Classification and Phylogenetic Significance of Molluscan Shell Microstructure

Abstract

Like most classifications of molluscan shell microstructure published during the past 25 years (e.g., MacClintock, 1967; Kobayashi, 1964, 1971; Taylor, Kennedy and Hall, 1969, 1973; Grégoire, 1972a), the present one is based largely on Bøggild's (1930) monographic work, redefined from a modern perspective of combined light and scanning electron microscopy. However, this is the first attempt to integrate shell microstructure terminology for mollusks with that employed by students of bryozoan and brachiopod shell microstructure (e.g., Williams, 1968a,b, 1970, 1973; Williams and Wright, 1970; Armstrong 1968, 1969; Sandberg, 1971, 1977; Brunton, 1972; MacKinnon, 1974, 1977; MacKinnon and Williams, 1974; Iwata, 1981, 1982). An integration of nomenclatorial schemes is desirable for purposes of interphylum comparison, and is presently needed because there is considerable overlap and inconsistency in the application of microstructural terminology even within single molluscan classes. The present synthesis of shell microstructure nomenclature is possible primarily because of the extensive data base of invertebrate shell mineralogy, microstructure and especially ultrastructure published in more than 300 references in the past 15 years. To these data, the authors have contributed original information of shell mineralogy and microstructure for scores of Recent and fossil mollusks, brachiopods and bryozoans, with a clear emphasis on bivalved mollusks. Many inadequately described microstructure terms have been reanalyzed during the course of this study, either by examining species cited in the literature, or by using closely related species. Perhaps because they are better studied, but probably for other reasons as well, the diversity of molluscan shell microstructures is considerably greater than that of brachiopods and bryozoans combined (Carter, 1979). Consequently, most of the present nomenclature is based on mollusks, and only three of the major microstructural arrangements described in this guide (crossed bladed, semi-nacreous and semi-foliated) were known first in brachiopods or bryozoans and later recognized in molluscs.