viscous populations and the selection for altruistic behaviors

part of william hamilton‘s theory of inclusive fitness/kin selection, which explains how altruism ever could’ve arisen at all (altruism here having a very specific definition), is that it should be possible for genes for altruism to be selected for if close kin interact regularly. kin don’t need to recognize one another for altruism to be selected for. as long as closely related individuals don’t move far from one another — that is, if a population is viscous — selection for altruism might happen.

i can’t see why this couldn’t also apply to lesser forms of altruism, not just the kind where you sacrifice your life for two brothers or eight cousins. you know what i mean. like: reciprocal altruism or nepotistic altruism. or just pro-social behaviors. whatever you want to call them. seems to me that nepotistic behaviors ought to be selected for more easily in viscous populations (if they increase fitness, of course).

and some populations are more viscous than others:

1) inbreeding populations where close relatives marry frequently over the long-term. mating with relatives must be highly viscous [insert sweaty/sticky incest joke here]. not only do the individual members of the population likely interact fairly regularly (can depend on your mating pattern), they pass many of the genes they share in common on to the next generations — who then also interact and mate. that’s what i call viscous! and, as you all know by now, some human populations inbreed more than others, and some have been doing so for longer than others. and vice versa. (see: entire blog.)

2) populations where extended families are the norm. societies where two or three generations of families all stay together, work together, play together. viscous. plenty of opportunity for nepotistic behaviors to be selected for. on the other hand, societies of nuclear families where more distant relatives are seen only once a year on thanksgiving, and then only to argue, and where your your heir is your pet cat…not very viscous. (see: family types and the selection for nepotistic altruism.)

3) socio-economic systems which push for close relatives to remain together rather than dispersing. if that sounds vague, that’s ’cause it is. sorry. i haven’t thought through it all yet. i do have an example of the opposite for you — a socio-economic system which pushed for close relatives to disperse — and that is the post-manorialism one of northwest europe. already by the 1500s, it was typical for individuals in northwest europe to leave home at a young age (as teenagers) and live and work elsewhere — often quite long distances away (several towns over) — before marrying. then it was not unusual for them to marry someone from their new locale. not viscous. conversely, many societies outside of the hajnal line (northwest europe) have had systems which encouraged the opposite.

food for thought.

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family types and the selection for nepotistic altruism

it finally clicked in my head while thinking about polygamy what the importance of family types — nuclear vs. extended, etc. — might be in the selection for altruistic behavioral traits, especially nepotistic altruism or clannishness. i should’ve thought through polygamy sooner instead of putting it off, but hey — procrastination is heritable, too, so in the words of h. solo, it’s not my fault! (~_^)

the logic of the mating patterns/inbreeding-outbreeding theory goes that, given the right set of circumstances (i.e. certain sorts of social environments), selection for nepotistic altruism/clannishness ought to go quicker or be amplified by inbreeding (close cousin marriage or uncle-niece marriage) simply because there will be more copies of any nepotistic altruism genes (alleles) that happen to arise floating around in kin groups. in other words, inbreeding should facilitate the selection for clannishness…if clannish behaviors are being selected for in a population.

the thing is, though: the individuals carrying certain versions (alleles) of nepotistic altruism genes need to direct their nepotistic behaviors towards other individuals carrying those same alleles, otherwise their actions will be for naught. (yeah. kin selection.) if they direct their nepotistic actions towards people who don’t share the same alleles, then the actions will be “wasted” and the behavioral traits won’t be selected for — or at least not very strongly — and they might fizzle out altogether.

let’s take an imaginary society as an example: say everyone in our pretend population always marries their first cousins. their father’s brother’s daughters (fbd) even, so that we get a lot of double-first cousin marriage. h*ck! let’s throw in some uncle-niece marriages on top of it all. the inbreeding coefficients in such a society would be very high, and if clannishness was being selected for in our highly inbred population, the selection ought to move pretty quickly.

but suppose we separated all the kids at birth from their biological families and set them out for adoption by unrelated individuals — people with whom they likely did not share the same nepotistic altruism alleles. think: the janissary system, only on a population-wide scale. if we did that, there should be virtually no selection for clannishness despite all the inbreeding since pretty much no one’s nepotistic behaviors would be directed towards other individuals with the same nepotistic altruism genes. in this case, kin selection would just not be happening.

