Scoyenia burrows from Ordovician palaeosols of the Juniata Formation in Pennsylvania

Scoyenia beerboweri is a new ichnospecies of burrow from the late Ordovician (Ashgill) Juniata Formation in central Pennsylvania, USA. The burrows are abundant in red calcareous palaeosols, and were created by animals living at the time of soil formation, because they are filled with red sediment like that of the palaeosol matrix, and both cut across, and are cut by, nodules of pedogenic carbonate. The isotopically light carbon and oxygen of carbonate in the palaeosols indicate a terrestrial ecosystem of well-drained floodplains in a tropical seasonally-dry semi-arid palaeoclimate. Backfill layering within the burrows is evidence of a bilaterally symmetrical animal. Size distribution of the burrows reveals discontinuous growth, as found in arthropods. Ferruginized faecal pellets in the burrows indicate that they ingested sediment. For these reasons the burrows of Scoyenia beerboweri are most likely to be the work of millipedes. The nature of vegetation supporting them is unknown, although a single problematic plant-like fossil cast was found, and liverwort spores are widespread in rocks of this age. Vegetative biomass was limited judging from the degree of chemical weathering, extent of burial gleization and isotopic composition of carbon in the palaeosols. These distinctive respiration-dominated liverwort-millipede polsterlands lived at a time of global greenhouse climate, following Precambrian–Cambrian lichen-algal microbial earths and supplanted by Silurian brakelands of early vascular land plants.