  Horseshoe crabs are marine arthropods with a fossil record extending back
approximately 450 million years. They exhibit remarkable morphological
stability over their long evolutionary history, retaining a number of ancestral
arthropod traits, and are often cited as examples of "living fossils." As
arthropods, they belong to the Ecdysozoa}, an ancient super-phylum whose
sequenced genomes (including insects and nematodes) have thus far shown more
divergence from the ancestral pattern of eumetazoan genome organization than
cnidarians, deuterostomes, and lophotrochozoans. However, much of ecdysozoan
diversity remains unrepresented in comparative genomic analyses. Here we use a
new strategy of combined de novo assembly and genetic mapping to examine the
chromosome-scale genome organization of the Atlantic horseshoe crab Limulus
polyphemus. We constructed a genetic linkage map of this 2.7 Gbp genome by
sequencing the nuclear DNA of 34 wild-collected, full-sibling embryos and their
parents at a mean redundancy of 1.1x per sample. The map includes 84,307
sequence markers and 5,775 candidate conserved protein coding genes. Comparison
to other metazoan genomes shows that the L. polyphemus genome preserves
ancestral bilaterian linkage groups, and that a common ancestor of modern
horseshoe crabs underwent one or more ancient whole genome duplications (WGDs)
~ 300 MYA, followed by extensive chromosome fusion.