such a society does not exist, and i don’t think ever has. but there are societies out there with certain family types — namely nuclear families (or even post-nuclear family societies!) — which ought to have a similar dampening effect on any selection for clannishness.

northwestern “core” europe has had very low cousin marriage rates since around the 800s-1000s, but it has also, thanks to manorialism, had nuclear families of one form or another (absolute or stem) since the early medieval period — nuclear families are recorded in some of the earliest manor property records in the first part of the ninth century from northeastern france [see mitterauer, pg. 59]. on the other hand, eastern europeans, like the russians and greeks, while they also seem to have avoided very close cousin marriage for several hundreds of years (which is not as long as northwestern europeans, but is quite a while), have tended to live in extended family groupings. you would think that nepotistic altruism could be selected for, or maintained more readily, in populations where extended family members lived together and interacted with one another on a more regular basis than in societies of nuclear family members where individuals interact more with non-kin. societies comprised of nuclear families are more like my hypothetical janissary society above where the altruism genes that might’ve been selected for via kin selection instead fade away in the wash.

we have to be careful, though, in identifying nuclear family societies. the irish of today, for instance, are typically said to be a nuclear family society, but the extended family does still interact A LOT (i can tell you that from first-hand experience). same holds true for the greeks and, i suspect, the southern italians. i would say that these populations have residential nuclear families, but not fully atomized nuclear families which have infrequent contact with extended family (think: the english). the early anglo-saxons in england were also characterized by residential nuclear families — the extended family (the kindred) was still very important in that society. the individuals in a residential nuclear family society probably do interact with non-family more than individuals in a society structured around extended families or clans, but less so than a true nuclear family society.

the thought for the day then?: family types can also affect the selection for clannishness/nepotistic altruism.

that is all! (^_^)

previously: polygamy, family types, and the selection for clannishness and “l’explication de l’idéologie”

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human biodiversity and culture and history

william hamilton, considered to be one of the — if not the — greatest evolutionary theorists since darwin, had this to say:

“The incursions of barbaric pastoralists seem to do civilizations less harm in the long run than one might expect. Indeed, two dark ages and renaissances in Europe suggest a recurring pattern in which a renaissance follows an incursion by about 800 years. It may even be suggested that certain genes or traditions of pastoralists revitalize the conquered people with an ingredient of progress which tends to die out in a large panmictic population for the reasons already discussed. I have in mind altruism itself, or the part of the altruism which is perhaps better described as self-sacrificial daring. By the time of the renaissance it may be that the mixing of genes and cultures (or of cultures alone if these are the only vehicles, which I doubt) has continued long enough to bring the old mercantile thoughtfulness and the infused daring into conjunction in a few individuals who then find courage for all kinds of inventive innovation against the resistance of established thought and practice. Often, however, the cost in fitness of such altruism and sublimated pugnacity to the individuals concerned is by no means metaphorical, and the benefits to fitness, such as they are, go to a mass of individuals whose genetic correlation with the innovator must be slight indeed. Thus civilization probably slowly reduces its altruism of all kinds, including the kinds needed for cultural creativity (see also Eshel 1972).”

so hamilton clearly thought that biology and human biodiversity strongly influence culture and history, including the broad movements of history like renaissances or maybe even reformations or enlightenments, etc. and he thought that outbreeding, specifically too much outbreeding (i.e. panmictic populations), and presumably inbreeding too, relate to the selection for altruistic behaviors…and, therefore, certain aspects of cultures and history, etc. (remember that there’s more to hbd than just iq. (~_^) ) i dunno, maybe i and other hbd-ers are crazy (if so, we’re in GOOD company!), but this just makes intuitive sense to me. as john derbyshire said [15:00]:

“…if dimensions of the individual human personality are heritable, then society is just a vector sum of a lot of individual personalities.”

i like the big, probably impossible to answer fully questions: where does culture come from? where do institutions come from? where do renaissances come from? i don’t have the answers to those questions. nor am i under any illusions that i’ll ever be able to answer them. but am i very certain that they cannot be answered without taking into consideration human biology and biodiversity along with more conventional explanations drawn from history, economics, etc., and so i like to periodically bring them up.

