New evidence for five cephalic appendages in trilobites and implications for segmentation of the trilobite head

Abstract

Head segments play a critical role in revealing the relationships within major arthropod groups but the accurate determination of their number and relationship is challenging because of the decoupling of dorsal tergites and ventral appendages. In recent decades, trilobites have been considered to have four pairs of cephalic appendages and, commonly, five dorsal segments, which has served as the basis for understanding euarthropod cephalic evolution. Based on multiple analytical techniques applied to well-preserved soft-bodied specimens of the Upper Ordovician olenid Triarthrus eatoni and middle Cambrian corynexochid Olenoides serratus, we argue that an additional pair of cephalic biramous appendages occurred just behind the antennae, indicating that trilobites had five pairs of cephalic appendages and six dorsally expressed segments. We propose that the mismatch between the four dorsal furrows and five ventral appendages in Olenoides may be reconciled by considering the morphology of the hypostome that marks the position of the antenna and the bifurcation of the first glabellar furrow that has two muscle attachment sites fused. The fusion of clustered attachment sites of the cephalic biramous appendages may have aided feeding ability by gathering the limb bases close to the mouth.