Diversity and distribution patterns in high southern latitude sponges

Sponges play a key role in Antarctic marine benthic community structure and dynamics and are often a dominant component of many Southern Ocean benthic communities. Understanding the drivers of sponge distribution in Antarctica enables us to understand many of general benthic biodiversity patterns in the region. The sponges of the Antarctic and neighbouring oceanographic regions were assessed for species richness and biogeographic patterns using over 8,800 distribution records. Species-rich regions include the Antarctic Peninsula, South Shetland Islands, South Georgia, Eastern Weddell Sea, Kerguelen Plateau, Falkland Islands and north New Zealand. Sampling intensity varied greatly within the study area, with sampling hotspots found at the Antarctic Peninsula, South Georgia, north New Zealand and Tierra del Fuego, with limited sampling in the Bellingshausen and Amundsen seas in the Southern Ocean. In contrast to previous studies we found that eurybathy and circumpolar distributions are important but not dominant characteristics in Antarctic sponges. Overall Antarctic sponge species endemism is ～43%, with a higher level for the class Hexactinellida (68%). Endemism levels are lower than previous estimates, but still indicate the importance of the Polar Front in isolating the Southern Ocean fauna. Nineteen distinct sponge distribution patterns were found, ranging from regional endemics to cosmopolitan species. A single, distinct Antarctic demosponge fauna is found to encompass all areas within the Polar Front, and the sub-Antarctic regions of the Kerguelen Plateau and Macquarie Island. Biogeographical analyses indicate stronger faunal links between Antarctica and South America, with little evidence of links between Antarctica and South Africa, Southern Australia or New Zealand. We conclude that the biogeographic and species distribution patterns observed are largely driven by the Antarctic Circumpolar Current and the timing of past continent connectivity.     

How isolated is Antarctica?

The traditional view of Antarctica and the surrounding Southern Ocean as an isolated system is now being challenged by the recent discovery at the Antarctic Peninsula of adult spider crabs Hyas areneus from the North Atlantic and of larvae of subpolar marine invertebrates. These observations question whether the well described biogeographical similarities between the benthic fauna of the Antarctic Peninsula and the Magellan region of South America result from history (the two regions were once contiguous), or from a previously unrecognized low level of faunal exchange. Such exchange might be influenced by regional climate change, and also exacerbated by changes in human impact.     

Biodiversity and biogeographic affiliation of Bryozoa from King George Island (Antarctica)

King George Island (KGI), which is located between the Antarctic and South American continents, may play a crucial role in the exchange of Bryozoa amongst the various Antarctic sectors and across the Polar Front. Knowledge regarding the biological diversity of this area could help us understand the evolution of the Antarctic ecosystem and its connectivity to the South American continent as well as the colonization ability of particular species. Here, we investigate the patterns of diversity and biogeographic affiliation of the cheilostome Bryozoa from KGI and the surrounding areas. Of 114 identified taxa from a depth range of 6–492 m, 26 species were reported for the first time in KGI. The most speciose genera were Camptoplites, Osthimosia, Smittina, and Cellarinella. Species richness at KGI consisted of 70% of the total bryozoans at the South Shetland Islands (SSI). Fifty-nine per cent of the bryozoans from KGI are endemic to Antarctica, which closely reflects the previously estimated endemism rate for bryozoans and other Antarctic taxa. Cluster analysis indicated that the strongest faunal links of SSI bryozoans were with Antarctic Peninsula assemblages, corresponding to the physical distance between both locations. The biogeographic similarities between SSI and South America confirm the broad trend of existing Antarctic–South American faunal links previously observed in bryozoans and many other taxa and indicate that SSI might be an important transitional zone between Antarctica and South America.     

Diversity in a Cold Hot-Spot: DNA-Barcoding Reveals Patterns of Evolution among Antarctic Demosponges (Class Demospongiae,Phylum Porifera)

Background

The approximately 350 demosponge species that have been described from Antarctica represent a faunistic component distinct from that of neighboring regions. Sponges provide structure to the Antarctic benthos and refuge to other invertebrates, and can be dominant in some communities. Despite the importance of sponges in the Antarctic subtidal environment, sponge DNA barcodes are scarce but can provide insight into the evolutionary relationships of this unique biogeographic province.