so, if you happen to be new here, if you don’t like questioning — on every level — or biological explanations applied to The Big Questions, i’m afraid you’ve come to the wrong place. sorry. for my fellow hbd-ers — see you back here later in the week! (^_^)

previously: renaissances
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p.s. – btw, my vacation has been extended by a week (long story), so i’ll be back properly next week. sorry for the delay! (*^_^*)

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inclusive fitness

there’s some amount of confusion out there in the hbd-o-sphere (and beyond!) about inclusive fitness, which is understandable since the concept is not that straightforward — especially for those of us who are not scientists. i thought it’d try to dispel some of that misunderstanding by sharing my layman’s understanding of the concept — i think i grok the idea pretty well now (in a basic sorta way) — hope i don’t add to the confusion!

to start with, inclusive fitness is NOT some sort of biological law that organisms (including humans) will automatically be altruistic towards other individuals with whom they share a lot of genes (or vice versa if vice versa). if you hold that idea — and i get the impression that a lot of people do — get it out of your mind right now! you’ll feel better for it, trust me.

inclusive fitness is simply a concept or model which explains HOW certain social behaviors — especially altruism — might’ve evolved at all. period. full stop.

to understand inclusive fitness, we need to back up a sec first and think about fitness and what that is. very (very!) simply, fitness refers to an organism’s ability to survive and reproduce in a particular environment. traits — including behaviors — that enable an organism to survive and successfully reproduce will be selected for simply because that organism *is* able to survive and reproduce in its environment. this is natural selection. pretty simple, really, darwin’s dangerous idea.

when it comes to certain social behaviors in humans, it’s readily understandable why many of them were selected for. for example, mothers who devote a lot of time and energy to care for their infants — who obviously can’t take care of themselves and would die without any care — will be more fit than those mothers who don’t. the genes that predispose for those behaviors get selected for since children get half of their dna from their mothers, and the ones that are cared for are much more likely to survive.

what was — and to some extent still is — a big mystery is why other sorts of altruistic behaviors were ever selected for even though they hurt an organism’s fitness. how would self-sacrificing altruistic behaviors directed towards non-descendants ever be selected for? for instance, why on earth would somebody feel compelled to run into a burning building to save a neighbor (who wasn’t their child) at great risk to their survival and, therefore, to their fitness? we can see how “genes for altruistic behaviors towards offspring” could be passed down from mother (or parents) to kids, but how were genes for more general altruistic behaviors selected for?

here is where william hamilton‘s absolutely genius idea — inclusive fitness — comes in: perhaps certain social behaviors, which on the surface appear to reduce an organism’s fitness, and so shouldn’t get selected, might’ve been selected for if those behaviors were directed toward other close kin with whom individuals also share much dna in common.

everybody gets half of their dna from each of their two (for now, anyway) parents. but we also share dna with siblings and (blood-related) aunts and uncles and (wait for it…) cousins. given this inheritance pattern, probability says, for instance, that, in a randomly mating population, an individual should share 12.5% of their dna with a first cousin. so, if an individual with certain “genes for altruism” behaves altruistically toward their first cousins, odds are not bad that those first cousins might also have those same “genes for altruism.” here, then, we have a mechanism for how apparently self-sacrificing social behaviors can be selected for: since the altruistic individual 1) aids close kin with whom he shares much of his dna AND 2) probably in many instances shares the same “genes for altruism,” his being altruistic toward those kin 1) does not reduce his fitness AND 2) the “genes for altruism” get selected for, too. mystery solved. (see also: kin selection.)

one way i like to think of inclusive fitness — which, perhaps, isn’t entirely the right way to look at it, but i feel it helps my understanding — is that if you wanted to calculate an individual’s total fitness by adding up how many actual copies of his genes he passed on, you need to add together those found in his offspring and those in his close relatives’ offspring. in other words, you need to add together his own direct fitness plus his close relatives’ fitness to get his inclusive fitness (or his total fitness).

it seems likely that many of the altruistic (or spiteful, etc.) behaviors we’re talking about are pretty general in nature, i.e. not that specific behaviors like “be altruistic to your close kin” were selected for, but rather more like “be altruistic to the people around you, because they’re probably your close kin” were. it remains to be seen how much kin recognition plays a role in altruism in humans, but that’s a topic for another post anyway. for right now, i just wanted to make clear what inclusive fitness is — and isn’t. again, inclusive fitness is a concept which explains HOW altruistic behaviors MIGHT be selected for. it does NOT predict that individuals will DEFINITELY be more altruistic toward those with whom they share much dna.

