Ceridwen I. Fraser

Long-distance dispersal: a framework for hypothesis testing

Tests of hypotheses about the biogeographical consequences of long-distance dispersal have long eluded biologists, largely because of the rarity and presumed unpredictability of such events. Here, we examine data for terrestrial (including littoral) organisms in the Pacific to show that knowledge of dispersal by wind, birds and oceanic drift or rafting, coupled with information about the natural environment and biology of the organisms, can be used to generate broad biogeographic predictions. We then examine the predictions in the context of the origin, frequency of arrival and location of establishment of dispersed organisms, as well as subsequent patterns of endemism and diversification on remote islands. The predicted patterns are being increasingly supported by phylogenetic data for both terrestrial and littoral organisms.

Kelp genes reveal effects of subantarctic sea ice during the Last Glacial Maximum

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.

Oceanic rafting by a coastal community

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.

Contemporary habitat discontinuity and historic glacial ice drive genetic divergence in Chilean kelp

BackgroundSouth Americas western coastline, extending in a near-straight line across some 35 latitudinal degrees, presents an elegant setting for assessing both contemporary and historic influences on cladogenesis in the marine environment. Southern bull-kelp (Durvillaea antarctica) has a broad distribution along much of the Chilean coast. This species represents an ideal model taxon for studies of coastal marine connectivity and of palaeoclimatic effects, as it grows only on exposed rocky coasts and is absent from beaches and ice-affected shores. We expected that, along the central Chilean coast, D. antarctica would show considerable phylogeographic structure as a consequence of the isolating effects of distance and habitat discontinuities. In contrast, we hypothesised that further south - throughout the region affected by the Patagonian Ice Sheet at the Last Glacial Maximum (LGM) - D. antarctica would show relatively little genetic structure, reflecting postglacial recolonisation.ResultsMitochondrial (COI) and chloroplast (rbcL) DNA analyses of D. antarctica from 24 Chilean localities (164 individuals) revealed two deeply divergent (4.5 - 6.1% for COI, 1.4% for rbcL) clades from the centre and south of the country, with contrasting levels and patterns of genetic structure. Among populations from central Chile (32° - 44°S), substantial phylogeographic structure was evident across small spatial scales, and a significant isolation-by-distance effect was observed. Genetic disjunctions in this region appear to correspond to the presence of long beaches. In contrast to the genetic structure found among central Chilean populations, samples from the southern Chilean Patagonian region (49° - 56°S) were genetically homogeneous and identical to a haplotype recently found throughout the subantarctic region.ConclusionsSouthern (Patagonian) Chile has been recolonised by D. antarctica relatively recently, probably since the LGM. The inferred trans-oceanic ancestry of these Patagonian populations supports the notion that D. antarctica is capable of long-distance dispersal via rafting. In contrast, further north in central Chile, the correspondence of genetic disjunctions in D. antarctica with long beaches indicates that habitat discontinuity drives genetic isolation among established kelp populations. We conclude that rafting facilitates colonisation of unoccupied shores, but has limited potential to enhance gene-flow among established populations. Broadly, this study demonstrates that some taxa may be considered to have either high or low dispersal potential across different temporal and geographic scales.

Poleward bound: biological impacts of Southern Hemisphere glaciation

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.

The changing form of Antarctic biodiversity

Antarctic biodiversity is much more extensive, ecologically diverse and biogeographically structured than previously thought. Understanding of how this diversity is distributed in marine and terrestrial systems, the mechanisms underlying its spatial variation, and the significance of the microbiota is growing rapidly. Broadly recognizable drivers of diversity variation include energy availability and historical refugia. The impacts of local human activities and global environmental change nonetheless pose challenges to the current and future understanding of Antarctic biodiversity. Life in the Antarctic and the Southern Ocean is surprisingly rich, and as much at risk from environmental change as it is elsewhere.

Geothermal activity helps life survive glacial cycles

Significance The evolution and maintenance of diversity through cycles of past climate change have hinged largely on the availability of refugia. Geothermal refugia may have been particularly important for survival through past glaciations. Our spatial modeling of Antarctic biodiversity indicates that some terrestrial groups likely survived throughout intense glacial cycles on ice-free land or in sub-ice caves associated with areas of geothermal activity, from which recolonization of the rest of the continent took place. These results provide unexpected insights into the responses of various species to past climate change and the importance of geothermal regions in promoting biodiversity. Furthermore, they indicate the likely locations of biodiversity “hotspots” in Antarctica, suggesting a critical focus for future conservation efforts. Climate change has played a critical role in the evolution and structure of Earth’s biodiversity. Geothermal activity, which can maintain ice-free terrain in glaciated regions, provides a tantalizing solution to the question of how diverse life can survive glaciations. No comprehensive assessment of this “geothermal glacial refugia” hypothesis has yet been undertaken, but Antarctica provides a unique setting for doing so. The continent has experienced repeated glaciations that most models indicate blanketed the continent in ice, yet many Antarctic species appear to have evolved in almost total isolation for millions of years, and hence must have persisted in situ throughout. How could terrestrial species have survived extreme glaciation events on the continent? Under a hypothesis of geothermal glacial refugia and subsequent recolonization of nongeothermal regions, we would expect to find greater contemporary diversity close to geothermal sites than in nongeothermal regions, and significant nestedness by distance of this diversity. We used spatial modeling approaches and the most comprehensive, validated terrestrial biodiversity dataset yet created for Antarctica to assess spatial patterns of diversity on the continent. Models clearly support our hypothesis, indicating that geothermally active regions have played a key role in structuring biodiversity patterns in Antarctica. These results provide critical insights into the evolutionary importance of geothermal refugia and the history of Antarctic species.

