Palaeoecology and diversity of endosymbionts in Palaeozoic marine invertebrates: Trace fossil evidence

Endosymbionts are organisms that live within the growing skeleton of a live host organism, producing a cavity called a bioclaustration. The endosymbiont lives inside the bioclaustration, which it forms by locally inhibiting the normal skeletal growth of the host, a behaviour given the new ethological category, impedichnia. As trace fossils, bioclaustrations are direct evidence of past symbioses and are first recognized from the Late Ordovician (Caradoc). Bioclaustrations have a wide geographic distribution and occur in various skeletal marine invertebrates, including tabulate and rugose corals, calcareous sponges, bryozoans, brachiopods, and crinoids. Ten bioclaustration ichnogenera are recognized and occur preferentially in particular host taxa, suggesting hostspecificity among Palaeozoic endosymbionts. The diversity of bioclaustrations increased during the Silurian and reached a climax by the late Middle Devonian (Givetian). A collapse in bioclaustration diversity and abundance during the Late Devonian is most significant among endosymbionts of host coral and calcareous sponge taxa that were in decline leading up to the Frasnian–Famennian mass extinction.