Tardigrade stowaways: Literature review of Propyxidium tardigradum (Ciliophora, Peritrichia) and its first record in Scotland

Tardigrades are a phylum of microscopic invertebrates with a global distribution. Although our understanding of their systematic position and taxonomy has increased and continues to grow, their relationship with the other organisms that share their habitat remains poorly studied. One such organism is Propyxidium tardigradum, a peritrich ciliate which uses tardigrades for dispersion and as a substrate for reproduction. Here, we present the first Scottish record and tenth global occurrence of Propyxidium tardigradum, thereby expanding our knowledge of its poorly understood zoogeographic distribution. We also summarise the literature concerning P. tardigradum biology, proffer hypotheses regarding the Propyxidium-tardigrade relationship, and the apparent lack of heterotardigrade ciliate infestation. Additionally, we indicate a number of recommendations for the direction of future studies regarding the ciliate. Finally, we add a further three species, Milnesium variefidum, Hypsibius cf. scabropygus and Macrobiotus scoticus to the list of Propyxidium host species.