Spikey bivalves: intra-periostracal crystal growth in anomalodesmatans

The external shell surfaces of most anomalodesmatan bivalves are studded with small spikes, particularly at the posterior end. We have studied the morphology, mode of growth, and distribution among taxa of these spikes. In this study we found that spikes vary widely in morphology, from acute spikes to flat plaques. Optical and electron microscopy has revealed that the periostraca of Laternula, Myadora, and Thraciopsis consist of an outer dense layer and an inner translucent layer. The dense layer grows at the expense of the inner layer as it progresses toward the shell edge. The spikes begin to grow in the free periostracum, within the translucent periostracal layer, immediately below the dense layer. With growth, they push the dense periostracal layer upward but without penetrating it. Those parts of the spike in contact with this layer cease to grow, which explains the typical conical shape of spikes. When fully grown, spikes reach the base of the translucent layer, becoming incorporated into the outer shell layer. Scanning electron microscopy and electron backscatter diffraction analysis reveal that the spikes of Lyonsia norwegica and Lyonsiella abyssicola are prisms of aragonite composed of twinned crystals, with the c-axis vertical. A survey of the occurrence of spikes within the anomalodesmatans shows that they are present in all but a few families. Elsewhere within the closely related palaeoheterodonts, intra-periostracal calcification is also known in Neotrigonia and unionids, which indicates that this character may be plesiomorphic for these bivalves. The present data do not support the homology of spikes in other bivalve groups (e.g., veneroids) or in the aplacophorans or polyplacophorans.