Compared to modern cephalopods, the fossil record reveals 20 times the number of species. The first cephalopods appeared during the late Cambrian (about 500 million years ago). They looked like squid, but with long, cone-shaped shells from which they extended their tentacles. Typically, the shells were divided into chambers (septa), added on as the animal grew. Some early cephalopods were enormous, as long as a school bus. For animals without backbones (invertebrates), they were sophisticated, with nervous systems and the ability to swim by jet propulsion. By the end of the Paleozoic (250 million years ago), cephalopods had diversified to include coiled shells like a modern nautilus. While the extinction event at the end of the Paleozoic impacted many species, cephalopods flourished again during the Triassic and Jurassic. The extinction event at the end of the Cretaceous (65 million years ago) left only the nautiloids and the coleoids (octopus and squid).

Mollusks have inhabited the Earth for at least 540 million years, dating back to the early Cambrian period. The fossil record of mollusks consists mostly of shells. Rarely, it includes mouthparts (radulas) or the trapdoors (opercula) that cover shell openings. The rest of the body is soft and usually does not fossilize. Before the Cambrian, taxa such as Kimberella had many traits expected of early molluscs, but no shells or spines. Cambrian, worm-like animals with spiny scales, such as Wiwaxia, had mollusk-like mouthparts. The earliest agreed-upon mollusk looked like a snail with a single, cap-like shell that curved at the tip. More than 90,000 living species of mollusk have been described, plus another 70,000 species known only from fossils. Because many mollusks live in remote places such as in the bottom sediments of deep ocean, and new species are being discovered at a rapid rate, scientists think that many more species remain to be described.