VI.—Features of Placoderm Diversification and the Evolution of the Arthrodire Feeding Mechanism.

Synopsis The initial adaptive radiation of the Placodermi took place rapidly following the development of the basic placoderm adaptive complex after the ancestral scale covering of the trunk fused into a rigid shield, and not long before the group appears in the fossil record in the Lower Devonian. The radiation was mainly concerned with different ways of living in the benthos of a variety of marine and fresh-water environments; a few nektonic species appear late in the history of the Arthrodira. The fossil record shows the evolution of the orders in their adaptive zones. The zones become increasingly distinct as the orders evolve and become more specific in their adaptations, and the arthrodire, antiarch and rhenanid zones segregate into successively occupied sub-zones. The evolution of the Placodermi has been previously described in terms of improvements in the locomotor mechanism by an analysis of changes in the trunk-armour and pectoral fins. A more detailed description can be given by considering the feeding mechanism as well; this is particularly true of the largest order, the Arthrodira. Study of the feeding mechanism involves the cervical joints as well as the jaws and gnathals. The cervical joints had the same functions in feeding as the anterior part of the vertebral column (“the neck”) in many higher fish. In arthrodires jaw action involved vertical movements of the mandibular lever; the upper jaw apparatus is comparable to the rigid palatoquadrate-maxillary complex of primitive bony fish. The mandible was transformed into a bent lever in Brachyosteus by the development of a small “coronoid” process, but the arthrodire jaw apparatus remained undeveloped in comparison with Actinopterygii and Elasmobranchii. Arthrodire jaw suspension was autostylic. Evidence from the Rhenanida and Ptyctodontida has been interpreted to suggest that this condition was secondary, and that primitive placoderms had an elasmobranch or holocephalan-like palatoquadrate with hyostylic suspension. This view is not entirely supported by the state of the palatoquadrate in primitive arthrodires, but there is no good evidence that placoderms had a complete, open spiracular gill-slit (the aphethoyoid condition). Arthrodire phylogeny cannot yet be described in vertical lines, but four successive levels of organization of increasing efficiency can be recognized; the actinolepid, phlyctaenaspid, coccosteomorph and pachyosteomorph levels. These levels can be defined by simple characters relating to broad adaptations in the locomotor and feeding mechanisms. Evolutionary trends in the Arthrodira include the enlargement of the scapulocoracoid and base of the pectoral fin and the reduction of the spinal plate and flank armour, as the fish gain better control in the water and more myomeres become available for use in swimming; and the enlargement of the nuchal gap and development of the cranio-thoracic joint as powerful muscles are developed to raise the head to give a wide gape, accompanied by the specialization of the gnathals for different modes of feeding. Some of these trends are reversed in compressed, nektonic species. The description of arthrodire phylogeny in terms of changes that can be understood from a functional point of view reveals interesting examples of mosaic and parallel evolution. Parabelosteus n.gen. is erected.