The interrelationships of major clades within the Arthropoda remain one of
the most contentious issues in systematics, which has traditionally been th
e domain of morphologists(1,2). A growing body of DNA sequences and other t
ypes of molecular data has revitalized study of arthropod phylogeny(3-7) an
d has inspired new considerations of character evolution(8,9). Novel hypoth
eses such as a crustacean-hexapod affinity(4,10-12) were based on analyses
of single or few genes and limited taxon sampling, but have received recent
support from mitochondrial gene order(13), and eye and brain ultrastructur
e and neurogenesis(14,15). Here we assess relationships within Arthropoda b
ased on a synthesis of all well sampled molecular loci together with a comp
rehensive data set of morphological, developmental, ultrastructural and gen
e-order characters. The molecular data include sequences of three nuclear r
ibosomal genes, three nuclear protein-coding genes, and two mitochondrial g
enes (one protein coding, one ribosomal). We devised new optimization proce
dures(16,17) and constructed a parallel computer cluster with 256 central p
rocessing units(18) to analyse molecular data on a scale not previously pos
sible. The optimal 'total evidence' cladogram supports the crustacean-hexap
od clade, recognizes pycnogonids as sister to other euarthropods, and indic
ates monophyly of Myriapoda and Mandibulata.