Growth increments in the shell of the living brachiopod Terebratalia transversa

Scanning electron micrographs revealed growth increments in the primary shell layer of the extant terebratulid brachiopod Terebratalia transversa collected from Anacortes, Washington, USA, during the summers of 1982–1984. The increments extend into the secondary shell layer, but only as poorly-defined continuations. Growth increments narrow and widen cyclically, producing patterns similar to tidally-induced, fortnightly growth patterns in bivalve molluscs. Asymmetric increments or doublets that consist of paired wide and narrow bands present additional evidence that the growth pattern is produced by the interference of tidal (24 h 50 min) and solar (24 h) daily environmental influences on growth, as is the case in bivalves. Growth increments are relatively well-defined in specimens from tidal habitats and are poorly-defined in subtidal specimens. Specimens grown in laboratory aquaria and subjected to simulated tidal emersions and submersions had well-defined increments, whereas those that were continuously submerged in aquaria had poorly-defined increments, paralleling the distinctness in growth increments produced at various depths in the natural habitat. Production of growth increments in T. transversa may be related to valve movement and changes in mantle-cavity pH. When the valves are closed, pH drops and, when they are open, pH rises. Highperformance liquid chromatography (HPLC) of mantle tissue and fluid failed to detect any single organic acid that could have been responsible for the pH changes, confirming that intermediate metabolism in T. transversa is very low compared to that of bivalve molluscs. Calculations indicate that cycles in intermediate metabolism may nevertheless be responsible for cycles in growth-increment production and shell dissolution.Please address all correspondence to Dr. G.D. Rosenberg in Indianapolis