Architecture of the integument in lower teleostomes: functional morphology and evolutionary implications

A bony ganoid squamation is the plesiomorphic type in actinopterygians. During evolution, it was replaced by weak and more flexible elasmoid scales. We provide a comparative description of the integument of "ganoid" fishes and "nonganoid" fishes that considers all dermal components of mechanical significance (stratum compactum, morphology of ganoid scales, and their regional differences) in order to develop a functional understanding of the ganoid integument as a whole. Data were obtained for the extant "ganoid" fishes (Polypteridae and Lepisosteidae) and for closely related "lower" actinopterygians (Acipenser ruthenus, Amia calva) and "lower" sarcopterygians (Latimeria chalumnae, Neoceratodus forsteri). Body curvatures during steady undulatory locomotion, sharp turns, prey-strikes, and fast starts in "ganoid" fishes were measured from videotapes. Extreme body curvatures as measured in anesthetized specimens are never reached during steady swimming, but are sometimes closely approached in certain situations (sharp turns, prey-strike). During extreme body curvatures we measured high values of lateral strain on the convex and on the concave side of the body. Scale overlap changes considerably (66-127% in Lepisosteus, 42-140% in Polypterus). The ganoid squamation forms a protective coat, but at the same time it permits extreme body curvatures. This is reflected in characteristic morphological features of the ganoid scales, such as an anterior process, concave anterior margin, and peg-and-socket articulation. These characters are most pronounced in the anterior body region, where maximum changes in scale overlap are required. The anterior processes and anterior concave margin, together with the attached stratum compactum, guide movements in a horizontal plane during bending. Displacements of scales relative to each other are possible for scales of different scale rows, but are impeded in scales of the same scale row due to the peg-and-socket articulation. Furthermore, ganoid scale rows, fibers of collagen layers of the stratum compactum, and the lateral myoseptal structures follow the same oblique orientation, which is needed to achieve extreme body curvatures. There is no evidence that body curvatures are limited by the ganoid squamation in Polypterus or Lepisosteus to any larger extent than by a type of integument devoid of ganoid scales in teleostomes of similar body shape. Our results essentially contradict former functional interpretations: 1) Ganoid scales do not especially limit body curvature during steady undulatory locomotion; 2) They do not act as torsion-resisting devices, but may be able to damp torsion together with the stratum compactum and internal body pressure.