The biology and functional morphology of Humphreyia strangei (Bivalvia: Anomalodesmata: Clavagellidae): an Australian cemented ‘watering pot’ shell

Abstract

Amongst the watering pot shells of the Clavagellidae (Anomalodesmata), the least well known is Humphreyia, with a single species H. strangei. Unlike representatives of the other extant clavagellid genera, i.e. Clavagella and Brechites, however, where juveniles are unknown, there is a single juvenile (plus adult specimens) of H. strangei available for study. The Clavagellidae contains the weirdest of all bivalves, encased in a huge (in proportion to the tiny juvenile shell) adventitious tube with an anterior ‘watering pot’. The juvenile H. strangei is dimyarian with a large foot and byssal groove, and a large pedal gape; the entire body is enclosed within a bag-like periostracum-covered mantle cavity. Upon permanent residence, however, the anterior watering pot component of the adventitious tube is cemented to the chosen substratum, the adductor muscles are lost and the foot and pedal gape greatly reduced; the fourth pallial aperture closes and the pedal gape and the tubules of the watering pot are similarly occluded. Once cementation is achieved, further growth anteriorly is impossible and the adventitious tube is secreted posteriorly, to house the long siphons, and subsequently can be added to as growth increments. Anatomically, H. strangei has a typical anomalodesmatan arrangement of mantle cavity and digestive organs so that it is, probably, a suspension feeder. Inside the pericardium, however, is a pair of unique proprioreceptors which probably serve to monitor rectal tonus and thus control defecation, co-ordinated with siphonal retraction and extension. They probably also prevent over-filling of the capacious rectum. It is believed that cementation and adventitious tube formation occur at the time of sexual maturity and this change in lifestyle between juvenile and adult represents a form of metamorphosis – quite distinct from that which occurs in all bivalves between the pediveliger and juvenile stages – and seems to be unique to Humphreyia and probably Brechites. This family of anomalodesmatan bivalves is thus actually stranger than the already aberrant watering pot would suggest.