population movements of insects), but I kept following it. What I heard during the 1980s and 1990s was a relentless drumbeat against group selection. As a result of books by such luminaries as G. C. Williams and Richard Dawkins, evolutionary biology at the time thoroughly repudiated this idea.
Later, during the 1990s I became interested in scientific study of history (so much that I eventually switched from biology to historical social science) and discovered that the theory of group selection, which by then became the theory of multilevel selection, could provide a very productive conceptual framework for the study of the evolution of complex societies. It turned out that multilevel selection yielded predictions that could actually be tested with historical data. What was particularly exciting was that theoretical predictions, time and again, yielded novel insights that were, amazingly enough, supported empirically (when I started working on historical dynamics I did not realize how much data there are to test different theories).
Meanwhile, there was a glacial, but also tectonic shift (sorry about mixing metaphors) in social evolution, especially in human social evolution. Gradually an increasing number of researchers came over to the view that cultural and genetic group selection provides a very viable theoretical framework for the study of evolution of human sociality. Colleagues like Pete Richerson and Rob Boyd, Sam Bowles and Herb Gintis, Elliott Sober, Joe Henrich, and many others became seriously engaged with models and empirical analyses framed within the multilevel selection paradigm.
The signal event in this tectonic shift was the “defection” of E. O. Wilson to the group selection side in 2007. The two Wilsons co-authored a Quarterly Review in which they proposed the dictum, “Selfishness beats altruism within groups. Altruistic groups beat selfish groups. Everything else is commentary.”
A month ago we ran a workshop at NIMBioS (the National Institute for Mathematical and Biological Synthesis; I will be reporting on it later this week). The workshop was attended by a truly multidisciplinary group of more than 40 scientists. What was really surprising was that the idea of multilevel selection was something all of us could agree upon.
I believe that we are in a Kuhnian paradigm shift, and I fully expect that multilevel selection will become the reigning paradigm in the next 5-10 years. But the transition is not going to happen without pain.
This paradigm shift is associated with a remarkable degree of acrimony (although I did not experience at first hand other paradigm shifts, so perhaps they all are similar in this respect). The case in hand is the exchange over the weekend between David Wilson and Jerry Coyne. It started with a post by Wilson When Richard Dawkins is not an Evolutionist. Coyne shot back with David Sloan Wilson loses it again, and Wilson blasted away with Pugilistic Science.
In the process, Wilson intimated that Dawkins is not really a scientist, while Coyne suggested that “Wilson is totally over the waterfall.” At the end of his post, Coyne added: “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.” Now I don’t want to paint Jerry as a blackguard, all parties of this debate have transgressed over the norms of polite discourse, and in a different age this could easily lead to a dawn encounter with matched weapons.
But let’s return to the substantive issue, that of group, or better multilevel selection. Specifically, not whether it is prevalent in the animal world, but its role in the human social evolution. I am of course a partisan in this debate, but I don’t understand the extreme position taken by Richard Dawkins and Jerry Coyne, who deny any role for multilevel selection. If we want to understand the evolution of large-scale complex societies, what is the alternative? How could human ultrasociality, our ability to cooperate in huge groups of genetically unrelated individuals, evolve? The two theories discussed in The Selfish Gene, reciprocity and kin selection, fail utterly on both conceptual and empirical grounds, as has been abundantly demonstrated by the likes of Richerson and Boyd.
To make this question even more specific, we know that humans are capable of sacrificing their lives for the sake of huge groups consisting of unrelated strangers. As an example, think of the Southerners and Northerners volunteering for the Confederate and Union armies during the American Civil War. It is a particularly well-documented case, because those volunteers were literate, and they left behind thousands of letters explaining their motivations. Hundreds of thousands of them died on the Civil War battlefields. So how can you possibly explain such remarkable properties of human sociality, other than with a group selectionist model?
This is not a rhetorical question – I genuinely would like to see what theoretical alternatives there are, so that we can start figuring out how to test them empirically. So let’s hear them.
You raise just one of the many features of human behavior that are readily explicable via multi-level selection but inexplicable via “gene” only selection. My view of Dawkins is that what he has produced is either obvious/definitional or wrong. But in his efforts to address your question I guess he would have to throw his meme idea at the wall and hope it sticks. So the men dying in the Civil War were victims of “memes” that hijacked their brains in some unknown way in order to “replicate,” or something, somehow. That is about as precise as Dawkins and the meme camp get about it.
Then looking at Political Science and International Relations there are various notions assigning agency to “the state,” which apparently has “interests,” or ascribing causation to various constructs such as Capitalism, the interstate system, Patriarchy, etc. The notable thing about all of these notions is that they are absurd, and yet they dominate the discipline to the exclusion of all others.
Peace Studies briefly summarized asserts that war mostly is about fear of the unknown “other” (so each combatant is motivated not for the group but for their own safety) and if only we get to know each other better we will be peaceful. Of course, brothers fought on opposite sides in the Civil War.
Of course, psychology has plenty of wild ideas about why men go to war as well. Staub comes to mind as a representative sample, asserting that in order for men not to wage war we need to have “love and affection…constructive fulfillment of basic human needs; humanizing the other…self-awareness…healing from past victimization…” Of course, we must also toss in “Helping young people who come from difficult backgrounds or who have had painful experiences…”
Carmi,
Yes, “memes made me do it” is one possible response from Richard Dawkins and his followers. It’s a variant of what might be called the “great deception” hypothesis, e.g. Marxist explanations how people are fooled to fight and die for the sake of the ruling class’ interests. The problem is that fighting in war has very significant fitness consequences (roughly one-quarter of male deaths in small-scale societies is due to warfare). So by the “selfish gene” logic such fierce selection should result in the evolution of very effective resistance to such lethal memes.
Peter, while I agree with you, I think it is a little more complex. A Dawkins meme counter would be that the increased fitness of the males that survive the war (via greater access to females) and propagate the meme allows for positive selection. Further, if the meme includes severe punishment of those males who do not fight in the war (as is common), then the relative fitness calculations again must be redone. On the other hand, this is all fancy dancing since these same arguments can be made without using memes at all.
Put more simply, the average fitness of the men going to war only changes if women die or territory is lost. Otherwise, even if 80% of the men die the average fitness of the men of the group remains essentially constant.
I elected to respond to the “memes made me do it” point on my own blog. The article is titled: “Necrotrophic memes”. However, I’m not sure if posting offsite links here is cool.
