From the mountains to the coast and back again: Ancient biogeography in a radiation of short-range endemic harvestmen from California.

The harvestmen genus Calicina is represented by 25 short-range endemic species occurring in the western Sierra Nevada, Transverse and Coast Ranges of California. Our principal aim was to reconstruct the temporal and spatial biogeographic history of this arachnid lineage. We inferred a time-calibrated species tree for 21 of 25 described Calicina species using multiple genes and multilocus coalescent-based methods. This species tree was used as a framework for algorithmic biogeographic and divergence time analyses, and a phylogenetic canonical correlation analysis (CCA) was used to examine the relationship between morphological evolution and environmental variables. Species tree and biogeographic analyses indicate that high-elevation Sierran taxa are early-diverging in Calicina, with subsequent biogeographic "criss-crossing" of lineages from the Sierra Nevada to the Coast Ranges, back to the Sierra Nevada, then back to Coast Ranges. In both the Sierra Nevada and Coast Ranges, distantly-related parapatric lineages essentially never occur in sympatry. CCA reveals that in both the Coast Ranges and the Sierra Nevada, distant phylogenetic relatives evolve convergent morphologies. Our evidence shows that Calicina is clearly dispersal-limited, with an ancient biogeographic history that provides unique insight into the complex geologic evolution of California since the mid-Paleogene.