Growth and function of the micro-frills present on the Devonian brachiopod Athyris campomanesi (Verneuil & Archiac)

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Abstract
Minute, lamellose concentric micro-frills are described on the external surfaces of the Devonian brachiopod Athyris campomanesi (Verneuil & Archiac). The micro-frills are very regularly spaced with a separation of approximately 0.5 mm, are inclined anteriorly and antero-laterally, and become more or less recurved peripherally down towards the valve surface. These micro-frills must have been secreted by outward extensions of epithelium, presumably resulting from an increase in the rate of cell proliferation or extension in the marginal zone. Extrapolating from the known growth rates of living brachiopods, it seems probable that the micro-frills formed very rapidly, within a matter of days or hours. In life the micro-frills would have formed minute channels around the peripheries of both valves, similar in form and function to rain gutters. As such they probably functioned as baffle chambers, within which excess paniculate material from the inhalant feeding currents would have accumulated. Such particulate material is likely to have been moved laterally from the median axis of the shell down to the sediment surface under the effects of gravity and in vitro shell movements. Each micro-frill may only have functioned in this fashion for a matter of weeks before being rendered ineffective by the forward growth of the valve margin, at which stage it would have been abandoned and replaced by a subsequently formed micro-frill.