I find Richard D. Alexander’s approach helpful. Much of the debate is semantic. Humans compete via coalitions, and likely have done so with increasing significance for much of our evolutionary history. Our neurobiology provides some interesting clues.
Alexander RD (1987) Biology of moral systems. Hawthorne, NY: Aldine Press.
Alexander RD (2006) The challenge of human social behavior. Evolutionary Psychology, 4(2), 1-28.
De Dreu CKW, Greer LL, Handgraaf MJJ, Shalvi S, Van Kleef GA, Baas M, Ten Velden FS, Van Dijk E, & Feith SWW (2010). The neuropeptide oxytocin regulates parochial altruism in intergroup conflict among humans. Science, 328 (5984), 1408-1411.
Flinn MV, Geary DC, & Ward CV (2005). Ecological dominance, social competition, and coalitionary arms races: Why humans evolved extraordinary intelligence. Evolution and Human Behavior, 26, 10-46.
Flinn MV, Ponzi D, & Muehlenbein MP (2012). Hormonal mechanisms for regulation of aggression in human coalitions. Human Nature, 22(1), DOI 10.1007/s12110-012-9135-y
I don’t disagree with this, but semantics are important. It is important to use words carefully, especially in multi-disciplinary settings (such as social evolution) that bring specialists from wildly different disciplines, each with its own language. So evolutionary theorists may know that models of kin selection and group selection are mathematically equivalent, but it is extremely unhelpful to tell sociologists, for example, that kin selection explains the evolution of macrosocieties consisting of millions of individuals, in which average degree of relatedness is indistinguishable from 0.
If individuals within groups really are unrelated, then group selection models don’t work either. Group selection relies on the individual being more like members of the group than like a randomly-chosen member of the species. Such relationships are normally expressed in terms of relatedness. However, note that average relatedness is not what matters. The population may well contain two brothers, or eight cousins – even if the average relatedness is zero. What matters is more like “total relatedness”. If the total relatedness is “indistinguishable from 0”, then group selection models will fail as well – for the same reason that kin selection models do.
I am not sure what you mean by: “reciprocity and kin selection, fail utterly”. These are, in fact, widely accepted theories. You mean that they don’t account for ultrasociality? That’s because there are also other causes of ultrasociality – including manipulation (e.g. see “parental manipulation”) and (in humans) effects to do with culture.
People going to war often don’t think they are going to die. Their reasons for going vary from obeying authority to earning money. Sometimes they are manipulated into it. Sometimes they miscalculate the risks and benefits involved. Sometimes their brains are hijacked by cultural symbionts that use fictive kin effects to induce patriotic fervour. In short, there isn’t just one reason.
Human ultrasociality is largely the product of culture – as can be seen in the difference in group size between hunter-gatherer tribes and agriculture-capable communities. The primary explanation for this can be found in my “memes and the evolution of human ultrasociality” article. Perhaps check it out.
The group selection controversy is partly a misunderstanding over terminology. The group selection bashed by Dawkins and Maynard Smith and Williams excluded kin selection. The group selection promoted more recently by Wilson and Wilson includes kin selection (and lots of other things). The same name – but radically different theories. The first kind of group selection is still mostly wrong – but it is important not to muddle it up with the second kind of group selection. If you have 20 minutes, perhaps try watching “Group selection debate – Stuart West” for how and why group selection lost out historically.
Regarding cultural influences: there are quite a few lethal diseases out there. Fierce selection against them is hindered by the fact that they are not that common – and the fact that the diseases have a shorter generation time and evolve more quickly than their hosts.
FWIW, Richard Dawkins does not “deny any role for multilevel selection”. Multi-level selection applies practically any form of selection acting on individuals in groups (as is explained in “Unto others”) – and thus includes kin selection and reciprocal altruism – which Dawkins regards as being important. What he apparently has a problem with is the rechristening of the term “group selection”:
“The so-called “new group selection” is just kin selection or in some cases reciprocal altruism under another name. For reasons best known to himself (which I can’t understand) D. S. Wilson thinks it’s helpful to rephrase it in terms of group selection. How it can be helpful when he’s reviving a word which has been debunked and is simply grafting that word onto the very thing that did the debunking – namely kin selection and reciprocal altruism and various other things – it seems to me be to be utterly unhelpful, to be totally misleading to students and it’s deeply regrettable that E. O. Wilson should have teamed up with him in this way.” – Richard Dawkins.
Thanks for this quote – it’s very representative of what I had in mind when writing the post. Just to clarify things, kin selection and reciprocal altruism fail in explaining the evolution of cooperation in large-scale human societies (millions and more people); for other questions they could be quite frutiful approaches.
Most multi-level selection models can be “rexpressed” in terms of inclusive fitness. E.g. see: “Quantitative genetics, inclusive fitness, and group selection” by Queller (1992) in “American Naturalist”.
Instead of saying that genes in altruists within groups spread because they are present in other group members, an inclusive fitness approach would say that they spread because other group members are more closely related to them than individuals randomly sampled from the entire species. It’s another way of looking at things that has been much more popular historically – since inclusive fitness also explains phenomena such as parental care and nepotism. Since evolutionary biologists had already adopted inclusive fitness, group selection was seen as adding little.
An inclusive fitness approach becomes more obviously pertinent when considering small tribes and the EEA – where the chance of someone you randomly encounter being a relative are higher. That’s where our cooperative instincts were forged.
Kin selection and reciprocal altruism are actually very important theories when it comes to explaining human ultrasociality. Human social instincts mostly evolved in small and more inbred tribes – where kin relationships were common, and cooperation was mostly either with kin or those you would have repeated reciprocal interactions with. The exact same instincts that caused cooperation in the past now cause with strangers in the modern environment – even in cases where such cooperation is maladaptive. Humans account poorly for group size – because they were only ever in small groups historically. These ideas do not “fail utterly on both conceptual and empirical grounds” – though they aren’t the sole cause of human ultrasociality (there’s also “virtue signalling” – and several other effects).
Kin selection, for all its appeals, does not fit the facts on the ground when it comes to human sociality. Humans are joiners, and the huge number of experiments in minimal group dynamics done by social psychology show that our brains automatically have positive bias towards in-group, even when expressly grouped together as the group that does not like each other. Mathematically, kinship fails in groups larger than forty individuals. The R values are too low. Humans evolved to live in groups larger than forty. The obvious fact that humans define in-group from out-group culturally far more than by kinship also indicates the weakness of kin selection. Finally, there are a large number of historical and anthropological examples from disparate times and cultures where brothers fought each other on opposite sides of a war.
