Brachiopods: biomechanical interdependences governing their origin and phylogeny.

The adaptive advantage of epibenthic articulate brachiopods over inarticulate forms resulted from a modification of the mechanics of shell opening from an indirect hydraulic system to a direct muscular one. As a consequence, the articulate brachiopods were able to reduce the complex muscular system of the ancestral inarticulates, freeing two-thirds of the space within the shell for enlargement of the feeding apparatus. The original hydraulic mechanism of the inarticulate brachiopods most likely evolved from the hydrostatic skeleton of metameric lower invertebrates, probably polychaete-like annelids, as shown by a biomechanical analysis. The transitional stages between such annelids and inarticulate brachiopods are presented and explained as adaptive improvements in body construction.