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The Shell Structure of the Recent Patellogastropoda (Mollusca: Gastropoda)

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The Shell Structure of the Recent Patellogastropoda (Mollusca: Gastropoda) Paleontological Research, vol. 9, no. 2, pp. 143–168, June 30, 2005 6 by the Palaeontological Society of Japan The shell structure of the Recent Patellogastropoda (Mollusca: Gastropoda) TAKESHI FUCHIGAMI1 AND TAKENORI SASAKI2 1Department of Earth and Planetary Sciences, Faculty of Science, University of Tokyo, 7-3-1 Hongo, Bunkyo-ku, Tokyo 113-0033 2The University Museum, The University of Tokyo, 7-3-1 Hongo, Bunkyo-ku, Tokyo 113-0033 ([email protected]) Received March 25, 2005; Revised manuscript accepted March 28, 2005 Abstract. The shell microstructure of 44 species belonging to 19 genera and 5 families of Patellogas- tropoda was observed by scanning electron microscopy on the basis of material mainly from the Northwest Pacific. As a result, 17 microstructures of prismatic, crossed, and lamellar structures were recognized. The comparison among species revealed 20 shell structure groups which are defined by microstructures and shell layer arrangement. The relations between taxa and shell structural composition indicate that the Recent patellogastropods generally have distinctive and stable shell structures at the genus level. This high level of consistency provides a firm basis for the application of shell structural characters to identify fossil pa- tellogastropods. However, the evolutionary process of microstructures and homology across different shell layers are mostly ambiguous in the absence of robust phylogeny and undoubted positional criteria for comparison. More studies from phylogenetic, ontogenetic and mineralogical viewpoints should be under- taken to discuss the process of shell structure diversification in patellogastropods. Key words: Shell microstructure, new definition, taxonomic occurrence, Patellogastropoda, Recent Introduction tropods using soft-part characters have repeatedly corroborated many cases in which macroscopic tele- Studies of shell microstructure are particularly im- oconch morphology is useless in determining higher portant because they provide information for the tax- systematic position (Haszprunar, 1988; Ponder and onomic and phylogenetic analysis of molluscs includ- Lindberg, 1997). Limpets, including patelliform gas- ing fossil taxa. Detailed comprehensive works have tropods and also monoplacophorans, are particularly been published for various shell-bearing molluscs, es- notorious as a case of multiple convergence. Several pecially bivalves (e.g., Taylor et al., 1969, 1973; Taylor, different major taxa are inferred to have evolved 1973; Carter, 1990a, b), gastropods (Carter and Hall, similar-looking limpet-form shells. Among them pa- 1990; Bandel, 1990a), and cephalopods (e.g., Bandel, tellogastropods are one of the outstanding groups 1990b; Kulicki, 1996). However, studies of shell struc- comprising limpet-shaped species exclusively. ture in gastropods are limited to a relatively small Patellogastropods share many apomorphic charac- number of specific taxonomic groups (e.g., Taylor and ters anatomically, and their monophyly is strongly Reid, 1990 for Littorinidae; Bandel and Geldmacher, supported (Ponder and Lindberg, 1997; Sasaki, 1998). 1996 for Patella; Sasaki, 2001 for Neritoidea; Kiel, Recent phylogenetic studies revealed that the patello- 2004 for gastropods from vents/seeps) and their taxo- gastropods constitute a well-defined independent clade nomic coverage is insufficient compared to other ma- (Eogastropoda), being clearly separated from the jor conchiferan molluscs. Therefore, the gastropod rest of the gastropods (Orthogastropoda) (Ponder shell structure is not well understood in general over- and Lindberg, 1997; Sasaki, 1998). When the shell view even at present. is perfectly intact, the patellogastropods can be dis- One of the difficulties in systematic investigations of tinguished from other gastropods by (1) a conical fossil gastropods is the low preservative potential of shell with an anteriorly positioned apex (except lep- soft parts. Meanwhile, gastropod higher taxa are de- etid Propilidiinae having a posteriorly situated apex: fined primarily by the anatomy of soft tissue in the Lindberg, 1998: 649), (2) a thick horseshoe-shaped Recent groups. Phylogenetic analyses on Recent gas- muscle scar of the shell muscle with constricted outline 144 Takeshi Fuchigami and Takenori Sasaki and a thin scar of the pallial retractor muscle, (3) a to a thickness of 400 A˚ . The three-dimensional shell symmetrically little coiled protoconch (Sasaki, 1998), microstructures were observed in their radial and and (4) a characteristic scar in the apex of the tele- commarginal sections and inner shell surface with oconch after the protoconch is detached (cf. Sasaki, scanning electron