Aplacophora

Abstract: Molluscs (snails, octopuses, clams and their relatives) have a great disparity of body plans and, among the animals, only arthropods surpass them in species number. This diversity has made Mollusca one of the best-studied groups of animals, yet their evolutionary relationships remain poorly resolved. Open questions have important implications for the origin of Mollusca and for morphological evolution within the group. These questions include whether the shell-less, vermiform aplacophoran molluscs diverged before the origin of the shelled molluscs (Conchifera) or lost their shells secondarily. Monoplacophorans were not included in molecular studies until recently, when it was proposed that they constitute a clade named Serialia together with Polyplacophora (chitons), reflecting the serial repetition of body organs in both groups. Attempts to understand the early evolution of molluscs become even more complex when considering the large diversity of Cambrian fossils. These can have multiple dorsal shell plates and sclerites or can be shell-less but with a typical molluscan radula and serially repeated gills. To better resolve the relationships among molluscs, we generated transcriptome data for 15 species that, in combination with existing data, represent for the first time all major molluscan groups. We analysed multiple data sets containing up to 216,402 sites and 1,185 gene regions using multiple models and methods. Our results support the clade Aculifera, containing the three molluscan groups with spicules but without true shells, and they support the monophyly of Conchifera. Monoplacophora is not the sister group to other Conchifera but to Cephalopoda. Strong support is found for a clade that comprises Scaphopoda (tusk shells), Gastropoda and Bivalvia, with most analyses placing Scaphopoda and Gastropoda as sister groups. This well-resolved tree will constitute a framework for further studies of mollusc evolution, development and anatomy.

Abstract: Articulated halkieriids of Halkieria evangelista sp. nov. are described from the Sirius Passet fauna in the Lower Cambrian Buen Formation of Peary Land, North Greenland. Three zones of sclerites are recognizable: obliquely inclined rows of dorsal palmates, quincuncially inserted lateral cultrates and imbricated bundles of ventro-lateral siculates. In addition there is a prominent shell at both ends, each with radial ornamentation. Both sclerites and shells were probably calcareous, but increase in body size led to insertion of additional sclerites but marginal accretion of the shells. The ventral sole was soft and, in life, presumably muscular. Recognizable features of internal anatomy include a gut trace and possible musculature, inferred from imprints on the interior of the anterior shell. Halkieriids are closely related to the Middle Cambrian Wiwaxia, best known from the Burgess Shale: this clade appears to have played an important role in early protostome evolution. From an animal fairly closely related to Wiwaxia arose the polychaete annelids; the bundles of siculate sclerites prefigure the neurochaetae whereas the dorsal notochaetae derive from the palmates. Wiwaxia appears to have a relic shell and a similar structure in the sternaspid polychaetes may be an evolutionary remnant. The primitive state in extant polychaetes is best expressed in groups such as chrysopetalids, aphroditaceans and amphinomids. The homology between polychaete chaetae and the mantle setae of brachiopods is one line of evidence to suggest that the latter phylum arose from a juvenile halkieriid in which the posterior shell was first in juxtaposition to the anterior and rotated beneath it to provide the bivalved condition of an ancestral brachiopod. H. evangelista sp. nov. has shells which resemble those of a brachiopod; in particular the posterior one. From predecessors of the halkieriids known as siphogonuchitids it is possible that both chitons (polyplacophorans) and conchiferan molluscs arose. The hypothesis of halkieriids and their relatives having a key role in annelid-brachiopod-mollusc evolution is in accord with some earlier proposals and recent evidence from molecular biology. It casts doubt, however, on a number of favoured concepts including the primitive annelid being oligochaetoid and a burrower, the brachiopods being deuterostomes and the coelom being an archaic feature of metazoans. Rather, the annelid coelom arose as a functional consequence of the transition from a creeping halkieriid to a polychaete with stepping parapodial locomotion.

Abstract: Stasek (1) theorized that the extant mollusks are the progeny of three separate lineages that separated before the phylum was well established. He wrote that no known intermediate forms, fossil or living, bridge the "enormous gaps between any two of the three lineages," and therefore treated each as a separate subphylum. These subphyla are (i) the subphylum Aculifera Hatscheck 1891, containing only the class Aplacophora, derived from the most primitive ancestors of the Mollusca; (ii) the subphylum Placophora von Jhering 1876, containing only the class Polyplacophora, and emphasizing the pseudometamerism of its more advanced premollusk ancestor; and (iii) the subphylum Conchifera Gegenbaur 1878, containing the Monoplacophora and the other classes derived from it. We point out that the Polyplacophora may be derived from the Monoplacophora instead of a more primitive ancestral stock. We also suggest that the Conchifera can be separated into two major lineages, each worthy of the rank of subphylum. The fossil record indicates that the Monoplacophora gave rise to the Gastropoda, Cephalopoda, Rostroconchia, and possibly Polyplacophora, and that the Pelecypoda and Scaphopoda are derived from the Rostroconchia. These last three classes thus form a lineage that diverged from the Monoplacophora in the Early Cambrian. They emphasized a shell form that in all groups is primitively open at both ends, allowing the gut to remain relatively straight, with an anterior mouth and posterior anus. They became burrowing (infaunal) deposit or filter feeders. We coin the term Diasoma (through-body) for the subphylum containing these three classes (Rostroconchia, Pelecypoda, and Scaphopoda). The remaining three classes (Monoplacophora, Gastropoda, and Cephalopoda) emphasize a conical univalved shell, usually twisted into a spiral. The relatively small single aperture forces the anus to lie close to the mouth, and the gut is bent into a "U." Most are surface-dwelling (epifaunal) grazers or carnivores. We coin the name Cyrtosoma (hunchback-body) for the subphylum containing these three classes. Strictly speaking, the cyrtosomes are the ancestors of the diasomes but, in fact, both subphyla appeared and began to diversify within a few million years in the Early Cambrian. Note added in proof: After proofs were corrected we were informed that the new genus Opikella (40) is preoccupied by (Opikella = Oepikella) Thorslund 1940, an Ordovican ostracod. We rename the mollusk genus Oepikila.