19 October 2009

Social termites team up with non-relatives

This post was chosen as an Editor's Selection for ResearchBlogging.orgIn social insects, colonies of hundreds or thousands of workers and soldiers forgo reproduction to support one or a few "reproductives" -- drones and a queen. In most cases, this isn't as selfless as it might seem. Because the workers in a colony are all offspring of the queen, they're really reproducing through her -- because the queen shares genes with the workers, when she reproduces it contributes to their evolutionary fitness.

This is called kin selection, and in many cases it's a good explanation for the way the interests and behavior of individual workers are overridden by the interests of the colony. There are, however, exceptions -- and an open-access paper in the latest issue of PNAS describes what looks like a good case: mergers between unrelated colonies of termites.


Zootermopsis nevadensis, a social insect inclined to negotiated settlements. Photo by BugGuide/ Will Chatfield-Taylor.
The termite Zootermopsis nevadensis lives in small, socially-stratified colonies that tunnel through rotting logs. Each colony has a pair of reproductive individuals, a king and queen, served by sterile workers and soldiers. Multiple unrelated colonies usually nest in a single log, and when they encroach on each other's territory, something interesting happens -- they merge.

In what the authors refer to obliquely as the "interaction" that precedes a merger, the king and queen of one or both colonies may die. Mergers occur in the aftermath, as workers from the two colonies began to work in concert, and one or a few of them become replacement reproductives. This ability of sterile workers to start reproducing in the absence of a king and queen is unique to termites. DNA analysis shows what happened after mergers -- new reproductives could arise come from either or both colonies, and that in some cases they interbred.

It's this possibility to become genetically invested in the newly merged colony, the authors say, that motivates workers from two unrelated colonies to work together. If this is the case, it means that kin selection is not what keeps merged colonies together. Group selection might be a better explanation. Kin selection is often contrasted with group selection, in which unrelated individuals sacrifice their own interests to those of a larger group, so that their colony can better compete against rival colonies. In a classic 1964 Nature paper [$-a], John Maynard Smith discussed the conditions under which kin selection operates well:
By kin selection I mean the evolution of characteristics which favour the survival of close relatives of the affected individual, by processes which do not require any discontinuities in population breeding structure.
And contrasts them to conditions necessary for group selection to work:
[Under group selection] ... If all members of a group acquire some characteristic which, although individually diadvantageous, increases the fitness of the group, then that group is more likely to split into two, and in this way bring about an increase in the proportion of individuals in the whole population with the characteristic in question. The unit on which selection is operating is the group and not the individual.
The ecology of Zootermopsis nevadensis may set the stage for group selection to overpower kin selection. With many small colonies competing for a single rotting log, the benefits of possibly contributing to the reproduction of a larger, more competitive colony make mergers worthwhile. Something similar has been documented in ants, which can form supercolonies of unrelated colonies if there is some external threat (another ant species) to force them to band together -- you can find discussion of a recent paper on this case over at Primate Diaries.

References

SMITH, J. (1964). Group selection and kin selection Nature, 201 (4924), 1145-1147 DOI: 10.1038/2011145a0

Johns, P., Howard, K., Breisch, N., Rivera, A., & Thorne, B. (2009). Nonrelatives inherit colony resources in a primitive termite Proc. Nat. Acad. Sci. USA, 106 (41), 17452-6 DOI: 10.1073/pnas.0907961106

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