Methodology/Principal Findings

We sequenced the standard barcoding COI region for a comprehensive selection of sponges collected during expeditions to the Ross Sea region in 2004 and 2008, and produced DNA-barcodes for 53 demosponge species covering about 60% of the species collected. The Antarctic sponge communities are phylogenetically diverse, matching the diversity of well-sampled sponge communities in the Lusitanic and Mediterranean marine provinces in the Temperate Northern Atlantic for which molecular data are readily available. Additionally, DNA-barcoding revealed levels of in situ molecular evolution comparable to those present among Caribbean sponges. DNA-barcoding using the Segregating Sites Algorithm correctly assigned approximately 54% of the barcoded species to the morphologically determined species.

Conclusion/Significance

A barcode library for Antarctic sponges was assembled and used to advance the systematic and evolutionary research of Antarctic sponges. We provide insights on the evolutionary forces shaping Antarctica''s diverse sponge communities, and a barcode library against which future sequence data from other regions or depth strata of Antarctica can be compared. The opportunity for rapid taxonomic identification of sponge collections for ecological research is now at the horizon.     

Biogeographical and bioclimatic outline of Antarctica

Abstract

This study proposes a bioclimatic characterization and a new biogeographic division for the Antarctic territories up to the province level following the criteria and models of Rivas-Martínez et al. The Antarctic Kingdom comprises the continent of Antarctica, the surrounding ice-covered Antarctic islands, and the associated cold oceanic islands and archipelagos. It has two biogeographic regions: the Antarctic Region and the Subantarctic Insular Region. The Antarctic Region includes the entire pergelid Antarctic continent and the surrounding islands and archipelagos, and is characterized by upper suprapolar hyperoceanic and oceanic or Polar pergelid bioclimates on the coasts. The region has been divided into three pr6ovinces: Maritime Antarctica, West Antarctica and East Antarctica. The Subantarctic Insular Region comprises the circumantarctic islands and archipelagos that are widespread at the southern tip of the planet’s most important oceans, mostly in the subtemperate latitudinal zone inside or not far from the Antarctic Convergence. Bioclimatically, all insular subantarctic territories (excluding the South-American Tierra de Fuego, Terra Magellanica and large islands) are characterized by thermo-suprapolar and semipolar antarctic hyperoceanic bioclimates on the coasts. Four provinces – Falklandian-South Georgian, Kerguelenian, Macquarian and Aucklandian-Campbellian – have been recognized in this region. All these units are characterized by floristic bioindicators.     

Extinction and recolonization of maritime Antarctica in the limpet Nacella concinna (Strebel, 1908) during the last glacial cycle: toward a model of Quaternary biogeography in shallow Antarctic invertebrates

Quaternary glaciations in Antarctica drastically modified geographical ranges and population sizes of marine benthic invertebrates and thus affected the amount and distribution of intraspecific genetic variation. Here, we present new genetic information in the Antarctic limpet Nacella concinna, a dominant Antarctic benthic species along shallow ice‐free rocky ecosystems. We examined the patterns of genetic diversity and structure in this broadcast spawner along maritime Antarctica and from the peri‐Antarctic island of South Georgia. Genetic analyses showed that N. concinna represents a single panmictic unit in maritime Antarctic. Low levels of genetic diversity characterized this population; its median‐joining haplotype network revealed a typical star‐like topology with a short genealogy and a dominant haplotype broadly distributed. As previously reported with nuclear markers, we detected significant genetic differentiation between South Georgia Island and maritime Antarctica populations. Higher levels of genetic diversity, a more expanded genealogy and the presence of more private haplotypes support the hypothesis of glacial persistence in this peri‐Antarctic island. Bayesian Skyline plot and mismatch distribution analyses recognized an older demographic history in South Georgia. Approximate Bayesian computations did not support the persistence of N. concinna along maritime Antarctica during the last glacial period, but indicated the resilience of the species in peri‐Antarctic refugia (South Georgia Island). We proposed a model of Quaternary Biogeography for Antarctic marine benthic invertebrates with shallow and narrow bathymetric ranges including (i) extinction of maritime Antarctic populations during glacial periods; (ii) persistence of populations in peri‐Antarctic refugia; and (iii) recolonization of maritime Antarctica following the deglaciation process.     

