there were a handful of science news stories out this past week about how a couple of researchers reportedly discovered a case of “group selection” in certain spiders (Anelosimus studiosus or tangle web spiders): for example, see Proving ‘group selection’: Spider colonies need the correct mix of personalities to survive and Elusive Form of Evolution Seen in Spiders. a bunch of people on twitter got all excited about this finding, because they wonder if (some of them i think hope that) group selection might also apply to groups of humans. i agree: that would be very interesting to know one way or the other. so i went and read the original paper — Site-specific group selection drives locally adapted group compositions — to see what these guys had found.

before i offer up my admittedly layman’s thoughts on this paper, let me first say what a really neat piece of research this was! if there existed a nobel prize for geeky dedication and sheer nerdiness, these guys would’ve won it! — and i mean that as a compliment! the researchers, pruitt and goodnight, studied groups of tangle web spiders in the wild, captured some and brought them back to their lab, conducted personality tests on the spiders (yes! there are apparently personality tests for spiders!), painstakingly painted those little dots on the backs of individuals to keep track of them (you know, like how they sometimes do with bees), bred the spiders, released new groups of them back into the wild, and checked up on them one and two generations later to see how they fared. this is some really cool research! nerds ftw! (^_^)

but did they find evidence for group selection?

weeeellll, no, i don’t think so.

to begin with, right at the start of the paper pruitt and goodnight (p&g) define group selection as “selection caused by the differential extinction or proliferation of groups.” eeeehhhh, as far as i understand it, that’s not really the definition of group selection, and even the authors admit that their definition is a “broad” one.

group selection is more accurately defined as when “natural selection [operates] between groups of organisms, rather than between individuals.” in contrast, p&g’s broad definition could theoretically include cases in which natural selection worked between individuals (individual selection) which also just incidentally happened to result in the proliferation of the group to which the lucky selected individuals belonged. an example of this is the selection for lactase persistence in some humans in which those individuals who could drink milk as adults were able to leave behind more descendants than those individuals who could not. while lactase persistence might indeed have benefitted groups of milk-drinking individuals, natural selection did not act on the group, but rather on the individuals in that group. (pretty sure i stole this example from @supermisdreavus, but i can’t find where he said that right now.)

in other words, you always need to work out what the target of selection is: the group or the individuals that make up the group. (really it’s ultimately the genes, but — oh, nevermind.) remember that “‘a fleet herd of deer’ is really just a herd of fleet deer.”

so, really, the discussion could end right here, because i don’t think the authors are talking about group selection proper. but, since i’ve read the whole paper, i’ll carry on. (yes, i’m one of those people who’s never learned to quit while they’re ahead!)

a. studiosus spiders live either as solitary individuals or in groups where they cooperate on tasks like hunting and the raising of young. the individuals that live together in groups are, on average, more closely related to one another than those that live alone [pdf] — they’re generally as related to one another as though they were half-siblings. one reason why they’re probably not more related to one another in these groups — like to the degree that ants or bees in colonies often are — is that the males move between groups. remember that.

the personality types of the individual spiders in a. studiosus groups come in two sorts: docile and aggressive. the docile spiders are typically pretty laid back and aren’t much bothered by the presence of other spiders (even spiders from other species), whereas the aggressive individuals like their space — they’ll chase off other individuals. individuals of both types are found in groups of a. studiosus, but the frequencies vary. from the paper:

“At…high-resource sites, small colonies were dominated by docile females and the frequency of aggressive individuals increased with colony size. By contrast, at low-resources sites, small colonies were dominated by the aggressive phenotype and the frequency of the docile phenotype increased with colony size.”

well, that doesn’t sound too surprising at all. in locales where there is plenty of resources, there are more laid back individuals in the colonies, prolly ’cause being laid back works just fine. in areas where resources are lacking, more aggressive individuals do better. btw, they found that the heritability of these personality types in the spiders is 0.66.

groups that have more docile individuals (i.e. the ones in high-resource areas) are at a greater risk of invasion by other types of spiders which, over the long-term, tends to be a really bad thing for an a. studiosus colony (i.e. it’s usually destroyed). groups that have more aggressive individuals (i.e. the ones in low-resource areas) tend in bad times to experience too much “egg case cannibalism.” needless to say, that’s not a good thing over the long-term either.

what p&g did in their study was to introduce into the wild — into differing environments — groups having varying frequencies of these personality types [source]:

“He [pruitt] took spiders from warrior-heavy colonies and used them to assemble new groups that were heavy on the nannies. He also used spiders from mostly docile colonies to create warrior-laden groups. In addition, he assembled control groups that matched the composition of their original groups.”

what they found was that after three generations:

“60 percent of the colonies were extinct. Control groups that returned to their ancestral homes tended to do well, and those that were transplanted into a new environment generally died. Neither of these outcomes was much of a surprise.

