The pattern of variation in centipede segment number as an example of developmental constraint in evolution

Abstract

The range of animal morphologies observed in nature is partly determined by natural selection. However, there is no agreement yet regarding whether it is also partly determined by developmental constraint. Testing for the effects of constraint has been difficult due to the lack of both an appropriate null model and a sufficiently simple system capable of yielding unambiguous results regarding the model's plausibility. Here we examine the case of variation in segment number in geophilomorph centipedes. Curiously, while this ranges between 29 and 191, there are no species in which an even number of segments is observed, in contrast to about 1000 species with odd numbers of segments. It seems unlikely that this distribution of character values is determined by selection alone. Using an approach based on Bayesian inference, we attempt to quantify the probability of obtaining the observed distribution of values given a null model in which developmental constraint is absent. Since this probability is in the region of 10-20, we conclude that constraint must be involved. We discuss various implications of this conclusion, and comment on the unexpected absence of neoteny and progenesis in centipede evolution.