School of Life Sciences, The University of Warwick, Gibbet Hill Campus, Coventry, United Kingdom
a School of Biological Sciences, The University of Hong Kong, Hong Kong, SAR, China
b Department of Animal Biology, University of Campinas (Unicamp), Brazil
Abstract
Most bivalves are suspension feeders. On the deep sea floor, however, some are predators, typically of meiobenthic crustaceans: copepods, cumaceans and ostracods. Propeamusiid scallops are one such group of predators. The largest numbers of predators, however, belong to the bivalve subclass Anomalodesmata and constitute, as currently recognised, some 500 species belonging principally to the Verticordioidea (120), Poromyoidea (75) and Cuspidarioidea (304) with four, two and four constituent families, respectively. A further family, the Parilimyidae, is considered to be derived from the Pholadomyoideathe anomalodesmatan ancestor. These, generally small (< 60 mm shell length), nacreous and thin-shelled predators share many anatomical features that formerly allowed them to be collectively classified as the Septibranchia. Although this name is now rarely used, it refers to their possession of a ctenidially-derived septum in the mantle cavity and functioning in prey capture. Generally, there is a trend, possibly evolutionary, from a typical bivalve ctenidium (Parilimyidae and some Verticordioidea) to a complete septum (other Verticordioidea, Poromyoidea and Cuspidarioidea). In addition, the inhalant siphon, foot, labial palps, mouth and its lips play a role in prey capture, and ingestion. Similarly, the stomach is modified to digest such, typically chitinous, ingested prey. Most septibranchs are either consecutive or simultaneous hermaphrodites with self-fertilisation possibly usual and with some evidence in a few of larval brooding.
Notwithstanding, the deep sea septibranch species are poorly studied with virtually nothing being known about their wider distributions, ecology, detailed reproductive strategies and life history traits.
Keywords
Anomalodesmata; Bentholyonsiidae; Carnivorous lifestyle; Ctenidia; Evolution; Prey; Phylogeny; Septum
1 Introduction
Considered the second most diverse class of the Mollusca, it is currently estimated that there are between 8000 and 20,000 species of Bivalvia Linnaeus, 1758 contained in 106 families (. Of the, as currently considered, 106 families there are, at present, 1257 known species (13.7%) belonging to 12 (11.3%) families occurring in freshwater habitats. The remainder (86.3%) is marine and here they occupy all latitudes and virtually all habitats from the intertidal to the abyss.
The bivalves first evolved in the geologically brief period of the Cambrian explosion, which took place between 540 and 520 million years ago ().
Subsequent to the Ordovician, the bivalves have radiated to occupy most aquatic habitats and on hard and soft substrata the numbers of individuals can be enormous. For example, on a large beach in South Wales in the United Kingdom, reported that on a North Yorkshire rocky coast in the United Kingdom, the numbers of newly settled plantigrade individuals among an established population of Mytilus edulis Linnaeus, 1758 varied between 100 and 1000 per 5 cm2. Much of this abundance and success relates to the filter feeding and gregarious behaviour of many bivalves and a generalised reproductive strategy of lecithotrophy in which the external fertilisation of huge numbers of gametes and the development of planktonic larvae can produce an equally large number of settled juveniles.
The great majority of bivalves are restricted to fresh waters and intertidal and shallow subtidal habitats and ).
In a major review of the living bivalves of the Eastern Pacific Ocean all the way from the Bering Strait to Cape Horn and extending out into waters around the Galapagos Islands, recording 168 bivalves from these cold waters and of these but 2% (~ 3) were septibranchs.
The waters around the Canary Islands support a large number of septibranchs: Verticordiidae (6), Poromyidae (1) and Cuspidariidae (15) ().
The septibranchs thus have a global distribution on and, especially, in the deep sea floor raising questions of as yet unstudied cosmopolitanism and genetic diversity at abyssal and, even, hadal depths. And, the septibranchs, plus other deep water but non-predatory anomalodesmatans, thus constitute some 22% of the deep water bivalve fauna with the predators making up approximately 20% of them. It is these that are the principal subject of this review.