Variation and specialisation of the forcipular apparatus of centipedes (Arthropoda: Chilopoda): a comparative morphometric and microscopic investigation of an evolutionary novelty.

The forcipules of centipedes are the only known example in the animal kingdom of an evolutionary transition from walking legs to venom-injecting appendages. They provide a classic case of an evolutionary novelty under most (but not all) definitions of that concept. Although there is a reasonable literature on forcipules, and on the forcipular segment more generally, it is fragmentary and scattered. Also, many previous studies have been based on a single species and hence have no comparative component. Here, we build on this earlier literature by providing detailed qualitative and quantitative information on the forcipular segments of representatives of the five extant orders of centipedes. Our results reveal notable differences between the orders - as well as considerable variation within some of them. The pattern of inter-group differences can be used to infer, albeit cautiously, a major evolutionary trend from a presumed scutigeromorph-like last common ancestor (LCA), in which the forcipules were probably leg-like (as in present-day scutigeromorphs) to a more specialized claw-like structure with movement restricted to the horizontal plane. This morphological trend may reflect an ecological trend from open-habitat ambush predation to leaf-litter and subterranean predatory opportunism.