the whole topic of inclusive fitness is, of course, much more complicated than all that, but i think this is a good basic intro to the concept. hope so, anyway! (^_^)

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reverse renaissance?

trigger warning: the following post contains much that is speculative. in fact, the entire post is one long speculation. if the thought of speculating when it comes to human biodiversity/sociobiology makes you queasy or fills you with existential angst, this might not be the blogpost for you. no, really. you might want to pass the time in some other way.
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i wrote about this once before, and since i’m extremely lazy, i’m just going to cut-and-paste from the previous post:

“in Innate Social Aptitudes of Man: An Approach from Evolutionary Genetics [pdf], william hamilton suggested that, perhaps, one gets a renaissance by (re-)introducing barbarian altruism genes into a too outbred population, letting the mixture ferment for ca. 800 years or so, and then enjoying the fruits of everyone’s labors. he’s talking here, of course, about the european renaissance of the fourteenth to seventeenth centuries…and classical greece/athens after the dorian invasion of ca. 800 years earlier? i *think*. if it happened at all (link inserted by me):

“‘The incursions of barbaric pastoralists seem to do civilizations less harm in the long run than one might expect. Indeed, two dark ages and renaissances in Europe suggest a recurring pattern in which a renaissance follows an incursion by about 800 years. It may even be suggested that certain genes or traditions of pastoralists revitalize the conquered people with an ingredient of progress which tends to die out in a large panmictic population for the reasons already discussed. I have in mind altruism itself, or the part of the altruism which is perhaps better described as self-sacrificial daring. By the time of the renaissance it may be that the mixing of genes and cultures (or of cultures alone if these are the only vehicles, which I doubt) has continued long enough to bring the old mercantile thoughtfulness and the infused daring into conjunction in a few individuals who then find courage for all kinds of inventive innovation against the resistance of established thought and practice. Often, however, the cost in fitness of such altruism and sublimated pugnacity to the individuals concerned is by no means metaphorical, and the benefits to fitness, such as they are, go to a mass of individuals whose genetic correlation with the innovator must be slight indeed. Thus civilization probably slowly reduces its altruism of all kinds, including the kinds needed for cultural creativity (see also Eshel 1972).'”

william hamilton — probably the greatest evolutionary theorist since darwin and an evil, evil speculator! not to mention crimethinker.

anyway…my own speculation re. the biological substrate of renaissances is that it’s not populations which experience an injection of barbarian altruism genes that wind up having a renaissance, but rather that populations which outbreed (i.e. quit marrying close relatives) for ca. 400 to 800 years (egs. medieval/renaissance europe and archaic/classical greece?) undergo a sort-of wikification of their society which drives intellectual openness and curiosity and sharing — the kinds of behavioral derring-do that you need in order to have a renaissance at all. see the previous post for more on all of those speculations.

today’s speculation is that perhaps the arabized world underwent a reverse renaissance process thanks to the introduction by the arabs of the most inbred form of cousin marriage — father’s brother’s daughter (fbd) marriage — to the populations of the middle east/maghreb (not to mention the introduction of arabs, themselves, who had probably been inbreeding closely for up to nine hundred years before their expansion).

the islamic golden age lasted for a good six hundred years or so, but instead of the scope of islamic philosophy and science and law widening over the time period — instead of a wikification process — the tendency was for thinking in the arabized world to narrow. ijtihad (“independent thinking”) was gradually replaced by taqlid (“imitation”). this narrowing of thought was already widespread in the muslim world by the twelfth century — just about 400-450 years after the arab conquests. (braudel puts the beginning and end dates of the islamic golden age as 813 and 1198, the beginning of al-ma’mun’s caliphate and the death averroes respectively. – pg. 202.)

irfan habib points out that the islamic golden age in science was very much founded on long-established traditions of free inquiry in the near east, from greece to persia [pg. 69 — link added by me]:

