Paula C. Dentzien-Dias, George Poinar, Ana Emilia Q. de Figueiredo, Ana Carolina L. Pacheco, Bruno L. D. Horn, Cesar L. Schultz

Remains of parasites in vertebrates are rare from the Mesozoic and Paleozoic. Once most parasites that live in - or pass through - the gastrointestinal tract of vertebrates, fossil feces (coprolites) or even intestinal contents (enterolites) can eventually preserve their remains. Here we announce the discovery of a spiral shark coprolite from the Paleozoic bearing a cluster of 93 small oval-elliptical smooth-shelled structures, interpreted as eggs of a tapeworm.The eggs were found in a thin section of an elasmobranch coprolite. Most of the eggs are filled by pyrite and some have a special polar swelling (operculum), suggesting they are non-erupted eggs. One of the eggs contains a probable developing larva. The eggs are approximately 145-155 µm in length and 88-100 µm in width and vary little in size within the cluster. The depositional and morphological features of the eggs closely resemble those of cestodes. Not only do the individual eggs have features of extant tapeworms, but their deposition all together in an elongate segment is typical to modern tapeworm eggs deposited in mature segments (proglottids). This is the earliest fossil record of tapeworm parasitism of vertebrates and establishes a timeline for the evolution of cestodes. This discovery shows that the fossil record of vertebrate intestinal parasites is much older than was hitherto known and that the interaction between tapeworms and vertebrates occurred at least since the Middle-Late Permian.