Either way, the ultimate “cause” of Dawkins disagreeing with group selection is that he has failed to understand evolution. One of the most basic assumptions/assertions of the Theory of Evolution is that competition for resources generates selective pressure. Humans clearly compete for resources at the group level as well as at the individual level. Competition for resources MUST generate selective pressure at whatever level it exists at, or the entire Theory of Evolution is wrong and must be rejected. Dawkins must either show how/why competition at the group level does not result in selective pressure at the group level or finally give up his stupid ideas. The above is true regardless of kinship or reciprocity or altruism. It really is this simple.
You can’t say that kinship models fail because relatedness is too low – and that the answer is group selection. The “new” group selection and inclusive fitness are broadly-equivalent theories. To quote from Queller (1992):
“The inclusive-fitness and group-selection models presented here do not represent alternative processes. Nor is one a subset of the other. Instead, they are simply alternative ways of viewing the same selection process.”
Humans defining in-group from out-group culturally more than by kinship does NOT indicate a weakness of kin selection models – any more than it implies a weakness of group selection models. It just means you need to apply the concept of cultural kin selection – kin selection acting on memes – not on genes. Shared military uniforms indicate shared memes. The memes are prepared to sacrifice themselves for copies of themselves in other bodies – and they manipulate their hosts to achieve that end, using the “fictive kin” phenomenon – to fool their hosts into believing that their fellows are their genetic kin – not just their memetic kin.
Tim, I am sorry I was not more explicit above. I am well aware of the standard “new” group selection as per the two Wilsons and others. I am disagreeing here and stating that none of that complex math about groups of altruists vs groups of selfish is required, nor is there any need to worry about how closely related we are. If my ability to acquire fitness limiting resources is partially or entirely controlled by my groups ability to control or gain access to those resources, then there is evolutionary selective pressures at the group level; my fitness as an individual is partially or entirely determined by the relative fitness of my group regardless of memes or kin or even reciprocity. This is true, or one of the most basic foundations of the Theory of Evolution is false.
Now, I know that the dominant paradigm here is that if the above was true then selfish would wipe out altruists within the group and blah blah blah. Wrong again, for two reasons. First, we need not assume naive altruists who are defenseless against selfish exploitation, both unable to detect it and unable to act against the selfish once they have been detected. We do not have to equate altruist with sucker, because in fact the evidence is ample that when altruists figure they are being made suckers they act quite decisively to become selfish or to rid themselves of the selfish, whichever is most to their advantage. In general, along this line of reasoning, anyone from a small town would be shocked at the Game Theory assumptions that selfish could blithely defect and get away with their reward. Privacy and secrecy are very rare in small groups living closely together. Remember also that our ancestors did not have a lot of choices about which group to live with; if you were in a lifeboat lost at sea, would you risk pissing off everyone else in the lifeboat? Our ancestors were no less dependent on their group for their survival.
Second, until just recently everyone but myself seemed to miss the fact that selfish are in fact selfish. The selfish who rids themselves of other selfish within the group of altruists will increase their fitness, and provide themselves with the sheepskin to cover their wolf natures. This is the fundamental process we call “politics” these days. Selfish competing for the ability to exploit the altruists. Because of the large sizes of human groups, kinship amongst selfish is too low for them to continue to cooperate with each other based on it well before they reach majority size in a group of 150. So altruists have an interest in getting rid of selfish, and selfish have a fitness interest in getting rid of selfish other than themselves and their close kin. Selfish punishment of selfish ends the infinite regression issue for enforcement of altruism. The worst outcome for selfish is that a majority of the group decides they are being made suckers and defect to the selfish side themselves.
As for your assertions about memes having “kin,” this is really stretching an absurdly bad idea to incredible lengths. The whole meme idea is readily disproven by playing the preschool game known as “telephone.” Did the meme manage to “replicate” more than a few times? Nope. Is the gibberish the fifteenth person says a “kin” of the original meme? Hardly. Memes cannot “sacrifice themselves,” they have no selves. Memes have no fitness or replication either.
You reject the idea of cultural kin selection – because of your unfamiliarity with, or dislike of the meme terminology? That seems pretty ridiculous to me.
The whole business where people think that kin selection doesn’t apply to large groups of humans – and therefore we need to invoke group selection is just down to ignorance of how cultural evolution works. Kin selection and group selection are broadly equivalent theories. Each applies to the organic realm of DNA, and the cultural realm. We have organic eusociality – with a sterile queen and lots of workers. And we have cultural eusociality with a fertile bank and lots of uncopyable bank notes. Altruism genes sacrifice for copies of themselves in relatives, and patriotism memes sacrifice themselves for copies of themselves in other humans. One could give many other examples. It is a different domain, but the exact same logic applies – since both are Darwinain systems.
The whole Meme notion actually, and interestingly, reduces to Standard Social Science Model when you think it through. Why do humans go off to war? “Memes”/”Culture” did it. What is the difference between the Memes assertion and the Culture assertion? Just some argumentation trappings and gibberish about “selection” from the Meme folks.
Lots of these ideas can seem to make sense when one is an academic with a highly limited imagination and not much knowledge. However, the reality is that all group social territorial species, possibly excepting Bonobos, engage in group level violence. Humans are not the exception at all, despite that idea being repeated by thousands of “scholars.” It remains vaguely possible that Memes/Culture are to blame, but given similar behaviors exhibited in lions, meerkats, many species of monkeys, ants, wild dogs, hyenas, chimps…I think the first place to look would be within our evolved behaviors.
The whole Memes idea is just a desperate attempt to explain what cannot be explained otherwise by those who maintain their religious belief that there is no group level selection. Just as SSSM folks who deny that our behavior evolved are forced to assert “Culture” as a one size fits all “answer,” people who deny group level selection are forced to assert “Memes” as their version.
But think about it just a little. Culture is highly transient in our society, changing quite rapidly these days. But we can name epochs of our ancestors based on their stone tool making methods. Changes happen very rarely; hundreds of thousands of years pass without any noticeable changes. Where is the “Meme” or “Cultural” selection pressure? Everyone had pretty much the same culture as their neighbors had. (http://phys.org/news/2013-03-ancient-stone-tools-pace-remarkable.html) Differences in culture or memes that would make a difference in group fitness were exceedingly rare.
