Cambrian origins and affinities of an enigmatic fossil group of arthropods.

Euthycarcinoids are one of the most enigmatic arthropod groups, having been assigned to nearly all major clades of Arthropoda. Recent work has endorsed closest relationships with crustaceans or a myriapod-hexapod assemblage, a basal position in the Euarthropoda, or a placement in the Hexapoda or hexapod stem group. Euthycarcinoids are known from 13 species ranging in age from Late Ordovician or Early Silurian to Middle Triassic, all in freshwater or brackish water environments. Here we describe a euthycarcinoid from marine strata in Argentina dating from the latest Cambrian period, extending the group's record back as much as 50 million years. Despite its antiquity and marine occurrence, the Cambrian species demonstrates that morphological details were conserved in the transition to fresh water. Trackways in the same unit as the euthycarcinoid strengthen arguments that similar traces of subaerial origin from Cambro-Ordovician rocks were made by euthycarcinoids. Large mandibles in euthycarcinoids are confirmed by the Cambrian species. A morphology-based phylogeny resolves euthycarcinoids as stem-group Mandibulata, sister to the Myriapoda and Crustacea plus Hexapoda.

Arthropod phylogeny based on eight molecular loci and morphology.

The interrelationships of major clades within the Arthropoda remain one of the most contentious issues in systematics, which has traditionally been the domain of morphologists. A growing body of DNA sequences and other types of molecular data has revitalized study of arthropod phylogeny and has inspired new considerations of character evolution. Novel hypotheses such as a crustacean-hexapod affinity were based on analyses of single or few genes and limited taxon sampling, but have received recent support from mitochondrial gene order, and eye and brain ultrastructure and neurogenesis. Here we assess relationships within Arthropoda based on a synthesis of all well sampled molecular loci together with a comprehensive data set of morphological, developmental, ultrastructural and gene-order characters. The molecular data include sequences of three nuclear ribosomal genes, three nuclear protein-coding genes, and two mitochondrial genes (one protein coding, one ribosomal). We devised new optimization procedures and constructed a parallel computer cluster with 256 central processing units to analyse molecular data on a scale not previously possible. The optimal 'total evidence' cladogram supports the crustacean-hexapod clade, recognizes pycnogonids as sister to other euarthropods, and indicates monophyly of Myriapoda and Mandibulata.

Head segmentation in early cambrian fuxianhuia: implications for arthropod evolution.

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.