Scotia Arc Acari: antiquity and origin

The Scotia Arc comprises South Georgia, South Sandwich, South Orkney and South Shetland Islands together with the Antarctic Peninsula. Free-living Acari of the Scotia Arc contain immigrant and endemic elements. Transport of immigrant species to and within the Maritime Antarctic has been via Holocene storms and ocean currents. Most immigrants are Gamasida, Oribatida and Acaridida while Actinedida dominate the endemic element. Immigrant species on South Georgia share common 'sub-Antarctic' affinities with South Indian and, to a lesser extent, South Pacific Ocean island faunas. In contrast, immigrants to the rest of the Scotia Arc and Bouvetøya on the mid-Atlantic Ridge, form a robust 'Maritime Antarctic' Province group. The endemic component is of largely Tertiary origin and, like that of Continental Antarctica, dominated by a few cosmopolitan families and genera of Actinedida. There are no bona fide pan-Antarctic species and little evidence that Continental and Maritime Antarctic faunas have a common ancestry, indeed the Continental endemic fauna is entirely montane while that of Maritime Antarctica is coastal. The presence of common Maritime Antarctic/South Pacific island genera corroborate the Antarctic Peninsula as being derived from a South Pacific island archipelago which collided with Continental Antarctica during the Tertiary period.     

Deep-Sea Polychaetous Annelida from Central America to the Antarctic Peninsula and South Sandwich Islands

Based on the 1958 Vema cruise and the 1965 Anton Bruun cruise 11 off the west coast of Central America and south through the islands east of the tip of South America, 143 trawl samples were examined for abyssal polychaetous annelida. The cruise tracks designated 7 major areas: Central America (1), Ecuador-Peru (II), Central and Southern Chile (III), Chile and the Magellan Strait (IV), Tierra del Fuego (V), the Island Complex (Falkland, South Georgia and South Sandwich Islands) (VI), the Drake Strait and Antarctic Peninsula (VII). The collections yielded 7,015 individuals and 322 species from 52 families. The relatively small number of species was attributed mainly to sampling effort, collecting gear and processing techniques. Each major area was characterized in terms of the number of individuals, the number of species, mean number of individuals, mean number of species and mean H′, SR, J′ and DI. Species diversity was significantly higher in Central America (I) than in the other areas. The relationships between biotic and abiotic measures (depth and areas of upwelling) were examined. There were no significant associations for the entire data base with depth. Moreover, there was no association between abundance of polychaetes and areas of upwelling which was hypothesized. In fact, relative abundance was highest in the deepest portions of Central America (I) (Guatemala Basin) and Ecuador—Peru (II) (Milne—Edwards Deep) compared to shallower areas closer to areas of upwelling. Quantitative collecting gear and refined processing techniques will be required to more accurately address these relationships. Dominant families were identified in terms of number of species and number of individuals. These families conformed well with world wide generalizations of dominant families. Dominant species were identified based on the Biological Index Value. The top twenty or dominant species for each major area were identified. A number of these dominant species occurred widely throughout the major areas. The dominant species for the entire data set were determined. In general these species were very eurybathic which was consistent with other worldwide depth distributions of abyssal polychaetes. Based on biotic measures (mean number of species, mean H', SR, J′ and DI) and the Simpson and Dice similarity co-efficients, several zoogeographic patterns emerged. The polychaete fauna from Central America (I) to southern Chile (II) formed a relatively homogeneous group. Polychaetes from Central America (I) had affinities with faunas studied off western Mexiko. The Chile and Magellan Strait (IV) area seemed to be a transition area which yielded to another relatively homogeneous fauna comprising areas V—VII. The latter fauna belonged to an Antarctic polychaete fauna which was documented earlier in other studies with a larger data base and more sophisticated analysis. It is hoped that the present study will help to provide a start to fill the zoogeographic gap of this enormous and bathymetrically diverse area.     