Another important point is that we can distinguish between what is cultural/meme based and what is biologically based by noting if something is universal. Cultures and memes are highly malleable especially in large societies. Yet things like “patriotism” which you suggest as a meme are in fact universal and they have components that are sub-conscious and automatic (positive bias for in-group even in extreme minimal group conditions). Of course, the empirical evidence only makes sense if we accept group level selection…
It is not that I am unfamiliar with memes or simply dislike them. I have thought the idea of meme selection through and see why it is wrong.
Levels of violence are rather obviously affected by both DNA and memes. The contribution of human DNA is evident in the widespread nature of violence the contribution of culture is evident in the decline of violence over time – as ably documented by Pinker in “The Better Angels of Our Nature”.
Cultural differences that contribute to group fitnesses are well known and well documented. If you look at “Guns Germs and Steel”, the Guns and Steel are cultural. Their historical effect on group fitnesses has been clear and dramatic.
Patriotism has a well-known cultural component. Governments actively attempt to influence their populations using culture to encourage patriotism and nationalism. Then threatened by aggressive neighbors they invest more in primary education, to influence their citizens when they are more impressionable. Patriotism memes are well-known – just think of “Your Country Needs You”, for example.
Just because something is universal, that doesn’t mean it doesn’t have a culturally-transmitted component. Language is universal, but that doesn’t mean that Japanese is universal. Japanese has many culturally-transmitted local quirks.
You argue that much culture is universal, cite the S.S.S.M. straw man approvingly and counter the “memes did it” idea with the idea that genes did it. That profile fits many who identify as being in the “evolutionary psychology” camp. While those folk are justifiably contemptuous of many of the social scientists who preceded them, those social scientists got one thing right that the evolutionary psychologists failed to notice – namely they were impressed by the extent and influence of human culture. Culture is not the answer to everything. It is a bit less significant than genes on the average phenotypic trait. However, memetics treats it seriously as its own evolutionary process – in line with what is currently known about cultural evolution – while evolutionary psychology has historically ignored and shunned culture in a manner that looks pretty ridiculous to those who understand its significance. You seem at risk of making this mistake.
“Culture did it” simply can’t be dismissed as an unscientific hypothesis that explains everything and anything – a la Tooby and Cosmides. Culture really *did* do it – maybe about 30% of the time. You often have to look at evidence such as transmission patterns to see whether the hypothesis is true or not.
Multi-level selection – as embodied by the Price equation – has surely been orthodoxy for decades now. I tend to regard the modern enthusiasm for cultural group selection partly as a Wynn-Edwards-style “bubble” – in the cultural realm. People are invoking cultural group selection without even properly considering cultural kin selection – despite the fact that inclusive fitness theory has been massively more successful in the organic domain. Essentially people are making much the same mistakes as we saw in the history of organic evolution – and they are doing so partly because the parallels between cultural and organic realms are not being properly appreciated.
Regarding the use of the term “group selection” by Richerson, Boyd, Bowles, Gintis, Sober and Henrich – that’s a rather incestuous group. Boyd and Richerson at least recognise the equivalence of inclusive fitness and group selection – but apparently prefer the “group selection” term – despite the fact that this is historically mired in confusion and misunderstanding. Practically the whole of the rest of the evolutionary biology establishment prefers to use the “inclusive fitness” term. It seems to be a straightforwards scrap over terminology. The “inclusive fitness” crowd looks a lot like it won the fight back in the 1980s to me. Maybe there will be some rear-guard action late in the day – but it doesn’t seem terribly likely. It also seems to me that the “cultural group selection” crowd have been led badly astray by their own framework. Where is their cultural parental care? Where is their cultural kin recognition? Why concentrate on group level effects – while ignoring the enormous significance of close cultural kin? It doesn’t seem to make any sense – scientifically speaking.
I’m rather sceptical about the possibility of a “tectonic” paradigm shift in this area. My reasons are: there’s no significant new science – the issue is about whether to describe a theory using the terms “inclusive fitness” or “group selection” – and scientists have been using the “inclusive fitness” terminology for decades now. What might happen is that the reputation of the “new” group selection could improve – as increased numbers of people understand its relationship with and equivalence to the mainstream inclusive fitness theory. However, that will just be business as normal for most evolutionary biologists who use the inclusive fitness theory – not really much of a revolution.
Carmi,
1. “even if 80% of the men die the average fitness of the men of the group remains essentially constant” – but the point is that if you are a free rider, you are going to be one of the surviving 20 percent.
2. When you invoke greater access to females or punishment of those who do not fight, you have to deal with the problem of second-order public goods. Why should females evolve a preference for mating with heroes?
3. The broader point here is that the evolution of cooperative traits is affected by a multitide of competing influences. When you deal with such complex nonlinear dynamics you have to use mathematical models. If you want to understand whether fighting for the group and punishing shirkers can evolve together, you need to model coevolution of these processes. Fortunately, this ground has been well plowed over the last decade or two, so now we have a fairly mature body of theory that tells us what works and what doesn’t. See, for example, the recent book by Bowles and Gintis for an overview. My statement that kin selection and reciprocal altruism fail to explain the evolution of human sociality is based on this body of theoretical results. In particular, reciprocal altruismm breaks down as soon as you get beyond tiny groups of of a few individuals, while kin selection breaks down in human groups of a few dozen people (as you say elsewhere).
“Virtue signalling” doesn’t break down in large groups – as reputation-based systems like eBay and Amazon abundantlly demonstrate.
“Manipulation” doesn’t break down in large groups either. When the king orders, his generals respond. When the general orders, his troops respond. No need to explain the benefits of dying in battle if there *are* no benefits – and soldiers are working for someone else’s benefit. Then you still need to explain why humans respect authority – but that seems to be a much easier task.
Inclusive fitness theory is isomorphic to multi-level selection theory – as you seem to agree. You can’t consistently claim that kin selection breaks down beyond a few dozen people while multi-level selection works – since these are the same theory in different outfits. Relatedness has a “long tail” – where much of your group is only distantly related to you – BUT there are a large number of individuals involved. The fact that *many* distant relatives are involved is why interdemic selection has any chance of working at all. However – as you seem to agree – the effects on close relatives seem to be dominant. We can empirically see adaptations for behaving altruistically towards close relatives – while adaptations for dealing altruistically towards distant group members remain unproven.
Peter,
1) I think we often lose sight of the realities here. It is pretty hard to free ride when the tribe is fielding twenty or thirty men in battle. It is possible, and I remember one account from anthropology where a man panicked and failed to aid his friends who were ambushed. He was after that labelled a “ghost” in his tribe, a non-person, a person who had no value or consequence. It is no accident that initiation rituals often test the courage and ability to endure pain of the males.
