a muscular gizzard lined with chitinous teeth that crush armoured prey such as diatoms. Wave-like peristaltic contractions then move the food through the stomach for digestion. Molluscs have a structure called a radula which bears a ribbon of chitinous teeth. However, these teeth are histologically and developmentally different from vertebrate teeth and are unlikely to be homologous. For example, vertebrate teeth develop from a neural crest mesenchyme-derived dental papilla, and the neural crest is specific to vertebrates, as are tissues such as enamel. The radula is used by molluscs for feeding and is sometimes compared rather inaccurately to a tongue. It is a minutely toothed, chitinous ribbon, typically used for scraping or cutting food before the food enters the oesophagus. The radula is unique to molluscs, and is found in every class of mollusc apart from bivalves. Within the gastropods, the radula is used in feeding by both herbivorous and carnivorous snails and slugs. The arrangement of teeth (also known as denticles) on the radula ribbon varies considerably from one group to another as shown in the diagram on the left. Predatory marine snails such as the Naticidae use the radula plus an acidic secretion to bore through the shell of other molluscs. Other predatory marine snails, such as the Conidae, use a specialized radula tooth as a poisoned harpoon. Predatory pulmonate land slugs, such as the ghost slug, use elongated razor-sharp teeth on the radula to seize and devour earthworms. Predatory cephalopods, such as squid, use the radula for cutting prey. In most of the more ancient lineages of gastropods, the radula is used to graze by scraping diatoms and other microscopic algae off rock surfaces and other substrates. Limpets scrape algae from rocks using radula equipped with exceptionally hard rasping teeth. These teeth have the strongest known tensile strength of any biological material, outperforming spider silk. The mineral protein of the limpet teeth can withstand a tensile stress of 4.9 GPa, compared to 4 GPa of spider silk and 0.5 GPa of human teeth. Fossilization and taphonomy Because teeth are very resistant, often preserved when bones are not, and reflect the diet of the host organism, they are very valuable to archaeologists and palaeontologists. Early fish such as the thelodonts had scales composed of dentine and an enamel-like compound, suggesting that the origin of teeth was from scales which were retained in the mouth. Fish as early as