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Some Aspects of the Biology and Functional Morphology of Trapezium (Neotrapezium) Sublaevigatum (Lamarck) (Bivalvia: Arcticacea)L

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Some Aspects of the Biology and Functional Morphology of Trapezium (Neotrapezium) Sublaevigatum (Lamarck) (Bivalvia: Arcticacea)L Pacific Science (1979), vol. 33, no. 2 © 1980 by The University Press of Hawaii. All rights reserved Some Aspects of the Biology and Functional Morphology of Trapezium (Neotrapezium) sublaevigatum (Lamarck) (Bivalvia: Arcticacea)l BRIAN MORTON 2 ABSTRACT: Trape::ium sublael'igalum is a widely distributed nestler in rocky intertidal and sublittoral crevices in Hong Kong. It occurs on a range of beach types and in waters of different salinity; it is byssally attached and weakly heteromyarian in form. The quadrangular shell is often distorted. The basic structure of the shell and the morphology of the organs of the mantle cavity, visceral mass, and pericardium are broadly similar to a wide range of veneroid relatives. However, the species is weakly heteromyarian, which results from the assumption of the byssally attached, epifaunal mode of life. There is thus a reduction of the anterior face of the shell and an enlargement of the posterior face with corresponding modifications to the shell (notably hinge teeth and ligament) and to musculature. Similar, convergent adaptations are seen in members of the veneroid superfamilies Carditacea, Gaimardiacea, and Dreis­ senacea, with which Trapezium can be compared. Further study of other members of the (isomyarian) Arcticacea might help to establish how the heteromyarian form arose in Trapezium. THE EPIFAUNAL BIVALVE colonizers of the (Carditacea) (Yonge 1969) and the cemented rocky intertidal and sublittoral are variously Spondylus (Spondylidae), Plieatula (Pli­ adapted to their mode of life. Many are catulidae) (Yonge 1973), and Dimya (Dimyi­ permanently byssally attached, whereas dae) (Yonge 1975) occur. others are only temporarily attached as larvae Aside from the bivalves associated com­ prior to the adoption of, for example, a mensally with other organisms, such as the cemented life-style. colonizers of compound ascidians, e.g., On exposed shores many genera are either Ryenella (Mytilacea) and Mytilimeria (Ano­ heteromyarian in form, e.g., Mytilus and malodesmata) (Yonge 1952), and the borers Septifer (Mytilacea) (White 1937, Yonge and of soft rocks and shales, e.g., Barnea and Campbell 1968); laterally flattened, e.g., Pholas (Pholadidae) (Purchon 1955b) and Anomia (Anomiacea) (Yonge 1977); or ce­ Petrieola (Petricolidae) (Purchon 1955a), a mented, e.g., Chama (Chamidae) (Yonge third major category of epifaunal bivalves 1967) and Cleidothaerus (Anomalodesmata) are the nestlers. Nestlers are usually small, (Morton 1974). On less exposed, often es­ secretive, byssally attached bivalves, which tuarine boulder- or cobble-strewn beaches, are hidden in crevices, in the holdfasts of bivalves with similar habits are found, e.g., algae or among masses of larger bivalves. the heteromyarian Braehidontes and Xeno­ Representatives from a wide range offamilies strobus (Mytilacea) (Wilson 1967) and the fall into this category, such as members of cemented oysters Ostrea, Crassostrea, and the Arcacea (Barbatia, Area), Mytilacea Saeeostrea (Ostreacea) (Yonge 1953). Simi­ (Modiolus), and Saxicavacea (Hiatella) larly in the sublittoral, the byssally attached (Yonge 1971), together with delegates from heteromyarian bivalves Cardita and Beguina a range of families belonging to the vast, cosmopolitan, and highly successful order Veneroida. These include members of the I Manuscript accepted 6 November 1978. 2 University of Hong Kong, Department of Zoology, Leptonacea (Lasaea) (Oldfield 1955), Gai­ Pokfulam Road, Hong Kong. mardiacea (Neogaimardia) (Morton 1979), 177 178 PACIFIC SCIENCE, Volume 33, April 1979 Veneracea (Venerupis) (Ansell 1961), Petri­ of Coralliophaga are coral-associated nes­ colidae (Claudiconcha) (Morton 1978), and tIers; Trapezium (Glossocardia) obesa is a Trapezium (Arcticacea, Trapeziidae). deepwater species while the two species of The arcticiid Trapezium sublaevigatum is Trapezium (Neotrapezium) inhabit crevices closely allied to a wide range of infaunal of rocks or oysters in shallow, intertidal animals which have not been adequately waters. studied. The present study seeks information on T. sublaevigatum in order to supply infor­ mation on the superfamily and also to com­ BIOLOGY pare the adaptations of T. sublaevigatum with those of other nestling bivalves about Solem (1954) considers Trapezium sub­ which more is known and which have a laevigatum to be a continental or large island similar mode of life. species because, although it is widely dis­ tributed in the Indo-Pacific, it has only rarely been recorded from offshore islands. Solem's review of the literature pertaining to the MATERIALS AND METHODS ecology of this species establishes that it is The ciliary currents ofdissected individuals byssally attached within the crevices of rocks were studied using carmine suspended in or oysters in the intertidal zone. Nothing is seawater. Specimens for sectioning were fixed known ofthe adaptations of T. sublaevigatum in Bouin's