A terrestrial fauna from the Scottish Lower Carboniferous

Abstract

Despite several important discoveries, extending over more than 120 years, our knowledge of early land vertebrates is still sparse. The earliest tetrapod remains are known from the Upper Devonian of East Greenland1–3 and Australia4–6, but the tetrapod fossil record does not become plentiful until Coal Measure times, in the Upper Carboniferous, some 50 Myr later. Finds in the Lower Carboniferous are very few indeed. Apart from two localities in West Virginia, USA7,8, and one in Nova Scotia, Canada9, all other Lower Carboniferous tetrapod sites are from the Visean of Fife and the Lothian Region, Scotland10,11. We report here the discovery of an assemblage of terrestrial animals from a new Lower Carboniferous locality in the Lothian Region. Specimens were collected from the East Kirkton Limestone in the Brigantian stage of the Scottish Visean, and include the first articulated amphibian skeleton to be found in the Lower Carboniferous of Europe in the twentieth century. This find is the earliest well-preserved amphibian skeleton ever discovered. The associated fauna is remarkable for the presence of myriapods, scorpions, the earliest known harvest-man and several other types of amphibian. The presence of such forms, together with the striking absence of fishes, suggests that the amphibians form an integral part of a terrestrial fauna; terrestrial amphibians are otherwise unknown before the Upper Carboniferous Coal Measures.