Mixoscyphus antarcticus gen. nov., sp. nov. (Cnidaria, Hydrozoa, Sertulariidae), the first truly endemic genus of Antarctic benthic hydroids

A genus and species of Antarctic benthic hydroids new to science, Mixoscyphus antarcticus gen. nov., sp. nov., is described and figured. Its systematic position amongst allied genera of the family Sertulariidae is discussed. The studied material originates from the South Shetland Islands area (West Antarctica), which was collected by several Spanish and US Antarctic expeditions. Mixoscyphus gen. nov. currently represents the only complete endemic genus of Antarctic benthic hydrozoans. A discussion of other genera of benthic hydroids that are largely endemic to the Antarctic is also provided.     

Accidental transfer of non-native soil organisms into Antarctica on construction vehicles

Antarctic terrestrial ecosystems currently include very few non-native species, due to the continent’s extreme isolation from other landmasses. However, the indigenous biota is vulnerable to human-mediated introductions of non-native species. In December 2005, four construction vehicles were imported by contractors to the British Antarctic Survey’s (BAS) Rothera Research Station (Antarctic Peninsula) from the Falkland Islands and South Georgia (South Atlantic) on board RRS James Clark Ross. The vehicles were contaminated with >132 kg of non-Antarctic soil that contained viable non-native angiosperms, bryophytes, micro-invertebrates, nematodes, fungi, bacteria, and c. 40,000 seeds and numerous moss propagules. The incident was a significant contravention of BAS operating procedures, the UK Antarctic Act (1994) and the Protocol on Environmental Protection to the Antarctic Treaty (1998), which all prohibit the introduction of non-native species to Antarctica without an appropriate permit. The introduction of this diverse range of species poses a significant threat to local biodiversity should any of the species become established, particularly as the biota of sub-Antarctic South Georgia is likely to include many species with appropriate pre-adaptations facilitating the colonisation of more extreme Antarctic environments. Once the incident was discovered, the imported soil was removed immediately from Antarctica and destroyed. Vehicle cleaning and transportation guidelines have been revised to enhance the biosecurity of BAS operations, and to minimise the risk of similar incidents occurring.     

Diversity and biogeography of the Antarctic flora

Aim To establish how well the terrestrial flora of the Antarctic has been sampled, how well the flora is known, and to determine the major patterns in diversity and biogeography. Location Antarctica south of 60° S, together with the South Sandwich Islands, but excluding South Georgia, Bouvetøya and the periantarctic islands. Methods Plant occurrence data were collated from herbarium specimens and literature records, and assembled into the Antarctic Plant Database. Distributional patterns were analysed using a geographic information system. Biogeographical patterns were determined with a variety of multivariate statistics. Results Plants have been recorded from throughout the Antarctic, including all latitudes between 60° S and 86° S. Species richness declines with latitude along the Antarctic Peninsula, but there was no evidence for a similar cline in Victoria Land and the Transantarctic mountains. Multi‐dimensional scaling ordinations showed that the species compositions of the South Orkney, South Shetland Islands and the north‐western Antarctic Peninsula are very similar to each other, as are the floras of different regions in continental Antarctica. They also suggest, however, that the eastern Antarctic Peninsula flora is more similar to the flora of the southern Antarctic Peninsula than to the continental flora (with which it has traditionally been linked). The South Sandwich Islands have a flora that is very dissimilar to that in all Antarctic regions, probably because of their isolation and volcanic nature. Main conclusions The Antarctic flora has been reasonably well sampled, but certain areas require further floristic surveys. Available data do, however, allow for a number of robust conclusions. A diversity gradient exists along the Antarctic Peninsula, with fewer species (but not fewer higher taxa) at higher latitudes. Multi‐dimensional scaling ordination suggests three major floral provinces within Antarctica: northern maritime, southern maritime, and continental. Patterns of endemism suggest that a proportion of the lichen flora may have an ancient vicariant distribution, while most bryophytes are more recent colonists.     

