Nacre was previously thought to be primitive in the Mollusca, but no convincing Cambrian examples are known. This aragonitic microstructure with crystal tablets that grow within an organic framework is thought to be the strongest, most fracture-resistant type of shell microstructure. Fossils described herein from the Ordovician of Iowa, Indiana, and Ohio provide supporting evidence for the hypothesis that sometime between the middle Cambrian and late Ordovician, nacre originated in cephalopod, bivalve, and possibly gastropod lineages. The correlation of independent origins of fracture-resistant nacre with increasing shell-crushing abilities of predators during the Cambrian-Ordovician suggests an early pulse in the evolutionary arms race between predators and molluscan prey.

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Commensal epizoozoans and episkeletozoans are rarely preserved attached to the external exoskeleton of the Late Ordovician trilobite Flexicalymene. Of nearly 15,000 Flexicalymene specimens examined, 0.1% show epizoozoans or episkeletozoans. Factors limiting Flexicalymene fouling include a shallow burrowing life style, frequent molting of the host, larval preference for other substrates, observational bias caused by overlooking small fouling organisms, and the loss of the non-calcified, outermost cuticle prior to fossilization or as the trilobite weathers from the encasing sediment. Trepostome bryozoans, articulate and inarticulate brachiopods, cornulitids, and a tube-dwelling/boring nonbiomineralized organism represent the preserved members of the Late Ordovician marine hard substrate community fouling Flexicalymene. This assemblage of organisms is less diverse than the hard substrate community fouling Late Ordovician sessile epibenthic organisms. Fouling is not restricted to only large Flexicalymene specimens as observed in previous studies but occurs in medium to large individuals interpreted as early to late holaspid specimens.Epizoozoans fouling the carcasses or molt ensembles of 16 Flexicalymene specimens provide insight into the life habits of the host and these fouling organisms. Trepostome bryozoans, articulate and inarticulate brachiopods, and cornulitids preferentially attached to elevated portions of the dorsal exoskeleton, and preferentially aligned in either the direct line or lee side of currents generated by Flexicalymene walking on the sea floor or swimming through the water column.

Exceptional sub-micrometer details of shell microstructure are preserved in phosphatic micro-steinkerns representing several phyla from shell beds of the Upper Ordovician of the Cincinnati Arch region, USA. These fossils provide the most detailed record of Ordovician mollusk shell microstructures, as well as exceptional details on the earliest cases of undisputed nacre. The trend towards nacre in the Mollusca is one aspect of the surge in escalation between mollusks and their predators during the Great Ordovician Biodiversification Event.

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