ABSTRACTThe Arachnomorpha or Arachnata concepts have resolved Trilobita as most closely relatedto Chelicerata amongst extant Arthropoda. An alternative position of trilobites in the stemlineage of Mandibulata is suggested by their pattern of head tagmosis. The antennae of trilobites and Mandibulata are considered non-homologous with the antennae of Onycho- phora and stem lineage Euarthropoda: they represent ‘secondary’ and ‘primary antennae’,respectively. In extant taxa, ‘secondary antennae’ are deutocerebral, post-ocular, and areconnected to deutocerebral olfactory neuropils, whereas ‘primary antennae’ are pre-ocular and connected to protocerebral olfactory neuropils. In fossils, an insertion at the antero-lateral margin of the hypostome rather than more anteriorly on the head allows ‘secondaryantennae’ to be identified. A deutocerebral mouthpart, of which the onychophoran jaw andthe chelicera are examples, is regarded as plesiomorphic for Arthropoda. A loss of ‘primaryantennae’ and modification of the deutocerebral mouthpart into a sensory antenna definesthe Mandibulata. Trilobites share a ‘secondary antenna’ and a clearly-delimited head tagmawith mandibulates. Given the extensive homoplasy forced by the Arachnata concept (rever-sals in pycnogonids and arachnids), a trilobite/mandibulate alliance may be better sup- ported.
Dedicated to Fred Schram on the occasion of his retirement. Our article challenges canonical viewsabout arthropods, and we were forced to question ideas that we have long considered the best expla-nation of facts. In doing so, we venture into a territory from which Fred Schram has never shied away. Fred’s synthesis of data from living and fossil arthropods, his efforts to integrate classical morphological and evo-devo perspectives, and his willingness to explore dangerous ideas haveinspired our reappraisal of the trilobite problem.
INTRODUCTIONThe last decade has seen dramatic changes of our views on arthropod development, mor- phology, palaeontology, phylogeny, and evolution (see, for example, the books edited byFortey & Thomas 1997; Edgecombe 1998; Deuve 2001; Scholtz 2004). The comparativemolecular approach to embryology, cell lineage studies, new microscopic techniques with ahigh morphological resolution, and phylogenetic analyses based on molecular and refined
Heads, Hox and the phylogenetic position of trilobites
& GREGORY D. EDGECOMBE
Institut für Biologie/Vergleichende Zoologie, Humboldt-Universität zu Berlin, Berlin, Germany
Australian Museum, Sydney, Australia