Phylogeography of sugar kelp: Northern ice-age refugia in the Gulf of Alaska.

Many Northeast (NE) Pacific fishes and invertebrates survived Pleistocene glaciations in northern refugia, but the extent that kelps survived in northern areas is uncertain. Here, we test the hypothesis that populations of sugar kelp () persisted in the Gulf of Alaska during ice-age maxima when the western margin of the Cordilleran ice sheet covered coastal areas around the NE Pacific Ocean. We estimated genetic diversities within and phylogeographical relationships among 14 populations along 2,800 km in the NE Pacific and Bering Sea with partial sequences of mitochondrial DNA 5'-cytochrome oxidase subunit I (COI,  = 624,  = 543), chloroplast DNA ribulose-1,5-bisphosphate carboxylase large subunit-3' (L,  = 735,  = 514), and 11 microsatellite loci. Concatenated sequences of L and showed moderate levels of within-population genetic diversity (mean  = 0.200) but substantial differences among populations (Φ = 0.834,  < .0001). Microsatellites showed moderate levels of heterozygosity within populations (mean  = 0.391). Kelps in the same organellar lineage tended to cluster together, regardless of geographic origins, as indicated in a principal coordinate analysis (PCoA) of microsatellite genotypes. The PCoA also showed evidence of nuclear hybridizations between co-occurring organellar lineages. Individual admixture plots with population clusters of  = 2, 6, and 9 showed increasing complexity with considerable historical admixture between some clusters. A time-calibrated phylogeny placed divergences between L- lineages at 1.4 million years at most. The time frames of mutation in the L- lineages and microsatellite population clusters differed among locations. The existence of ancient lineages in the Gulf of Alaska, moderate levels of genetic diversity, and the absence of departures from neutrality are consistent with northern refugia during multiple Croll-Milankovitch climate cycles in the Pleistocene Epoch.