2)Bravery is an honest signal of fitness, as much so in humans as in fish. Female fish of a certain species (forgive my early morning brain for not remembering the species) prefer males who show their bravery because it demonstrates fitness in an honest way. So too with humans.
3)For all their complexity, I feel that the models fail to include the role of selfish in encouraging cooperation. Even if I will free ride, it is in my interest to make sure no one else does. The margins between victory and defeat are often razor-thin, and my own wife and kids and my own hide depend on the group winning. At times I may even judge that free-riding is not in my own best interest either; better the risk of death than the certainty of it when my group is overrun.
Tim,
1. It’s entirely appropriate to provide an offsite link or two – after all, one of the goals of this site is to connect together people interested in social evolution. Just be parsimonious, because the spam-catching software will kill popsts with too many links.
2. I am not sure what “new group selection” is. I prefer multilevel selection.
3. All people working within the multilevel selection framework (at least, the ones that I explicitly named) recognize that mathematical models of inclusive fitness and group selection are equivalent at an abstract enough level. But your approach of simply sweeping all mechanisms into the kin-selection bin is not helpful at all. “Cultural kin”?
The general insight from a multitude of models is that in order for cooperative traits to spread, carriers of such traits should interact with each other more frequently than with carriers of noncooperative traits. Such nonrandom assortment may arise as a result of relatives cooperating within kin groups, or cooperative individuals building coalitions, or by the “Green Beard” effect, etc.
4. Finally, I am much more optimistic about the future of multilevel selection theory than you are. Within the field of human social evolution, multilevel selection seems to be the most productive research program. It’s actually driving empirical research (which cannot be said about other frameworks) – Sam Bowles and coworkers on genetic group selection, Richerson and Henrich on cultural group selection, Ara Norenzayan on evolution of religion, Joh Haidt within social psychology, and my own work on the evolution of large-scale societies. In my work I test predictions derived from the Price equation with data on temporal and spatial patterns in the rise of historical empires. See my cliodynamics.info site for refs.
Just as individuals tend to behave altruistically to those with shared genes, so they tend to behave altruistically to those with shared memes. We see that in police, military and religious groups. Shared memes are also why your computer cooperates with your printer. Inclusive fitness is a general aspect of Darwinian evolution. It is not confined to genes. It also applies in the cultural realm.
Multilevel selection theory is the same theory as kin selection. As we both keep repeating, these are broadly-equivalent frameworks. The issue is what to call it – and which set of models to use. Most evolutionary biologists (though not the handful you mention) are in the “kin selection/inclusive fitness” camp. If you are interested in understanding why that is, a good paper is: S. A. WEST, A. S. GRIFFIN & A. GARDNER (2009) “Social semantics: how useful has group selection been?” – http://www.zoo.ox.ac.uk/group/west/pdf/WestGriffinGardner_08.pdf
Multi-level selection may be driving empirical research – but to me it seems to be causing no end of problems. It is producing a lot of research based on interdemic selection – and practically none based on cultural kin selection among close kin. We already know from the organic realm that kin selection acting on close relatives is very important – while interdemic selection hasn’t been proved to be responsible for much at all – outside of the laboratory. That appears to be a massive and terrible research bias – from my perspective. If that is the effect of using group selection, social scientists should consider dropping the approach like a hot brick – before it causes them even more problems.
At the risk of stating the obvious, it really matters whether deaths in battle are glorious sacrifies for the good of the group – or represent the soldiers being psychologically manipulated against their own best interests by nationalistic propaganda into giving their lives for people who have been dressed up to look like their brothers, generals well behind the front lines – and stay-at-home fat cats in the government, who they really shouldn’t be caring two hoots about.
In the latter case there would seem to be more chance that warfare will be prevented once soldiers are better educated and have more chance of understanding the personal tragedy that they are getting involved in. The picture inspired by group selection is not just scientifically highly-dubious, it is research that could be used by unscrupulous governments to help manipulate soldiers into even deeper fighting frenzies.
I don’t think there can be much doubt that dying in battle is bad for your genes. However, what the deaths of soldiers *are* adaptive for is the nationalism memeplex. That exists in the form of other copies which directly benefit as a result of the sacrifices of the soldiers – and so the memeplex spreads. The instance of nationalism memeplex in the soldiers dies – but *copies* of that memeplex in the generals and politicians survive – and so nationalism spreads.
Seen from this perspective, the triumph of nationalism can be seen as the result of a form of cultural kin selection. Some of the nationalism memeplexes sacrifice themselves (and their hosts) for *copies* of themselves in *their* immediate relatives.
If you really, *really* want, cultural kin selection can be rephrased in terms of cultural group selection – since these are equivalent theories – but be warned, decades of muddle and confusion stem from attempts to apply group selection, while inclusive fitness is a well understood and uncontroversial theory which is widely accepted by practically all evolutionary biologists.
I note that Gordon Rakita (2009) gives the same sort of explanation that I gave above – in “Ancestors and Elites” – starting out by saying: “Kin selection does not seem to offer a suitable explanation for instances in which soldiers give their lives for their comrades. The conditions necessary for the model are not met, for example, soldiers are rarely genetically related. Given this, how are such acts to be explained?” The explanation given does not involve group level benefits – but rather benefits to immediate cultural relatives.
Hi, Everyone
The truth is I don´t really understand the controversy between selfish dynamics and the multilevel selection approach. It seems obvious to me that they are somehow linked: Selfish replicator selection leads to multilevel selection.
I think the problem is Dawkins´s atheist activism. When I started to read about darwinian cultural evolution I immediately saw that it explained religion as an adaptation at the group-selection level. Dawkins must have seen this too (along with D.S Wilson who wrote “Darwin´s cathedral”), and he must not have liked it. Therefore he has rejected it in a way which seems almost “religious”.
In the case of men going to war. It is not only a parasite meme which “manipulates” the individual minds of the soldiers to go willingly to war. The reason why most people go to war is because “they have to”. They will be punished if they don´t go. Muhammad Ali didn´t have a parasitized mind in this regard, and he paid for it: he spent 4 years in jail. Group selection in the cultural human level has led to the creation of institutions (memetic group organs) which have power over the individuals. This cultural constructs inforce their rules by delivering punishment. This is the mechanismo that Boyd aln Richerson called the “moralistic punishment”. This is the base for all human societies. Our co-evolved genes have made us conformists because it is the best way to thrive in a society with social rules and institutions, in a context of intergroup competition.
