Appendages of an early Cambrian metadoxidid trilobite from Yunnan, SW China support mandibulate affinities of trilobites and artiopods

Appendage anatomy contributes crucial data for understanding the evolution and ecology of Euarthropoda. The Palaeozoic trilobites show a great diversity of exoskeletons in the fossil record. However, soft parts, especially appendages, have only been discovered from a few trilobite species. Here we report extraordinarily preserved appendages in the trilobite species Hongshiyanaspis yiliangensis Zhang & Lin in Zhang et al. 1980 (Redlichiida, Metadoxididae) from a single mudstone layer of the Xiazhuang fossil assemblage within the Hongjingshao Formation (Cambrian Series 2, Stage 3) near Kunming, Yunnan, SW China. The appendages exhibit the common architecture revealed by other trilobites and artiopods by consisting of a pair of uniramous antennae followed by a series of paired homonomous biramous limbs. The antennae in holaspid individuals comprise up to 27 spinous podomeres and their ontogeny occurs by lengthening of the podomeres. The post-antennal biramous limbs are similar to those in other polymerid trilobites and artiopods by having a single-segmented protopodite and an endopodite comprising seven segments, but possess a unique wide tripartite exopodite with long setae. Sophisticated appendage anatomy, including the body-limb junction, fine setae, putative muscle bundles and duct-type tissues, are also revealed. Appendages of trilobites, artiopods and other upper stem-group euarthropods are compared and summarized. The H. yiliangensis appendages highlight the high morphological disparity of exopodites and the conservativeness of endopodites in trilobites and artiopods. This morphological pattern, together with similar body patterning seen in crustaceans but not in chelicerates, supports the mandibulate affinities of trilobites and at least some artiopods.