Paleozoic echinoderm hangovers: waking up in the Triassic

Echinoderms are among the marine invertebrates that underwent the most severe losses at the end-Permian extinction. The prevailing paradigm claims an extreme bottleneck with only very few, if not single, holdovers (“hangovers” herein) sparking the post-Paleozoic radiation. Here we identify previously overlooked Triassic echinoids, ophiuroids, and asteroids as unambiguous members of Paleozoic stem groups. These echinoderm hangovers occurred almost worldwide and had spread into a wide range of paleoenvironments by the Late Triassic. Our discovery challenges fundamentals of echinoderm evolution with respect to end-Permian survival and sheds new light on the early evolution of the modern clades, in particular on Triassic ghost lineages (i.e., inferred but undocumented fossil record) of the crown-group look-alikes of the Paleozoic hangovers.