A SPECIATIONAL HISTORY OF "LIVING FOSSILS": MOLECULAR EVOLUTIONARY PATTERNS IN HORSESHOE CRABS

Horseshoe crabs' exceptional morphological conservatism over the past 150 My has led to their reputation as "living fossils," but also has served to obscure phylogenetic relationships within the complex. Here we employ nucleotide sequences from two mitochondrial genes to assess molecular evolutionary rates and patterns among all extant horseshoe crab species. The American species Limulus polyphemus proved to be the sister taxon to a clade composed of the Asiatic species Tachypleus gigas, T. tridentatus, and Carcinoscorpius rotundicauda, whose relationships inter se were not resolved definitively. Both absolute and relative rate tests suggest a moderate slowdown in sequence evolution in horseshoe crabs. Nonetheless, dates of the lineage separations remain uncertain primarily because of reservations about molecular-clock calibrations resulting from large rate variances at examined loci across Arthropods and other animal lineages, as inferred in this and prior studies. Thus, ironically, separation dates as estimated by molecular evidence in general may remain most insecure in taxonomic groups for which such information is needed most-those lacking strong biogeographic or fossil benchmarks for internal-clock calibrations. In any event, the current results show that large numbers of molecular characters distinguish even these most morphologically conservative of organisms. Furthermore, comparisons against previously published mitochondrial sequence data in the morphologically dynamic hermit crab-king crab complex demonstrates that striking heterogeneity in levels of morphotypic differentiation can characterize Arthropod lineages at similar magnitudes of molecular divergence.