“[T]his particular phase in Islamic history was marked predominantly by the Mu’tazilite school of philosophy, which was based on freethinking and rationalism. It was an ecumenical setting for science, where savants of nearly all creeds and origins worked towards a common purpose. And this was not something new, it was a long established pre-Islamic tradition in the Near East, where translation of scientific and philosophical texts from Greek to Syriac took place….“

i wonder if what happened was that, with the establishment of the caliphate and all the civilized elements that went with it — good communications over long distances, (relative) peace within the realm, an excess of wealth — a “renaissance” was quickly established. however, that golden age — which happened in the early part of the era of the caliphs — was really a late flowering of whatever had been going on the region previous to the arabs (especially in persia). this renaissance was then reversed — stunted, really — as a result of the centuries of close inbreeding of the populations in the middle east and maghreb thanks to the introduction of fbd marriage by the arabs.

like i said — pure, unadultered speculation! (~_^)

previously: renaissances

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“genes for altruism”

in Genes underlying altruism published in october of last year, three biologists/researchers think through what “genes for altruism” ought to look like and how we will recognize them (“we” meaning teh scientists!):

(i) Genes underlying altruism should satisfy Hamilton’s rule of rb > c, where r is relatedness of actor to recipient, b is benefits to the recipient and c is costs to the actor. Altruism exists, and to the extent that this type of behaviour has evolved, we expect genetic variation to underlie it. In this sense, there must be genes ‘for’ altruism (genes showing allelic variation that is statistically associated with variation in altruistic behaviour) that are potentially detectable….

(ii) Genes underlying altruism should be environmentally sensitive. If genes for altruism are to evolve, then at least some carriers must reproduce. This inference implies that genes underlying altruism should be conditionally expressed as a function of their social environment….

(iii) Genes underlying altruism should increase in number and complexity with social-behavioural sophistication….

(iv) Genes underlying altruism should coevolve with, or depend on, the previous evolution of genes for kin recognition….

(v) Genes for altruism may reside in regions of low recombination, exhibit co-expression and show modular genetic architecture….

(vi) Genes underlying altruism should be at least partially additive. The evolution of altruism requires heritable variation, and we therefore expect genes for this and other evolved social traits to have significant additive effects that are responsive to kin-mediated selection….

(vii) Genes underlying altruism should exhibit strong pleiotropy. Pleiotropy (multiple phenotypic effects of alleles) should be fundamental to the evolution of altruism, given that it involves combinations of costs and benefits that may be simultaneously physiological, morphological, reproductive and behavioural….

the authors offer some candidate “genes for altruism” (this is a truncated version of their table. i’ve only included the ones for humans here — they also suggested some for eusocial insects):

i think teh scientists should also look for some genes related to violence — particularly tempermental, hotheaded sorts of violence — the kind that raises the testosterone levels of scots-irish, but not yankee, folks when they’re insulted. those types of fly off the handle behaviors, i think, are often altruistic in nature, since the hotheaded individual can be more willing to sacrifice himself in a fight or in battle for his kin.

also, i guess that “genes for altruism” ought to be found in differing frequencies — even variations — in different human populations, especially long-term inbred versus long-term outbred ones.

(^_^)

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inbreeding, digit ratio, and altruism

in his post A pathway to pro-social behavior, peter frost references this article Second-to-Fourth Digit Ratio Has a Non-Monotonic Impact on Altruism in which the authors say:

“We find an inverted U-shaped relation for left and right hands, which is very consistent for men and less systematic for women. Subjects with both high and low digit ratios give less than individuals with intermediate digit ratios.“

the subjects were w.e.i.r.d.-ish, btw — university students from granada, spain.

peter comments:

“From one population to the next, digit ratios tend to cluster around different means, perhaps because altruism has been favored or disfavored to different degrees.”

i’m not familiar with those differences (will have to investigate), but i did post this from turkey in a recent linkfest:

“Inbreeding is associated with lower 2D:4D digit ratio”

“We compared the 2D:4D ratios of 122 male and 108 female consanguineous (children of first cousin marriages) high school and university students to those of 142 male and 122 females controls. Across hands and sex, consanguineous parentage was consistently associated with lower, more masculine-typical, digit ratios. Digit ratios were 1.3–1.9 times more variable among the consanguineous group than the control group. While socio-economic status cannot explain the effects seen in our data, we found that lower, more masculinized, digit ratios were associated with lower family income.”

inbreeding leads to…whatever genotypic/phenotypic package that results in lower 2d:4d digit ratios *and* lower rates of altruistic behaviors? dunno. Further Research is Required^TM.

ftr — i come from a population that has been inbreeding up until fairly recently, and i’ve got a low 2d:4d digit ratio. uh oh.

