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Fossil record

Good evidence exists for the appearance of gastropods (e.g., Aldanella), cephalopods (e.g., Plectronoceras, Nectocaris ?) and bivalves (Pojetaia, Fordilla) towards the middle of the Cambrian period, c. 500 million years ago, though arguably each of these may belong only to the stem lineage of their respective classes. However, the evolutionary history both of the emergence of molluscs from the ancestral group Lophotrochozoa, and of their diversification into the well-known living and fossil forms, is still vigorously debated.

Debate occurs about whether some Ediacaran and Early Cambrian fossils really are molluscs. Kimberella, from about 555 million years ago, has been described by some paleontologists as "mollusc-like", but others are unwilling to go further than "probable bilaterian ", if that.

There is an even sharper debate about whether Wiwaxia, from about 505 million years ago, was a mollusc, and much of this centers on whether its feeding apparatus was a type of radula or more similar to that of some polychaete worms. Nicholas Butterfield, who opposes the idea that Wiwaxia was a mollusc, has written that earlier microfossils from 515 to 510 million years ago are fragments of a genuinely mollusc-like radula. This appears to contradict the concept that the ancestral molluscan radula was mineralized.

However, the Helcionellids, which first appear over 540 million years ago in Early Cambrian rocks from Siberia and China, are thought to be early molluscs with rather snail-like shells. Shelled molluscs therefore predate the earliest trilobites. Although most helcionellid fossils are only a few millimeters long, specimens a few centimeters long have also been found, most with more limpet -like shapes. The tiny specimens have been suggested to be juveniles and the larger ones adults.

Some analyses of helcionellids concluded these were the earliest gastropods. However, other scientists are not convinced these Early Cambrian fossils show clear signs of the torsion that identifies modern gastropods twists the internal organs so the anus lies above the head.

Volborthella, some fossils of which predate 530 million years ago, was long thought to be a cephalopod, but discoveries of more detailed fossils showed its shell was not secreted, but built from grains of the mineral silicon dioxide (silica), and it was not divided into a series of compartments by septa as those of fossil shelled cephalopods and the living Nautilus are. Volborthella ' s classification is uncertain. The Middle Cambrian fossil Nectocaris is often interpreted as a cephalopod with 2 arms and no shell, but the Late Cambrian fossil Plectronoceras is now thought to be the earliest undisputed cephalopod fossil, as its shell had septa and a siphuncle, a strand of tissue that Nautilus uses to remove water from compartments it has vacated as it grows, and which is also visible in fossil ammonite shells. However, Plectronoceras and other early cephalopods crept along the seafloor instead of swimming, as their shells contained a "ballast" of stony deposits on what is thought to be the underside, and had stripes and blotches on what is thought to be the upper surface. All cephalopods with external shells except the nautiloids became extinct by the end of the Cretaceous period 65 million years ago. However, the shell-less Coleoidea (squid, octopus, cuttlefish) are abundant today.

The Early Cambrian fossils Fordilla and Pojetaia are regarded as bivalves. "Modern-looking" bivalves appeared in the Ordovician period, 488 to 443 million years ago. One bivalve group, the rudists, became major reef -builders in the Cretaceous, but became extinct in the Cretaceous–Paleogene extinction event. Even so, bivalves remain abundant and diverse.

The Hyolitha are a class of extinct animals with a shell and operculum that may be molluscs. Authors who suggest they deserve their own phylum do not comment on the position of this phylum in the tree of life.