Occipital spine of Orthacanthus (Xenacanthidae, Elasmobranchii): structure and growth

The morphology of 16 occipital spines of the xenacanthid Orthacanthus from Upper Carboniferous deposits of Robinson (Kansas, USA), Nýran (Czech Republic) and Puertollano (Spain) is described. The nonreplaced spines reveal the growth pattern of the shark. Moreover, the relationship between growth and paleoenvironmental conditions can be used to determine paleoecological conditions. Both external and internal morphology indicate that the spine was superficially inserted in the skin. During growth, the spine moved from a deep position in the dermis, in which trabecular dentine is formed, to a more superficial location in which centrifugally growing lamellar dentine was formed. Centripetally growing lamellar dentine was deposited more slowly than the centrifugally growing dentine; it obliterated the pulp cavity. The denticles are independent dermal elements formed by a dermal papilla and secondarily attached by dentine to the spine proper. The number of denticles per annual cycle and the density of denticulation vary with the growth rate. Moreover, the ratio of length of denticulated region to total length of the spine changes throughout ontogeny. In consequence, those features cannot be used for systematic purposes without a careful analysis of the variability. Centrifugally growing lamellar dentine in spines from Robinson shows a regular alternation of layers, suggesting tidal conditions in the environment in which the sharks lived. Monthly and seasonal cycles also occur. Tidal (lunar) cyclicity is also observed in the denticles: size and distance between denticles increase and decrease gradually, forming waves that are considered seasonal and yearly cycles. The observed regularity could be related to the variation in calcium phosphate deposition following the cyclical changes in water temperature produced in the tidal zone. Monthly and seasonal cycles are the result of the interaction of the solar and tidal (lunar) cycles. The cyclical pattern of growth is used to determine the age and growth rates. Orthacanthus was a fast-growing shark like the Recent sharks Isurus, Mustelus, and Negaprion.