Both theories: the evolutionary psychology Big Mistake hypothesis and the darwinian cultural evolution theory are both correct and complementary in order to explain ultrasociality and group-selection in humans. In this case, memes (mostly parental) which have evolved thanks to intergroup competition “parasitize” and maniplulate the genes for kin altruism and reciprocal altruism. This would make a “memeticly guided” Big Mistake hypothesis.
Besides, whenever the neolythic arrived, humans started to live in a neatly marked territory, a source of sources, whose populations had it easier to achieve at least a certain degree of cultural (and genetic) relatedness. Few animals have territories as well defined as humans. The only ones I can think of are social insects. Bees have beehives and humans have towns and cities. Has this been taken into account?
Juan Alfonso del Busto
Juan, the problem with the work of Boyd and Richerson and with all this meme stuff is that it puts the cart well before the evolution of the horse. “This cultural constructs inforce their rules by delivering punishment” – here again we have, as I described in an earlier post, magical constructs “acting” without corporal bodies. How can this be described as how it always works? Why is it the case that “Our co-evolved genes have made us conformists because it is the best way to thrive in a society with social rules and institutions, in a context of intergroup competition.(?)” The fact is that one can readily have culture without conformity. Why should any of your claim above be true? We need to really examine the ultimate selective forces here.
What is the role of culture in human society and evolution? Critically, we define our in-group and exclude our out-group using culture. Why? Because it allows for larger groups to maintain solidarity and cohesion, and larger groups have a huge fitness advantage over smaller groups, like those which are limited by bonds of kinship. Why conformity? Well, obviously if we are defining the group via cultural means those who do not conform to that culture threaten group unity; either a schism or complete collapse of group cohesion can result. So non-conformity threatens to lower group fitness, and we all have an interest in keeping group fitness high. So it is not some random meme or the fact that “it is the best way to thrive in society,” rather it is because of group level selection that we have enforcement of cultural conformity in all human societies and groups. Importantly, in order for groups defined by culture to begin to exist, we had to have enforcement of conformity already in place. It is logically impossible for the “meme” or specific “culture” to create the genetics of conformity after they must have already existed.
Culture also helps humans deal with their environment (think clothes and weapons) and display their fine qualities to mates (think songs, humor). It isn’t just about marking group boundaries. Indeed group boundaries often follow cultural markers because similar memes clump together – into species groups of memes and families of memes. It’s rather like the way fish of the same species school together, or lions of the same family hunt together.
Re: “When I started to read about darwinian cultural evolution I immediately saw that it explained religion as an adaptation at the group-selection level. Dawkins must have seen this too (along with D.S Wilson who wrote “Darwin´s cathedral”), and he must not have liked it. Therefore he has rejected it in a way which seems almost “religious”.”
Dawkins offered cultural evolution as an explanation for religion long before “Darwin´s Cathedral” came out. For example, see “Virus of the mind” (1993) and “Memes: the new replicators”, 1976.
He mainly uses kin selection and reciprocity – not group selection – to explain religious co-operation. Since the “new” group selection is basically a synonym for kin selection, this isn’t such a big difference.
“He mainly uses kin selection and reciprocity – not group selection – to explain religious co-operation. Since the “new” group selection is basically a synonym for kin selection, this isn’t such a big difference.”
I know this is a popular dogma right now, but it is also very wrong. Human history is chock full of examples of group over kin, and there are anthropological examples as well. When forced to choose between loyalty to group or loyalty to kin, kinship mostly looses. In war, kin kill each other. In cults, religion wins over kin. In Rwanda, parents killed their own “mixed-group” children. In China and Russia and Nazi Germany, children turned in their own parents and siblings. In the Nika revolt in Byzantium, the contemporary historian Procopius tells us that kin killed kin over which chariot team they cheered for.
There is just no way to square the kin selection story with the actual empirical evidence of human behavior. We define groups culturally, not based on kinship, and have done so for quite a long time apparently. Kin selection is oh-so appealing and neat and tidy, but it is also wrong, not supported by the evidence especially in humans, and very much not the same thing as group selection in MLS.
Carmi, this is a very good point. To your list of examples, I would add fraternal conflict and homicide. History is full of examples when brothers fight civil wars with each other over who will inherit the kingdom. In some extreme cases, as in Ottoman Turkey, the winner had to personally kill his brothers. Group wins over kin.
Re: “History is full of examples when brothers fight civil wars with each other over who will inherit the kingdom. In some extreme cases, as in Ottoman Turkey, the winner had to personally kill his brothers. Group wins over kin.”
Peter, that’s argument by anecdote. Kin selection theory doesn’t claim that relatives cooperate on every single occasion. Indeed, the theory of “local competition” suggests that relatives that fail to disperse frequency come into conflicts over resources. Human history is full of cases where relatives kill each other – but so what? That isn’t a case of group selection and kin selection making different predictions. There have been many attempts to find such cases – and the efforts have consistently ended in failure. If you think you have found such a case by all means look into the literature on the topic – but history offers such efforts little encouragement.
Er, cultural kinship is not in conflict with kin selection theory – since kin selection applies to memes – as well as genes.
Have you looked at the literature on the equivalence of kin selection and MLS? Most of the MLS supporters endorse it these days. In particular S. Okasha, D. S. Wilson, R. Boyd, P. Richerson, M. Wade and H. Gintis are now supporters of the concept. It’s the consensus view.
To define our in-group and exclude our out-group is ONE of the roles of Culture, not THE role of Culture. And this role is not by any means a “default” function of Culture; It is an evolved adaptation through group level selection, and one which “coadyuvates” group-selection (a loop). I don´t see anything “magical” about this. Is a neurological module, which allegedly account for a given behaviour or emotion, a magical construct?… I don´t think so… We know quite a lot about DNA, but we don´t know much about genes (real genes, the kind of genes which account for a behaviour or an emotion, not the fragments of DNA between two codons). All right, we know a lot about the mendelian laws of gene transmission. Impressive. Culture has much more complicated laws of transmission. True. So what. It doesn´t make it “magical”. It makes Culture complex. And Culture and institutions HAVE bodies: ours. Police officers with clubs and guns are not magical.
