Article: A larval sea spider (Arthropoda: Pycnogonida) from the Upper Cambrian 'Orsten' of Sweden, and the phylogenetic position of pycnogonids

Among a set of small, secondarily phosphatised larval arthropods from the Upper Cambrian 'Orsten' of Sweden, described by Muller and Walossek in 1986, one form bears a remarkable resemblance to the hatching protonymph larva of extant Pantopoda. This 'larva D' shares with protonymphs their gross body form, the anteroventral mouth on a slightly off-set forehead region, the cheliceral morphology, two homeomorphic pairs of post-cheliceral limbs, and further detailed similarities. It is described herein as Cambropycnogon klausmuelleri gen. et sp. nov. and is proposed as the oldest unequivocal record of both Pycnogonida and Chelicerata. Plesiomorphic features such as a pair of rudimentary pre-cheliceral limbs and the gnathobasic basipods of the two post-cheliceral limbs distinguish it from all known larvae of extant Pantopoda and lead us to propose a phylogeny of the Pycnogonida of the form (Cambropycnogon klausmuelleri+ (Palaeoisopus+ (Palaeopantopus+ Pantopoda))). The fossil may help to resolve the long debate about the relationships of Pycnogonida to other Arthropoda and supports a (Pycnogonida + Euchelicerata) relationship within the Chelicerata. The pre-cheliceral limbs in this fossil support traditional morphological studies in which the chelicera represent the second (a2) head appendage, corresponding to the crustacean 'second antennae', and contradict recent data based on homeobox genes implying that the chelicerae are the first (a1) head appendages homologous with crustacean first antennae.