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renaissances

in Innate Social Aptitudes of Man: An Approach from Evolutionary Genetics [pdf], william hamilton suggested that, perhaps, one gets a renaissance by (re-)introducing barbarian altruism genes into a too outbred population, letting the mixture ferment for ca. 800 years or so, and then enjoying the fruits of everyone’s labors. he’s talking here, of course, about the european renaissance of the fourteenth to seventeenth centuries … and classical greece/athens after the dorian invasion of ca. 800 years earlier? i think. if it happened at all (link inserted by me):

“The incursions of barbaric pastoralists seem to do civilizations less harm in the long run than one might expect. Indeed, two dark ages and renaissances in Europe suggest a recurring pattern in which a renaissance follows an incursion by about 800 years. It may even be suggested that certain genes or traditions of pastoralists revitalize the conquered people with an ingredient of progress which tends to die out in a large panmictic population for the reasons already discussed. I have in mind altruism itself, or the part of the altruism which is perhaps better described as self-sacrificial daring. By the time of the renaissance it may be that the mixing of genes and cultures (or of cultures alone if these are the only vehicles, which I doubt) has continued long enough to bring the old mercantile thoughtfulness and the infused daring into conjunction in a few individuals who then find courage for all kinds of inventive innovation against the resistance of established thought and practice. Often, however, the cost in fitness of such altruism and sublimated pugnacity to the individuals concerned is by no means metaphorical, and the benefits to fitness, such as they are, go to a mass of individuals whose genetic correlation with the innovator must be slight indeed. Thus civilization probably slowly reduces its altruism of all kinds, including the kinds needed for cultural creativity (see also Eshel 1972).”

“self-sacrificial daring” is probably the equivalent of greying wanderer’s “aggression”, chris’ “drive”, staffan’s “persistence under negative reinforcement”, and/or my “contrarianism” or independent-mindedness.

the connection between these two renaissances might, indeed, be the reintroduction of some good altruism genes, but i think that maybe what these two “rebirths” have in common — what led to them occur at all — are the ca. 400-800 years of outbreeding which happened right before they began. in medieval europe we have the catholic church banning close cousin marriage around the year 500, and many secular authorities banned close cousin marriage at various points after that. and in archaic greece — the period just before classical greece/athens — we apparently have at least ca. 400 years of outbreeding — amongst the upper-classes most probably — and possibly amongst the lower classes, too (hesiod in his Works and Days recommends that a man — an ordinary man, a farmer — marry a nice girl from the neighborhood — from the kome or village — so, if archaic greeks actually did this, their mating patterns would’ve been quite endogamic, but not necessarily to close cousins — maybe third or fourth cousins or something — see A Companion to Archaic Greece).

i think you need some loosening of the genetic ties in populations — enough to get rid of a lot or most of the “clannishness” — so that you can have a “wikification” of those societies, i.e. societies where individuals are really willing to openly share their ideas with other like-minded people (see, for example, harold’s comment on the scientific revolution in england). but outbreed too much, and you might lose that “self-sacrificial daring” — because as hamilton said:

“…the benefits to fitness, such as they are, go to a mass of individuals whose genetic correlation with the innovator must be slight indeed.”

share your innovative ideas — your scientific inventions — with the entire world, and you might wind up benefitting all of those people more than your own descendents (if you’ve got any).

already at the start of the classical period in greece/athens, the mating patterns began to narrow [pg. 67]…

“[W]ith the emergence of the *polis*, exogamy began to give way in some places to endogamy — to marriage within the community. For the upper classes, this meant marriage within a tight circle of aristocratic families living in the same *polis*.”

…so it’s maybe no surprise that the athenians battled throughout the classical period against various aspects of clannishness (cleisthenes’ reforms are one huge example of this struggle) and that their renaissance didn’t last more than a couple hundred years. europeans, on the other hand — especially northern europeans — have continued to outbreed for something like over ca. 1000-1400 years — which, perhaps, is leading to another sort of problem for society — that it’s simplying fraying away at the seams because the weave is not tight enough.

maybe. dunno. all wild speculation on my part, obviously.

previously: archaic greek mating patterns and kinship terms

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