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The pterosaur is the earliest and largest powered flying vertebrate, even earlier and larger than the other extant archosaurian group, birds. However, evidence for this flying reptile, including the diversity of the small-sized pterosaur after the mid-Cretaceous, and their ecology, has remained elusive. Here we present numerous and dense pterosaur track assemblages from the Hwasun Seoyuri tracksite in the Upper Cretaceous Jangdong Formation of the Neungju Basin in Korea. The pterosaur track assemblage, assigned to Pteraichnus isp., consists of various sized, randomly oriented manus-dominated tracks with several pes claw marks. These features commonly indicate the semi-aquatic behavior and multi-age gregariousness of pterosaurs. The supposed trackmaker of pterosaur tracks would be the small-sized pterodactyloid that inhibited the Late Cretaceous Korean Peninsula, but that has not previously been reported. This ichnological evidence for the global distribution of small-sized pterosaurs could be interpreted to mean that the pterosaur fauna in the Late Cretaceous was more distributed and diverse than was previously known.

Here we describe a new pterosaur footprint assemblage from the Hwasun Seoyuri tracksite in the Upper Cretaceous Jangdong Formation of the Neungju Basin in Korea. The assemblage consists of many randomly oriented prints in remarkably high densities but represents a single ichnotaxon, Pteraichnus. Individuals exhibit a large but continuous size range, some of which, with a wingspan estimated at 0.5 m, are among the smallest pterosaurs yet reported from the Upper Cretaceous, adding to other recent finds which contradict the idea that large and giant forms entirely dominated this interval. Unusual features of the tracks, including relatively long, slender pedal digit impressions, do not match the pes of any known Cretaceous pterosaur, suggesting that the trackmakers are as yet unknown from the body fossil record. The Hwasun pterosaur footprints appear to record gregarious behavior at the exact location by individuals of different ages, hinting at the possibility that pterosaurs gathered in mixed-age groups.

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