Aim
Species or higher taxa that are obviously dispersal‐limited, but which occupy large geographical distributions, represent a biogeographical paradox. Dispersal must have happened, likely under special and infrequent environmental conditions, but details have been lost to history. The overarching goal of our research is to understand the details of a ‘common vicariance, rare dispersal’ biogeographical history in a widespread but habitat‐specialized harvestman species (Sabacon cavicolens) with a southern Appalachian centre of distribution.
Location
Eastern North America, southern Appalachians.
Methods
We assessed cryptic speciation using mitochondrial and nuclear gene DNA sequence data, testing alternative delimitation hypotheses using multispecies coalescent analyses. We also tested whether riverine barriers are associated with mitochondrial genealogical structuring, focusing on multiple rivers in the southern Blue Ridge physiographical province. Finally, we conducted population genetic analyses to assess female‐based range expansion out of the southern Blue Ridge.
Results
Genetic analyses suggest a large number of species‐level lineages within S. cavicolens, although we prefer a more conservative three‐species hypothesis. These putative species are geographically cohesive and allopatric (Ozarks, Cumberland Plateau, southern Blue Ridge), with the Blue Ridge species including multiple divergent mitochondrial haplogroups. Several genealogical breaks in the Blue Ridge species coincide with riverine barriers, separating mostly allopatric mitochondrial lineages. Contrasting with evidence for constrained gene flow and vicariance, two Blue Ridge haplogroups reveal extensive range expansion both northwards and westwards, resulting in the widespread distribution of closely related haplotypes, and occasional sympatry of dispersive haplotypes.
Main Conclusions
Hidden beneath the apparently widespread distribution of a single species is a history of old vicariance separating geographically disjunct cryptic species. How these lineages came to occupy such disparate geographies is illustrated by dynamics within the Blue Ridge species, where both in situ vicariance and long‐distance dispersal have shaped a ‘common vicariance, rare dispersal’ biogeographical history.
Our focus was to design harvestmen-specific PCR primers to target both introns and exons of the nuclear protein-coding gene Elongation Factor -1 alpha (EF-la). We tested this primer set on ten genera representing all primary iineages of Opiliones, with sets of close phylogenetic relatives (i.e., sets of several congeners) included to specifically assess utility at shallow phylogenetic levels. Our research also included the collection of parallel mitochondrial protein-coding DNA sequence datasets for the congeneric sets to compare relative rates of evolution and gene tree congruence for EF-la versus mitochondrial data. The harvestmen primers resulted in successful amplification for nine of ten tested genera. Exon sequences for these nine genera appear orthologous to previously-reported EF-la Opiliones sequences, which were generated using RT-PCR methods. Newly-generated exon sequences are interrupted by three separate spliceosomal introns; two introns are restricted to one or two genera, but a third intron is conserved in position across all surveyed genera. Phylogenetic analyses of EF-la nucleotide data for congeneric sets result in gene trees that are generally congruent with mitochondrial gene trees, with EF-la phylogenetic signal coming from both intron and exon sites, and resolving apparently recent divergences (e.g., as recent as one million years ago). Overall, the combination of gene orthology, conserved intron position, and gene tree congruence at shallow levels suggest that this gene region will prove generally useful for both phylogeographic and species-level phylogenetic analyses in Opiliones, complementing already-documented utility at higher taxonomic levels.
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