The biology and functional morphology of Nucula pusilla (Bivalvia: Protobranchia: Nuculidae) from Western Australia, Australia: primitive or miniature simplicity?

Abstract

The southern Australian nuculid Nucula pusilla is < 3 mm in shell length, making it one of the smallest known protobranchs and one of the smallest bivalves. It lives in clean, well-aerated, coarse, offshore marine sands in the southern half of the Australian continent. The bivalve lies in the sand with the pointed anterior end of the shell directed upwards at an angle of ~45° to the sediment surface and inhales water into the mantle cavity from this direction. The more rounded posterior shell margin sits within the sediment. The posterior exhalant current, created by the posterior, largely respiratory ctenidia assist in the removal, along with ciliary currents on the mantle surface and visceral mass, of unwanted particles. Nucula pusilla possesses a pair of large, closed statocysts with numerous statoconia, situated above the pedal ganglia and these are likely to be responsible for the bivalve’s orientation. There is also a minute Stempel’s organ located on the outer face of the anterior adductor muscle. Feeding is largely by means of palp proboscides that collect subsurface material and transport it along anterior ciliated grooves where it reaches and is sorted by the labial palps where some particles are accepted and others are rejected. The intestine is coiled complexly in the visceral mass but the style sac is much simplifi ed in comparison with N. sulcata. The