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Knowledge of Cambrian animal anatomy is limited by preservational processes that result in compaction, size bias, and incompleteness. We documented pristine three-dimensional (3D) anatomy of trilobites fossilized through rapid ash burial from a pyroclastic flow entering a shallow marine environment. Cambrian ellipsocephaloid trilobites from Morocco are articulated and undistorted, revealing exquisite details of the appendages and digestive system. Previously unknown anatomy includes a soft-tissue labrum attached to the hypostome, a slit-like mouth, and distinctive cephalic feeding appendages. Our findings resolve controversy over whether the trilobite hypostome is the labrum or incorporates it and establish crown-group euarthropod homologies in trilobites. This occurrence of moldic fossils with 3D soft parts highlights volcanic ash deposits in marine settings as an underexplored source for exceptionally preserved organisms.

As the largest predators of the Cambrian seas, the anomalocaridids had an important impact in structuring the first complex marine animal communities, but many aspects of anomalocaridid morphology, diversity, ecology, and affinity remain unclear owing to a paucity of specimens. Here we describe the anomalocaridid Hurdia, based on several hundred specimens from the Burgess Shale in Canada. Hurdia possesses a general body architecture similar to those of Anomalocaris and Laggania, including the presence of exceptionally well-preserved gills, but differs from those anomalocaridids by possessing a prominent anterior carapace structure. These features amplify and clarify the diversity of known anomalocaridid morphology and provide insight into the origins of important arthropod features, such as the head shield and respiratory exites.

The arthropod Fuxianhuia from the Chengjiang fauna displays primitive aspects of cephalic segmentation and trunk limb morphology that indicate a basal position within Euarthropoda. The cephalon consists of an eye-bearing sclerite that articulates with a head shield bearing antennules and subchelate appendages. Eye stalks, antennules, and subchelate appendages are proto-, deuto-, and tritocerebral limbs and organs, respectovely. The anterior position of the eye-bearing sclerite parallels the embryonic origin of arthropod eye lobes. The head of Fuxianhuia includes the acron and one somite and is regarded as a protocephalon. The definitive head of arthropods may have fused separate eye-bearing and appendage-bearing sclerites.