Protobranch bivalves

The subclass Protobranchia comprises more than 600 species of bivalves that occur throughout the world ocean. Mostly deposit feeders in soft sediments, they are abundant in the deep sea. Apomorphies that unite them as a group include gill structure, hinge conformation, shell microstructure, larval development, foot morphology, respiratory pigments, trophic mode and digestion. They are relatively small and highly conserved in form, originating in the Cambrian era. They may represent an ancestral, derived or paraphylectic group of the Bivalvia. The protobranchs include two orders, the Nuculoida and Solemyoida, which previously were classified separately in the subclasses Paleotaxodonta and Cryptodonta, respectively. They are of ecological interest and have a unique functional morphology. They feed mostly under the surface of the sediment with highly modified labial palps, but the degree to which they are selective in diet remains difficult to determine. They are important bioturbators in many soft-sediment assemblages; their feeding and locomotion affects sediment structure and community development. Solemyoids are unusual in inhabiting reducing environments and hydrocarbon seeps and in deriving their nutrition from endosymbiotic chemosynthetic bacteria. A variety of species of protobranchs are found in oceanic trenches, near hydrothermal vents, and in submarine caves. Protobranchs produce a lecithotrophic larval stage, the pericalymma, making their development unique among bivalves. The pericalymma remains in the plankton for a short time and presumably has low dispersal ability. Recruitment may be intermittent. Growth is rapid in post-larvae but decreases with age, though rates may not necessarily be slow, especially in continental shelf species. Life spans are commonly 1 to 2 decades, but deep-sea representatives may grow more slowly and live longer. Bottom fish, seastars and gastropods are their major predators and a few parasites and commensals have been documented. The predominance of protobranchs in deep-sea sediments may be a result of deep-sea origin or displacement from shallow waters by lamellibranchs. Their ability to deposit-feed, digest food extracellularly, and develop by means of lecithotrophic larvae make them particularly well adapted to cold and oligotrophic habitats.