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The end of the Last Glacial Maximum (LGM) dramatically reshaped temperate ecosystems, with many species moving poleward as temperatures rose and ice receded. Whereas reinvading terrestrial taxa tracked melting glaciers, marine biota recolonized ocean habitats freed by retreating sea ice. The extent of sea ice in the Southern Hemisphere during the LGM has, however, yet to be fully resolved, with most palaeogeographic studies suggesting only minimal or patchy ice cover in subantarctic waters. Here, through population genetic analyses of the widespread Southern Bull Kelp (Durvillaea antarctica), we present evidence for persistent ice scour affecting subantarctic islands during the LGM. Using mitochondrial and chloroplast genetic markers (COI; rbcL) to genetically characterize some 300 kelp samples from 45 Southern Ocean localities, we reveal a remarkable pattern of recent recolonization in the subantarctic. Specifically, in contrast to the marked phylogeographic structure observed across coastal New Zealand and Chile (10-to 100-km scales), subantarctic samples show striking genetic homogeneity over vast distances (10,000-km scales), with a single widespread haplotype observed for each marker. From these results, we suggest that sea ice expanded further and ice scour during the LGM impacted shallow-water subantarctic marine ecosystems more extensively than previously suggested.Durvillaea antarctica ͉ genetic ͉ phylogeography ͉ raft

Antarctica has long been considered biologically isolated 1 . Global warming will make parts of Antarctica more habitable for invasive taxa, yet presumed barriers to dispersal-especially the Southern Ocean's strong, circumpolar winds, ocean currents and fronts-have been thought to protect the region from non-anthropogenic colonizations from the north 1,2 . We combine molecular and oceanographic tools to directly test for biological dispersal across the Southern Ocean. Genomic analyses reveal that rafting keystone kelps recently travelled > 20,000 km and crossed several ocean-front 'barriers' to reach Antarctica from mid-latitude source populations. High-resolution ocean circulation models, incorporating both mesoscale eddies and wave-driven Stokes drift, indicate that such Antarctic incursions are remarkably frequent and rapid. Our results demonstrate that storm-forced surface waves and ocean eddies can dramatically enhance oceanographic connectivity for drift particles in surface layers, and show that Antarctica is not biologically isolated. We infer that Antarctica's long-standing ecological differences have been the result of environmental extremes that have precluded the establishment of temperate-adapted taxa, but that such taxa nonetheless frequently disperse to the region. Global warming thus has the potential to allow the establishment of diverse new species-including keystone kelps that would drastically alter ecosystem dynamics-even without anthropogenic introductions.Antarctica broke from its last Gondwanan connection (South America) over 40 million years ago 3 , and has long been considered biologically isolated from the rest of the world 1 . Antarctica's terrestrial and marine ecosystems have high levels of endemism 4 , and the Southern Ocean's strong, circumpolar winds and ocean currents have been inferred to preclude passive southward dispersal of organisms into the Antarctic 1,2 . The onset of deep-reaching flow of the Antarctic Circumpolar Current (ACC) created a barrier to poleward heat transport across the Southern Ocean 5 , and the strong, circumpolar thermal fronts have also been inferred to represent an impenetrable physical barrier for most organisms. Indeed, there has been almost no evidence of biological movement into the Antarctic from lower latitudes since the Last Glacial Maximum

Postglacial recolonisation patterns are well documented for the Northern Hemisphere biota, but comparable processes in the Southern Hemisphere have only recently been examined. In the largely terrestrial Northern Hemisphere, recession of ice after the Last Glacial Maximum (LGM) allowed various taxa, including slow-moving terrestrial species, to migrate poleward. By contrast, the Southern Hemisphere polar region is completely ringed by ocean, and recolonisation of Antarctica and the sub-Antarctic islands has thus presented considerable challenges. Although a few highly dispersive marine species have been able to recolonise postglacially, most surviving high-latitude taxa appear to have persisted throughout glacial maxima in local refugia. These contrasting patterns highlight the importance of habitat continuity in facilitating biological range shifts in response to climate change.

Knowledge of DNA evolution is central to our understanding of biological history, but how fast does DNA change? Previously, pedigree and ancient DNA studies--focusing on evolution in the short term--have yielded molecular rate estimates substantially faster than those based on deeper phylogenies. It has recently been suggested that short-term, elevated molecular rates decay exponentially over 1-2 Myr to long-term, phylogenetic rates, termed "time dependency of molecular rates." This transition has potential to confound molecular inferences of demographic parameters and dating of many important evolutionary events. Here, we employ a novel approach--geologically dated changes in river drainages and isolation of fish populations--to document rates of mitochondrial DNA change over a range of temporal scales. This method utilizes precise spatiotemporal disruptions of linear freshwater systems and hence avoids many of the limitations associated with typical DNA calibration methods involving fossil data or island formation. Studies of freshwater-limited fishes across the South Island of New Zealand have revealed that genetic relationships reflect past, rather than present, drainage connections. Here, we use this link between drainage geology and genetics to calibrate rates of molecular evolution across nine events ranging in age from 0.007 Myr (Holocene) to 5.0 Myr (Pliocene). Molecular rates of change in galaxiid fishes from calibration points younger than 200 kyr were faster than those based on older calibration points. This study provides conclusive evidence of time dependency in molecular rates as it is based on a robust calibration system that was applied to closely related taxa, and analyzed using a consistent and rigorous methodology. The time dependency observed here appears short-lived relative to previous suggestions (1-2 Myr), which has bearing on the accuracy of molecular inferences drawn from processes operating within the Quaternary and mechanisms invoked to explain the decay of rates with time.

Oceanic rafting is thought to play a fundamental role in assembling the biological communities of isolated coastal ecosystems. Direct observations of this key ecological and evolutionary process are, however, critically lacking. The importance of macroalgal rafting as a dispersal mechanism has remained uncertain, largely owing to lack of knowledge about the capacity of fauna to survive long voyages at sea and successfully make landfall and establish. Here, we directly document the rafting of a diverse assemblage of intertidal organisms across several hundred kilometres of open ocean, from the subantarctic to mainland New Zealand. Multispecies analyses using phylogeographic and ecological data indicate that 10 epifaunal invertebrate species rafted on six large bull kelp specimens for several weeks from the subantarctic Auckland and/or Snares Islands to the Otago coast of New Zealand, a minimum distance of some 400 -600 km. These genetic data are the first to demonstrate that passive rafting can enable simultaneous trans-oceanic transport and landfall of numerous coastal taxa.

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