fluid, sectioned at 6 .um, and (or of any other trapeziid) to its particular stained in Ehrlich's hematoxylin, Heiden­ mode of life. hain's hematoxylin, or Mallory's triple stain. Trapezium sublaevigatum is widely dis­ tributed in Hong Kong and is common in the many small bays characteristic of the coastline. Itis a nestler that is usually attached TAXONOMY by a stout byssus to the undersurface ofstones The veneroid superfamily Arcticacea New­ and boulders of the lower shore. Where it ton 1891 comprises nine families (Keen 1969) nestles in crevices there is often a resulting which have been little studied. In the Tra­ distortion of the shell. The habitat always peziidae Lamy 1920 there are seven genera, has a freshwater input and is sheltered from of which only Trapezium Megerle von Miihl­ strong wave action. The species is commonly feld 1811 and Coralliophaga de Blainville 1824 found attached to cultivated oysters in Deep are extant. Both genera have subgenera and Bay, where salinity can be very low and the the species reported upon here is the type sediment load high (Wong 1975); it also species for the subgenus Neotrapezium Habe occurs on experimental panels as a member 1951. This was originally described as Cardita of the fouling community. The specimens sublaevigata Lamarck 1819; the corrected reported here were collected mainly from name for the species is Trapezium (Neo­ sublittoral boulders at the mouth of the bay trapezium) sublaevigatum (Lamarck 1819). at Hoi Sing Wan, near the village of Wu Previous reviewers of the family include Kwai Sha, Tolo harbor, or from sublittoral Reeve (1843), Lamy (1920), and Solem (1954). dead coral heads, similarly collected from The latter author comprehensively described Tolo harbor. the known species and summarized data on the ecology of the family. Solem describes a total of nine extant species of Trapezium (6) and Coralliophaga (3) plus the somewhat FUNCTIONAL MORPHOLOGY dubious Isorropodon perplexum Sturany 1896 The Shell and Ligament from 2420 m in the Mediterranean, which has not been reported since. The three species The shell of Trapezium sublaevigatum of Trapezium (Trapezium) and three species (Figure lA) is approximately quadrangular . ;&1¥-@- W$eeIJ i R $i <$ J! W a &Z Trapezium (Neotrapezium) sublaevigatum (Lamarck)-MoRTON 179 A c I --- --- -- - -,------- I I I I POL HP I IL r I I I I I ,I I, CT PA--nr-~~ ABR AA 10mm FIGURE 1. Trapezium sublaevigatum. A, exterior view of the right shell valve; B, interior view of the left shell valve; C, end-on view showing the widest region ofthe shell. AA, anterior adductor muscle scar; ABR, anterior byssal retractor muscle; AOL, anterior outer ligament layer; CT, cardinal tooth; HP, hinge plate; IL, inner ligament layer; PA, posterior adductor muscle scar; PBR, posterior byssal retractor muscle; PL, pallial line; PLT, posterior lateral tooth; POL, posterior outer ligament layer; PS, pallial sinus. in appearance. There is no radial or con­ myarian form with the anterior slope of the centric sculpture, though there are often shell reduced relative to the posterior. Thus, distinct growth rings. The shell is delicately the umbones are located close to the anterior radially patterned, typically with nine dark­ end of the shell. In transverse section (Figure brown stripes. The shell is equivalve and I C) the greatest shell width is located dorsal markedly inequilateral, a condition resulting to the midpoint of the dorsoventral axis of from the assumption of a weakly hetero- the shell. Other epifaunal heteromyarian 180 PACIFIC SCIENCE, Volume 33, April 1979 HP CT IL CT POL u ~----u CT CT PLT POL HP FIGURE 2. Trapezium sub/aevigatum. The hinge plate ofthe right (above) and the left (below) shell valves. CT, cardinal tooth; HP, hinge plate; IL, inter ligament layer; PLT, posterior lateral tooth; POL, posterior outer ligament layer; U, umbo. bivalves, e.g., members of the Mytilacea and the cardinal teeth. Similarly, the absence of Dreissenacea, are much wider ventrally. anterior lateral teeth can be accounted for There is no byssal notch. by the reduction of the anterior slope of the Internally (Figure IB), the effect of the shell. heteromyarian form upon the shell is more The ligament (Figure 2) is opisthodetic and apparent. The posterior adductor (PA) and external. It comprises a posterior outer liga­ posterior byssal retractor muscles (PBR) are ment layer (POL) overlying the inner liga­ relatively larger than their anterior equiva­ ment layer (lL). This in turn overlies a small lents (AA and ABR). The pallial line (PL) anterior outer ligament layer that is often is thick and has a shallow pallial sinus (PS). obscured by erosion ofthis region ofthe shell. The hinge plate (Figures 1,2, HP) is narrow, with a number of hinge teeth that can be The Siphons interpreted as comprising either two (the posterior of which is bicuspid) or three The siphons of Trapezium sublaevigatum cardinal teeth (CT) and a single posterior (Figure 3) are relatively short and are re­ lateral tooth(PLT) located behind the liga­ tracted into a short pallial sinus. They are ment. The assumption of the heteromyarian formed by fusion of the inner folds only and form with the concomitant enhanced de­ are thus of type A (Yonge 1948, 1957).
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