The most interesting results came from colonies made up of spiders that had been forced into a composition different from the one they grew up in — warrior-majority colonies containing spiders from mostly docile groups, for example. The colonies whose composition fit the new environment tended to survive. But over time, surviving colonies reverted to their members’ original group composition. The warrior-majority colonies went back to having more nannies, for example. On the face of it, this is bizarre behavior; if the colonies are well-suited to their environment, why not maintain that ratio? It seems that some innate sense, perhaps encoded in the spiders’ genes, pulled the colony back to its original configuration, even though this change meant the colony would perish.”

well, i dunno. is that really “bizarre behavior?” i mean, if the personality types of a. studiosus are really highly heritable (0.66), is it strange that a population having come from a bunch of docile individuals should regress toward a docile mean? and vice versa? don’t forget, too, that the individuals in these groups are all related to one another as though they were half-siblings, so presumably individuals of either personality type might carry a great many genes of the other type in their genomes. (don’t know about that — i’m just guessing here, tbh.)

what really made me question whether or not this is “bizarre behavior” is the way in which the researchers bred the spiders when they had them in captivity [from the methods section at the end of the paper]:

“Females were mated randomly to a male of like behaviour type from their same source population, but which was collected from a source colony >5m distance.”

hmmmm. i dunno about that. they mated all the females with males of the same personality types, docile or aggressive? i’m guessing that they did this in order to reduce the number of possible confounding factors in the study, but i’m afraid they might’ve added something to the mix here that wouldn’t be found in nature, i.e. a 100% assortative mating rate (for personality type). mightn’t this almost guarantee that individual spider lineages would regress to their original personality-type means? docile females always mated with docile males and aggressive females always mated with aggressive males? that seems unlikely to happen in nature, especially given the fact that the males normally leave their colonies and move to others. (btw, male a. studiosus spiders prefer moving into colonies over mating with lone females. typical males, favoring harems! (~_^) )

p&g offer a number of explanations for how the frequencies of personalities in the groups might change over time:

“How native spiders are actually able to adjust their composition is unknown, but plausible regulatory mechanisms include developmental plasticity in the docile:aggressive phenotypes, policing of group membership, phenotype-biased dispersal, and/or selective cessation of reproduction.”

they reject the first explanation (the plasticity one) on the basis (in part) of the rather high heritability of spider personality types which they found. i’m inclined to agree with them on that.

out of their other reasons, policing of group membership and selective cessation of reproduction are behaviors that can be easily explained by natural selection between individuals, especially in populations that have rather highly related individuals so that levels of altruism are pretty high. the selective cessation of reproduction occurs, for instance, in some ant colonies since, due to the really high degrees of relatedness between individuals, the inclusive fitness payoffs are really large (eg. if you share three-quarters of your dna with your sister’s offspring, there’ll be a greater genetic payoff in helping her to reproduce rather than reproducing yourself, since you’d only share half of your genome with your offspring). that’s individual selection, not group selection. h*ck! both behaviors also occur in meerkat groups, although they, of course, show much less specialization of individuals than ants or bees. the policing of group membership can also be plausibly explained by natural selection between individuals — for example, aggressive individuals keep at bay all sorts individuals because that’s good for aggressive individuals (who are typically found in sparse environments).

so, i’m not at all convinced that pruitt and goodnight have found an example of group selection. i think they’ve found that genetics (as indicated by the heritability of the spiders’ personality traits) and natural selection certainly shape the average characteristics of groups, but it looks to me as though the seemingly “bizarre behaviors” that they found can easily be explained by individual selection. in fact, i’m more than a little concerned that due to the way they bred the spiders, p&g may have affected the outcomes of the reintroduced groups.

see also: The False Allure of Group Selection from steven pinker.

(note: comments do not require an email. a. studiosus group web!)