It happens the same with conformity. It is a group level adaptation. And group level selection in humans has a lot to do with gene-culture co-evolution. Whenever advanced imitation skills allowed cummulative Culture in a population, genes and Culture started to interact and co-evolve in a kind of positive feed-back loop of niche construction dynamics. Our genes (and instincts) started to adapt, through selection, to Culture. And vice-versa. Both replicators, genes and memes, have taken advantage of one another. Conformity bias, prestige bias, etc, are the result. They enhance in-group homogeneity and out-group heterogeneity, and therefore, group level selection. All groups as a whole, (as a “population” of groups) benefit from this, because another level of competition and selection (group selection) arises and makes it easier for all of them (as a population of populations) to add complexity to the solutions of old problems. With this, adaptation to harsh and diverse environments suddenly becomes possible. Since cultural evolution can be quite fast, it is logical to think that genetic adaptations have to catch up. On the other hand Culture evolves in the frame of previous genetic adaptations and instincts. Therefore Culture is somehow guided by biology. Memes are hardly ever random (their variation and evolution is). Well, at least parentally transmited memes are hardly ever random.
Of course we could have culture without conformity. For instance, if the environment favoured a kind of rapidly changing selfish culture, Culture would adapt (through multi-level selection) and conformity would vanish along with in-group detecting abilities as our genes would slowly adapt too, co-evolving with both the new environment and the new culture. Of course this would mean that Culture would lose a lot of the features that are most important (I think that Culture would resemble that of 100.000 years ago: very few innovations).
In my opinion the most important milestone achieved by Culture evolution is the creation of social conventions, norms and institutions which inforce these norms through punishment. This has set the base for a common trust atmosphere that has allowed societies to try new ways of specialization (specially since the neolythic with its permanent settlements) and put these new ways to test through intergroup tournament. This way institutions (norms, laws, etc) can evolve and adapt.
“It is logically impossible for the “meme” or specific “culture” to create the genetics of conformity after they must have already existed.” This sounds to me like the old creationist argument about the impossibilily of the evolution of the eye: How could creatures “know” that they were going to need eyes if they didn´t even “know” that light existed? Evolution is precisely about this. Chance and gradual variation can account for the development of an eye, or a beehive, or an instinct (like conformity), or a behaviour, or an emotion (like empathy) or an institution. Genes can reinforce what Culture started, and viceversa.
As I said in my previous comment, I don´t really see the controversy. We believe very much the same, but I think you happen to doubt about the existence of memes and/or the possibility of gene-culture co-evolution. But if that is the case, How can you solve the puzzle of ultrasociality in humans and explain the ultimate causes of human group selection? You seem to believe that everything discussed here can be explained by group selection… except group selection itself, I guess… but then, What was the driver which made human group selection possible in the first place?
For what it’s worth, conformity offers fairly strightforward individual-level benefits to social individuals. Conformity arises largely because other people (mostly) don’t like strangeness. They don’t like strange languages, strange food, strange music or strange customs. They like what they know (partly because they can then use cached responses, partly because weirdness is correlated with pathogens, with developmental problems, with being an outsider and with being a stranger). So: weirdos usually take a direct fitness hit. There’s no “disadvantageous within groups, advantageous between groups” business with conformity – it pays off directly in a simple and straightforwards way.
Juan, if “meme” is simply another word for concept, paradigm, belief, idea, then who could claim there is no such thing as a meme? If we look at gene-culture co-evolution then certainly changes in our behavior, such as cooking our food, which must be recognized as cultural have resulted in genetic changes.
But just as post-modernism has taken some simple ideas and gone way too far with them, so too in my view have the meme extremists and the gene-culture extremists. In current form the meme stuff is to the Blank Slate Standard Social Science Model what Intelligent Design is to Creationism. It predicts nothing, cannot be falsified, and can be used as a pat answer for every single behavior humans have. If we substitute “culture” or “learned pattern” or whatever similar term for “meme” then we are right back at SSSM and Blank Slate; largely or entirely excluding or ignoring the role of biological evolution in human behavior.
To my mind, the important task is to separate the cultural from the biological and then see how they interact with each other. Group level physical conflict shows many universal patterns in its expression and in the seemingly cultural spurs to encourage the men to go to war. Simply describing it as a “parasite meme” is both wrong and an unproductive path if we hope to create a world with less war in an intentional way. Claims like “The reason why most people go to war is because “they have to” ” simply beg the question and ignore our evolutionary past.
War is a pretty important topic, with untold numbers of lives literally on the line. Unlike most who venture an academic opinion on the topic, I have at least seen some of the effects of war firsthand and so I seem to be in a distinct minority of people who actually take war seriously in an academic way. What this has resulted in so far is mostly to be found here – http://theroadtopeace.blogspot.ca/ .
When it comes to war, in most cases culture simply serves as essentially random cover for behaviors that are far more based on evolved predispositions for their ultimate motivations than most care to admit. In most cases we do not go to war over religion, religion is simply a very useful tool which is often readily available and which has many features that fit our needs. Belief in religion X makes one a “good” person, according to the Western Religious models for example, and our readily dualistic brain is then quite happy to conclude that those who reject our religion are therefore “bad” people. Well, if they were good people, they would believe in our religion, belong to our group or political party….Killing bad people is a good thing to do. But absent the cover of religion, politics, all of the “major” causes, the same group conflict pattern still shows up. Instead we kill those with a different color bandana, those who cheer for a different chariot team or soccer team.
Most evolutionary theories predict gradualism. Sufficiently large saltations without any sign of the time or resources needed to execute a generate-and-test procedure would be evidence against evolutionary theories – and in favour of divine intervention. Most evolutionary theories also predict tree-like phylogenies in most cases where complex adaptations are involved. They also predict that adaptations with signs of historical development are likely to be found. It seems deeply unfair to criticise evolutionary theories as being “unfalsifiable”. Darwin had to put up with that sort of thing, but surely we know better these days – having had 150 years to come up with some tests.
Re: “if “meme” is simply another word for concept, paradigm, belief, idea, then who could claim there is no such thing as a meme?”
Nobody, of course. However note that this has never been the definition of a meme. Since 1976, the basic idea of a “meme” has included the idea that is is socially transmitted. Concepts, beliefs and ideas can be socially transmitted *or* the product of individual learning. Memes are a different concept. It’s basically the same thing as what some proponents of what they call “ideational culture” refer to as a “cultural variant”.
Well, I virtually agree on everything you say. It seems very difficult to present opinions on this matter without putting too much weight on one of the plates of the scale. Culture and genes are completely interconnected too such an extent that it is very difficult to state that a behaviour is either culture-induced or genes-induced. But, since meme theory is pretty new, it seems logical that “hard” versions of this theory happen to arise. Science usually follows that pattern; From the dialectic struggle between hard meme supporters and hard meme deniers will eventually arise a soft synthetic suitable theory. Right now Dual Inheritance Theory seems pretty much like it.
