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Most of these taxa (including trilobites) have traditionally been regarded as stem chelicerates under the Arachnomorpha hypothesis21,22, but more recent hypotheses regarding the organization of the arthropod head prompted their assignment to total group Mandibulata23. These and some subsequent studies considered deutocerebral antennae to be an autapomorphy of total group Mandibulata and the raptorial first pair of appendages of pycnogonids and euchelicerates to be a symplesiomorphy of Euarthropoda19,24. Under this scheme, the raptorial appendages of stem group euarthropods such as megacheirans (‘great appendage’ arthropods), fuxianhuiiids and bivalved stem group arthropods (for example, Odaraia, Canadaspis and Perspicaris) were considered homologous to the chelicerae of chelicerates, and any antenniform appendages anterior to this were considered segmentally homologous to the antennae of onychophorans, that is, protocerebral rather than deutocerebral.

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