Open-ocean barriers to dispersal: a test case with the Antarctic Polar Front and the ribbon worm Parborlasia corrugatus (Nemertea: Lineidae)

Open-ocean environments provide few obvious barriers to the dispersal of marine organisms. Major currents and/or environmental gradients potentially impede gene flow. One system hypothesized to form an open-ocean dispersal barrier is the Antarctic Polar Front, an area characterized by marked temperature change, deep water, and the high-flow Antarctic Circumpolar current. Despite these potential isolating factors, several invertebrate species occur in both regions, including the broadcast-spawning nemertean worm Parborlasia corrugatus. To empirically test for the presence of an open-ocean dispersal barrier, we sampled P. corrugatus and other nemerteans from southern South America, Antarctica, and the sub-Antarctic islands. Diversity was assessed by analyzing mitochondrial 16S rRNA and cytochrome c oxidase subunit I sequence data with Bayesian inference and tcs haplotype network analysis. Appropriate neutrality tests were also employed. Although our results indicate a single well-mixed lineage in Antarctica and the sub-Antarctic, no evidence for recent gene flow was detected between this population and South American P. corrugatus. Thus, even though P. corrugatus can disperse over large geographical distances, physical oceanographic barriers (i.e. Antarctic Polar Front and Antarctic Circumpolar Current) between continents have likely restricted dispersal over evolutionary time. Genetic distances and haplotype network analysis between South American and Antarctic/sub-Antarctic P. corrugatus suggest that these two populations are possibly two cryptic species.     

Allochronic taxa as an alternative model to explain circumantarctic disjunctions

Abstract Most biogeographical studies propose that southern temperate faunal disjunctions are either the result of vicariance of taxa originated in Gondwana or the result of transoceanic dispersal of taxa originated after the breakup of Gondwana. The aim of this paper is to show that this is a false dichotomy. Antarctica retained a mild climate until mid‐Cenozoic and had lasting connections, notably with southern South America and Australia. Both taxa originally Gondwanan and taxa secondarily on Gondwanan areas were subjected to tectonic‐induced vicariance, and there is no need to invoke ad hoc transoceanic dispersal, even for post‐Gondwanan taxa. These different elements with circumantarctic distributions are here called ‘allochronic taxa’– taxa presently occupying the same area, but whose presence in that area does not belong to the same time period. This model allows accommodation of conflicting sources of evidence now available for many groups with circumantarctic distributions. The fact that the species from both layers are mixed up in the current biodiversity implies the need to use additional sources of evidence – such as biogeographical, palaeontological, geological and molecular – to discriminate which are the original Gondwanan and which are post‐Gondwanan elements in austral landmasses.     

Multiple colonization and dispersal events hide the early origin and induce a lack of genetic structure of the moss Bryum argenteum in Antarctica

