Spatial and Temporal Distribution of the Island-Dwelling Kogaionidae (Mammalia, Multituberculata) in the Uppermost Cretaceous of Transylvania (Western Romania)

The latest Cretaceous kogaionid multituberculates from Transylvania (western Romania) were part of an endemic European clade of mammals that underwent an insular radiation at the end of the Cretaceous and then survived the end-Cretaceous mass extinction that extinguished many groups of contemporary therians. Transylvanian kogaionids lived on what was an island during the latest Cretaceous-"Hateg Island"-and their fossils are found in the uppermost Campanian to upper Maastrichtian deposits of the Hateg, Rusca Montana, and southwestern Transylvanian basins. This fossil record has improved dramatically over the past several decades, in part resulting from our decade-long joint Romanian-American-Scottish fieldwork, and comprises one of the most impressive and complete archives of Mesozoic mammals, including not only jaws and teeth but several incomplete skulls and partial skeletons. We here review the fossil record of kogaionids from Transylvania. We report four new occurrences from the Hateg Basin, update information on previously described ones, and use our database to reassess the chronostratigraphical and geographical distribution of kogaionids and their evolutionary patterns. Although it was previously suggested that large and small kogaionids had largely mutually exclusive spatial distributions, we recognize the cooccurrence of small and large taxa in various units, suggesting a sympatric distribution across their entire chronostratigraphic range. We also identify a novel pattern: small kogaionids appear somewhat earlier than their larger relatives in all well-sampled sedimentary successions, suggesting that kogaionid colonizations of Hateg Island and component regions took place at small body size and that body size increased only later through local evolution. We find correlations between body size, preservation style, and sedimentary context, which give insight into kogaionid paleobiology and diversity. Larger kogaionids are represented more often by partial skulls and occasionally skeletons compared with small kogaionids, which are usually represented only by isolated teeth, regardless of provenance. Larger kogaionids currently have a higher recognized local taxic diversity than their smaller relatives. We hypothesize that this may be in part a consequence of preservational bias related to body size, as more complete specimens may be more easily diagnosed as distinct taxa than those that are represented by more fragmentary and/or incomplete fossils. If true, the taxic diversity of smaller kogaionids may currently be underestimated. Finally, we identify correspondence between sedimentary facies and preservation style. Red-colored fine-grained rocks, suggestive of well-drained, oxidized floodplain paleoenvironments, yield more complete specimens than drab, greenish or grayish sediments deposited in more poorly drained parts of the floodplain. This pattern may suggest habitat preferences for better-drained floodplain environments and a semifossorial lifestyle for some taxa. As the kogaionid fossil record improves, we can further test the hypotheses and patterns outlined above. The pace of new kogaionid discoveries by our team and others indicates that a more complete picture of kogaionid distribution, paleobiology, and evolution will emerge in the coming years, contributing to a more profound understanding of this peculiar group of island-dwelling Mesozoic mammals.