Asymmetric dispersal of southern bull-kelp (Durvillaea antarctica) adults in coastal New Zealand: testing an oceanographic hypothesis

Coastal populations are often connected by unidirectional current systems, but the biological effects of such asymmetric oceanographic connectivity remain relatively unstudied. We used mtDNA analysis to determine the phylogeographic origins of beach‐cast bull‐kelp (Durvillaea antarctica) adults in the Canterbury Bight, a 180 km coastal region devoid of rocky‐reef habitat in southern New Zealand. A multi‐year, quantitative analysis supports the oceanographically derived hypothesis of asymmetric dispersal mediated by the north‐flowing Southland Current. Specifically, 92% of beach‐cast specimens examined had originated south of the Bight, many drifting north for hundreds of kilometres, and some traversing at least 500 km of ocean from subantarctic sources. In contrast, only 8% of specimens had dispersed south against the prevailing current, and these counter‐current dispersers likely travelled relatively small distances (tens of kilometres). These data show that oceanographic connectivity models can provide robust estimates of passive biological dispersal, even for highly buoyant taxa. The results also indicate that there are no oceanographic barriers to kelp dispersal across the Canterbury Bight, indicating that other ecological factors explain the phylogeographic disjunction across this kelp‐free zone. The large number of long‐distance dispersal events detected suggests drifting macroalgae have potential to facilitate ongoing connectivity between otherwise isolated benthic populations.

Glacial oceanographic contrasts explain phylogeography of Australian bull kelp

The evolutionary effects of Southern Hemisphere Pleistocene oceanographic conditions — marked by fluctuations in sea levels and water temperatures, and redirected currents — are poorly understood. The southeastern tip of Australia presents an intriguing model system for studying the biological impacts of palaeoceanography. In particular, contrasting oceanographic conditions that existed on eastern vs. western sides of the Bassian Isthmus during Pleistocene glacial periods allow for natural comparisons between putative refugial vs. re‐invading populations. Whereas many western Tasmanian marine taxa were likely eliminated by cold subantarctic water during the last glacial period, eastern Tasmanian populations would have persisted in relatively warm temperatures mediated by the ongoing influence of the East Australian Current (EAC). Here we test for the effects of contrasting palaeoceanographic conditions on endemic bull kelp, Durvillaea potatorum, using DNA sequence analysis (COI; rbcL) of more than 100 individuals from 14 localities in southeastern Australia. Phylogenetic reconstructions reveal a deep (maximum divergence 4.7%) genetic split within D. potatorum, corresponding to the ‘eastern’ and ‘western’ geographical regions delimited by the Bassian Isthmus, a vicariant barrier during low Pleistocene sea levels. Concordant with the western regions cold glacial conditions, samples from western Tasmania and western Victoria are genetically monomorphic, suggesting postglacial expansion from a mainland refugium. Eastern samples, in contrast, comprise distinct regional haplogroups, suggesting the species persisted in eastern Tasmania throughout recent glacial periods. The deep east–west divergence seems consistent with earlier reports of morphological differences between ‘western’ and ‘eastern’D. potatorum, and it seems likely that these forms represent reproductively isolated species.

GENETIC AND MORPHOLOGICAL ANALYSES OF THE SOUTHERN BULL KELP DURVILLAEA ANTARCTICA (PHAEOPHYCEAE: DURVILLAEALES) IN NEW ZEALAND REVEAL CRYPTIC SPECIES

Many macroalgae exhibit considerable intraspecific morphological variation, but whether such variation reflects phenotypic plasticity or underlying genetic differences is often poorly understood. We quantified both morphological and genetic variation of 96 plants from seven field sites across eastern South Island, New Zealand, to assess genetic differences between morphotypes of the southern bull kelp Durvillaea antarctica (Cham.) Har. Consistent DNA sequence differentiation across mitochondrial, plastid, and nuclear loci was correlated with two broadly sympatric morphotypes: “cape” and “thonged.” These ecologically, morphologically, and genetically distinct bull‐kelp lineages were previously considered to be environmentally determined phenotypes with no underlying genetic basis. Interestingly, the sheltered “cape” lineage appears essentially genetically uniform across its South Island range, whereas the exposed “thonged” lineage exhibits marked phylogeographic structure across its range. Results suggest that D. antarctica in New Zealand comprises two reproductively isolated species.