Symbiotic embedment structures in Silurian Caryocrinites (Echinodermata, Rhombifera, Hemicosmitida)

Abstract
A variety of pits representing symbiotic embedments, sometimes associated with pathological deformation in the host, are known from the skeletons of Paleozoic stalked echinoderms. These structures are well known from multiple genera of crinoids and a limited number of blastozoans but have not previously been described in detail from the skeletons of rhombiferans. This is surprising given the abundance of rhombiferans in certain deposits, the co-occurrence of rhombiferans with frequently infested taxa, including diploporitans, in multiple assemblages, and the morphological similarity between certain rhombiferan taxa and coeval infested crinoids. The common hemicosmitid rhombiferan Caryocrinites Say, 1825 is widespread throughout the middle Silurian of eastern North America and is herein reported to contain symbiotic (potentially parasitic) embedment structures. Specimens were collected from the lower portion of the mudstone lithofacies of the Massie Formation (Wenlock, Sheinwoodian) at the Napoleon quarry of southeastern Indiana, USA. Strong host specificity is indicated by the absence of pits in C. ornatus Say, 1825 and exclusive infestation of a smaller co-occurring species of Caryocrinites. Thecae with embedment structures are consistently smaller than thecae without such structures, with pitted specimens being restricted to a narrow range of thecal heights (20–24 mm). All embedment structures are present only on the proximal portion of thecae, with individual specimens containing between one and 30 pits. No elevated rims or significant swelling were observed on any specimens, and all pits are relatively small (~1 mm in diameter). The presence of symbiotic embedment structures represents an additional example of a crinoid-like aspect to the ecology of Caryocrinites.