Dual Inheritance Theory has very little to do with the blank slate. It has the same power of prediction in humans (and may be certain other species) than biological evolutionary theory in the rest of animals, if not more. In fact it requires less assumptions than regular biological evolutionary ecological behaviourism to explain human affairs.
Indeed War is an extremely important issue and worth of hard work investigation. War instincts are already present in animals, but “real” War self-sacrifice instincts are only present, as far as I know, in social insects… and humans. In both cases allegedly fueled by group level selection. As a matter of fact War is the most direct means by which groups compete with each other. Social insects have the logical of selfish gene theory behind them to explain the self-sacrifice behaviour of the insect soldiers. Humans must have something analogous to the kin affinity. It would be too easy and too quick to state that a kind of “selfish meme theory” is behind the human soldiers self-saccrifice behaviour.
Of course it is more complex than that. But it might be true that culture has started a positive feed back loop of gene-culture co-evolution in the individual level and specially in the group level. Of course the “memes” have had to work on previous old gene-induced instincts that are based in the ethnocentric instincts and the innate capacity to detect in-group from out-group. That is why I say these instincts are parasitized, although it really has been more like a mutualistic interaction in the group level (and probably in the individual level as well). It is mutualistic because this culturally guided instincts have been beneficial to the groups (and to group members) in the past. Therefore, and quoting Boyd and Richerson, we could have two sets of gene instincts towards War: the old primitive animal instincts (which we share with chimps) and a new set of gene-culture co-evolved instincts prone to interact with culture subtleties and “modeled” to maximize group level competition.
But in the present War has reached new levels of complexity, specially when the ones that decide to go to War are not in the front line of combat. At this point humans really resemble social insects societies, with their queens safely kept. At this point soldiers seem truly alienated. At this point you need more than group war instincts (either old and new) and social moralistic punishment. You need some degree of inforcement through actual institutions (not magical influences whatsoever) which deliver real punishment for the ones who refuse to go to war. And these institutions have evolved mainly through a cultural evolution process. Institutions are the cultural organs of the human super-organism. That is why I say that people go to war because “they have to”, a gross simplification.
And speaking of soccer (and other team competition sports) tradition. Couldn´t it be that these traditions are a kind of “escape valves” for both the original gene instincts and the gene-culture co-evolved genes and memes? How many wars have been prevented because we have this way of discharging our group agressivity?
I will visit your blog, by the way.
Re: “From the dialectic struggle between hard meme supporters and hard meme deniers will eventually arise a soft synthetic suitable theory. Right now Dual Inheritance Theory seems pretty much like it.”
“Dual inheritance theory” seems to have a dubious name. DNA and culture aren’t the only ways for information to be inherited in biology. Organisms inherit their position, their homes and their ecosystems. Calling all this “culture” would be special pleading. Dismissing it as irrelevant would be awful.
Memetics doesn’t have this problem. Overall, I think that few meme enthusiasts are going to ditch “meme” and “memetics” in favor of “cultural variant” and “dual inheritance theory”. That’s long-winded terminology made up by anthropologists to make culture seem less biological – in the hope of making their theories more digestible to their colleagues. By contrast, the “meme” and “memetics” terminology delight in cultural evolution’s links to biology. Plus the terminology is short and snappy. Most meme opponents don’t understand the idea. At the end of the day, ignorance isn’t much of an argument.
Tim, I am with you regarding Universal Darwinism and Memetics. It´s just that not everything that is inherited is automatically subject to darwinian natural selection. Dawkins coined the term “meme” in his book “The selfish gene” precisely to point out that the existance of other selfish replicators other than the genes is possible.
The controversy about the term “meme” is pointless. Memes are cultural selfish replicators that undergo the same selfish dynamics than genes. Or that ought to be the very definition. Not all cultural traits that are indeed inherited are capable of undergoing such dynamics since it requires fidelity, variation and selection. I don´t see how positions and homes can fit this model.
The term “Cultural variant” is broader than the term “meme” because there are values, attitudes, preferences and beliefs that are not complex enough to suffer mutations and further competition among kindred. However, individuals and even groups vehiculating some of these cultural variants, positions, extended phenotypes, etc, can overcome their competitor thanks to them.
Let´s not be extremists. Replicators do exist. Genes and memes do exist. And some other things that are inherited like wealth and social status. Let´s work with that.
I would claim that: everything that is inherited is automatically subject to natural selection. Natural selection applies to everything that comes into existence. If we disagree on this point, it’s probably over our use of the term “natural selection”.
The term “natural selection” has meant different things at different times to different people. I just use it to mean that nature selects (in some way) which things perish, and which things reproduce. These days lots of people think that the term “natural selection” refers to non-random change – to change not explicable to genetic drift. This usage is poor, in my opinion. I explain in more detail in the article “Selection vs drift – the conceptual mess at the heart of evolutionary theory”. A bunch of other people use “natural selection” to refer to processes that involve reproduction. However, things can be selected for death by nature without being copied. I find this usage too narrow.
You don’t see how position can evolve in a Darwinian manner? Perhaps look up my “Positional inheritance” article – it explains this point in some detail.
Animal homes are a bit different – they are not normally copied (except for human homes). What I said was that “Organisms inherit their position, their homes and their ecosystems”. This is true. For example, rabbit warrens are often inherited by one generation of rabbits from their predecessors. Rabbit warrens are also subject to selection – some are flooded, some collapse, some are destroyed by rabbit predators – and so on. However, nobody would claim that rabbit warrens are copied or reproduce. That means that rabbit warrens cannot exhibit cumulative adaptation (separate from cumulative adaptation in rabbit DNA). However, they *can* still exhibit adaptive fit – as a result of selection acting on them.
I don’t think that your argument that the term “cultural variant” is broader than the term “meme” is right. You seem to argue that memes have a higher minimum complexity than is implied by the concept of “cultural variant”. I don’t see it. Do you know where you got that idea from? Are you working from definitions of these concepts?
The concepts are certainly *extremely* similar. According to Runciman (2009, p.53), Boyd and Richerson initially drafted the book “Not by genes alone” using the term “meme” throughout – and then replaced it with their own term (“cultural variant”). However their “meme sweep” did not catch all the occurrences of the term “meme” – and there’s on left on page 244: “Modern societies, by vastly enlarging the scope for nonparental transmission have also increased the chance of choosing maladaptive memes”. This anecdote illustrates how interchangeable the terms are.