A pyritized Ordovician leanchoiliid arthropod.

Abstract

The "short-great-appendage" arthropods (Megacheira), such as Leanchoilia, have featured heavily in discussions of arthropod evolution, particularly related to the head and its appendages.^1,2,3,4 Megacheirans are subject to competing interpretations, either as a clade⁴ or a grade,⁵ in the stem group of Euarthropoda⁶ or, alternatively, Chelicerata.⁴ They are most diverse in Cambrian Burgess-Shale-type deposits, where the family Leanchoiliidae is represented by six genera,^{7,8,9,10,11,12} characterized by the presence of three distal flagella on the great appendage with a presumed sensory function. We describe the first post-Cambrian member of this family, Lomankus edgecombei gen. et sp. nov, from the Upper Ordovician (Katian) Beecher's Trilobite Bed site of New York State-the first post-Cambrian megacheiran with the exception of the Silurian and Devonian Enaliktidae. Micro-computed tomography (micro-CT) scanning reveals the morphology of the short great appendage with elongate flagella, four biramous cephalic limbs, 11 trunk segments with biramous limbs and dorsal tergites, and an elongate telson unique within Leanchoiliidae. The great appendage is also unique: the long endites that bear the flagella in other leanchoiliids are absent (or at least greatly reduced) and each flagellum appears to attach directly to an individual podomere, suggesting a sensory rather than a raptorial function. The remarkable preservation of a well-developed ventral plate (epistome-labrum complex) anterior of the mouth reinforces a deutocerebral origin^2,13 of the short great appendages. Lomankus edgecombei unveils the three-dimensional (3D) head morphology of leanchoiliids in unparalleled detail and demonstrates that these iconic fossil arthropods ranged into dysaerobic environments in the Ordovician, where Lomankus occupied a deposit-feeding niche.