Cladistic relationships of Trilobita, Naraoiidae (five ingroup taxa), Helmetiida (five ingroup taxa), Xandarellida, and the Cambrian arachnates Retifacies, Sinoburius, Emeraldella, and Sidneyia are investigated based on 29 characters. Documentation of appendage morphology and other ventral structures in Saperion from the Chengjiang fauna permits an appraisal of helmetiid relationships. A monophyletic Trilobita [=“Olenellida” (Emuellida + Eutrilobita)] is defined by numerous synapomorphies, including exoskeletal calcification and dorsal eyes with calcified lenses and circumocular sutures. Helmetiida is a robust clade, resolved as (Helmetiidae (Tegopeltidae (Saperiidae + Skioldiidae))). Naraoiid monophyly is well-supported, but neither a naraoiid-trilobite nor a naraoiid-Retifacies clade are parsimonious, the latter grouping (“Nectopleura”) being explicitly paraphyletic. A sister group relationship between Xandarellida and Sinoburius is endorsed, although character support is novel compared to previous groupings of these taxa. The fourth postantennal limb pair in trilobites, naraoiids, and apparently helmetiids is based beneath the cephalothoracic articulation. Reweighted characters favor Trilobita and Helmetiida as closest relatives, with Petalopleura and then Naraoiidae as sister groups.

Chelicerates are a diverse group of arthropods, represented by such forms as predatory spiders and scorpions, parasitic ticks, humic detritivores, and marine sea spiders (pycnogonids) and horseshoe crabs. Conflicting phylogenetic relationships have been proposed for chelicerates based on both morphological and molecular data, the latter usually not recovering arachnids as a clade and instead finding horseshoe crabs nested inside terrestrial Arachnida. Here, using genomic-scale datasets and analyses optimised for countering systematic error, we find strong support for monophyletic Acari (ticks and mites), which when considered as a single group represent the most biodiverse chelicerate lineage. In addition, our analysis recovers marine forms (sea spiders and horseshoe crabs) as the successive sister groups of a monophyletic lineage of terrestrial arachnids, suggesting a single colonisation of land within Chelicerata and the absence of wholly secondarily marine arachnid orders.

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Highlights d We describe a new early Cambrian fossil with a polypoid body plan from Chengjiang d It is related to the supposed cnidarian Xianguangia and the iconic Dinomischus d The fossils possess very large ciliary structures otherwise seen only in ctenophores d We show that these fossils form a grade of stem-group ctenophores