microscopes (SEM, HITACHI S- 1998: figure 21g, h). Unless these characters are ob- 2400S in Department of Earth and Planetary Science, servable, it is not possible to detect the systematic po- University of Tokyo and HITACHI S-4500 in Uni- sition of limpets in question based on shell morphol- versity Museum, University of Tokyo). All specimens ogy only. In such a case, the shell structure characters observed with SEM are registered and preserved in must be useful, if their original structures are well the Department of the Historical Geology and Pale- preserved. ontology, University Museum, University of Tokyo The systematization of patellogastropod shell (UMUT) (Table 1). structure was first established by MacClintock (1967). The descriptive terminology used in this study basi- He investigated shell structures of 120 Recent and cally follows MacClintock (1967) and Carter et al. fossil patellogastropod species by optical microscopy (1990). The shell layers are divided into outer and in- and proposed 9 microstructures and 17 shell structure ner layers by a myostracum which corresponds to the groups. He also clarified a high degree of consistency insertion of muscles on the shell. The position of each between taxonomic categories and shell structure shell layer is indicated by its arrangement relative to groups. Lindberg (1988a) further generalized the sig- the myostracum (abbreviated as M). In this nomen- nificance of shell structural characters in patellogas- clature outer shell layers are described as Mþ1, Mþ2, tropod systematics, and this methodology has been Mþ3, etc. from the inner to the outer side, and like- broadly applied to the identification of fossil patello- wise inner layers as MÀ1, MÀ2, MÀ3, etc. from the gastropods by Lindberg and Hickman (1986), Lind- outer to the inner side. Each layer reclines at small berg (1988a, b), Lindberg and Marincovich (1988), angles with the inner surface, and its distribution is Lindberg and Squires (1990), Kase (1994), Kase visible as a concentric ring in ventral view (Figure 1). and Shigeta (1996), Lindberg and Hedegaard (1996), Some layers with identical microstructure were de- and Hedegaard et al. (1997). However, the existing scribed separately as ‘‘concentric’’ or ‘‘radial’’ layers knowledge on patellogastropod shell structure still under the criterion of its first-order unit arrangement. largely depends on the descriptions of MacClintock To avoid confusion, the four terms regarding ‘‘shell (1967) at the level of optical microscopy, although structure’’ were used in the following sense in this recent standards in studies on calcified hard tissue study. (1) Microstructure – The morphology of crystal essentially require SEM-level information. Hence, units and their mode of aggregation. (2) Shell layer – the reinvestigation of patellogastropod shell struc- A sheet-like component consisting of single micro- ture at finer resolution has been an unquestionable structure. (3) Shell structure – The total composition of primary subject in the studies of molluscan shell mi- microstructures and shell layers constituting the shell. crostructure. In this study, we attempted to provide (4) Shell structure group – The group of species having detailed SEM-level descriptions to advance the the identical order of shell layer arrangement and knowledge on patellogastropod shell structure. microstructure. In the descriptions the dip angle in- dicates the angle between growth axis of crystal units and inner shell surface, unless otherwise mentioned. Materials and methods The identification of crystal forms of calcium car- bonate (aragonite or calcite) using X-ray diffraction The materials used in this study include 44 species was not carried out in this study. Feigl stain (see Car- belongingto19generaof5families.Theywerecol- ter and Ambrose, 1989 for details), which can distin- lected alive mainly around Japan and some non- guish aragonite from calcite by staining in black, was Japanese species, especially type species of several preliminarily tested for some species but generally this genera, were also included for comparison (Table 1). method yielded poor results. Most shallow-water spe- Immediately after capture, the soft parts were re- cies are originally pigmented a dark color on their moved from the shell. The shell of each species was inner surfaces, and such a dark background color broken in approximately radial or transverse direc- hindered a clear identification of a series of tightly tion, and pieces of shell fragments were treated with stratified thin shell layers. Therefore, determination of bleach for 24 hours and etched with 3% acetic acid crystal forms for each microstructure of the respective for three seconds. After cleaning with an ultrasonic species is a subject for future study and was not re- cleaner, they were coated with platinum – palladium solved in this study. Shell
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