The dispersal routes of taxa with transoceanic disjunctions remain poorly understood, with the potential roles of Antarctica not yet demonstrated. Mosses are suitable organisms to test direct intra‐Antarctic dispersal, as major component of the extant Antarctic flora, with the cosmopolitan moss Bryum argenteum as ideal target species. We analyzed the genetic structure of B. argenteum to provide an evolutionary time frame for its radiation and shed light into its historical biogeography in the Antarctic region. We tested two alternative scenarios: (a) intra‐Antarctic panmixia and (b) intra‐Antarctic genetic differentiation. Furthermore, we tested for evidence of the existence of specific intra‐Antarctic dispersal routes. Sixty‐seven new samples (40 collected in Antarctica) were sequenced for ITS nrDNA and rps4 cpDNA regions, and phylogenetic trees of B. argenteum were constructed, with a focus on its Southern Hemisphere. Combining our new nrDNA dataset with previously published datasets, we estimated time‐calibrated phylogenies based on two different substitution rates (derived from angiosperms and bryophytes) along with ancestral area estimations. Minimum spanning network and pairwise genetic distances were also calculated. B. argenteum was potentially distributed across Africa and Antarctica soon after its origin. Its earliest intra‐Antarctic dispersal and diversification occurred during a warming period in the Pliocene. On the same timescale, a radiation took place involving a dispersal event from Antarctica to the sub‐Antarctic islands. A more recent event of dispersal and diversification within Antarctica occurred during a warm period in the Pleistocene, creating favorable conditions also for its colonization outside the Antarctic continent worldwide. We provide evidence supporting the hypothesis that contemporary populations of B. argenteum in Antarctica integrate a history of both multiple long‐range dispersal events and local persistence combined with in situ diversification. Our data support the hypothesis that B. argenteum has been characterized by strong connectivity within Antarctica, suggesting the existence of intra‐Antarctic dispersal routes.     

Evidence of hidden biodiversity, ongoing speciation and diverse patterns of genetic structure in giant Antarctic amphipods

Recent molecular research on Antarctic benthic organisms has challenged traditional taxonomic classifications, suggesting that our current perceptions of Antarctic biodiversity and species distributions must be thoroughly revised. Furthermore, genetic differentiation at the intraspecific level remains poorly understood, particularly in eastern Antarctica. We addressed these issues using DNA sequence data for two sibling amphipod species that could be collected on a circum-Antarctic scale: Eusirus perdentatus and Eusirus giganteus. Haplotype networks and Bayesian phylogenies based on mitochondrial (COI, CytB) and nuclear (ITS2) DNA provided strong evidence of multiple cryptic species of Eusirus, with several occurring in sympatry and at least one likely to have a true circum-Antarctic distribution. Within species, gene flow was often highly restricted, consistent with a brooding life history and in some cases suggestive of current or future allopatric speciation. Patterns of genetic structure were not always predictable: one cryptic species showed preliminary evidence of high genetic differentiation across ～150 km in eastern Antarctica (F(ST) > 0.47, P < 0.01), yet another was remarkably homogenous across ～5000 km (F(ST) = 0.00, P = 1.00). Genetic diversity also varied among cryptic species, independent of sample size (π = 0.00-0.99). These results indicate several hidden levels of genetic complexity in these Antarctic amphipods that are neither apparent from previous taxonomic or ecological studies nor predictable from their life history. Such genetic diversity and structure may reflect different modes of survival for Antarctic benthic organisms during historic glacial cycles, and/or subsequent re-establishment of populations on the shelf, and highlight our misunderstanding of Antarctic marine species diversity.     

Invasive non‐native species likely to threaten biodiversity and ecosystems in the Antarctic Peninsula region

The Antarctic is considered to be a pristine environment relative to other regions of the Earth, but it is increasingly vulnerable to invasions by marine, freshwater and terrestrial non‐native species. The Antarctic Peninsula region (APR), which encompasses the Antarctic Peninsula, South Shetland Islands and South Orkney Islands, is by far the most invaded part of the Antarctica continent. The risk of introduction of invasive non‐native species to the APR is likely to increase with predicted increases in the intensity, diversity and distribution of human activities. Parties that are signatories to the Antarctic Treaty have called for regional assessments of non‐native species risk. In response, taxonomic and Antarctic experts undertook a horizon scanning exercise using expert opinion and consensus approaches to identify the species that are likely to present the highest risk to biodiversity and ecosystems within the APR over the next 10 years. One hundred and three species, currently absent in the APR, were identified as relevant for review, with 13 species identified as presenting a high risk of invading the APR. Marine invertebrates dominated the list of highest risk species, with flowering plants and terrestrial invertebrates also represented; however, vertebrate species were thought unlikely to establish in the APR within the 10 year timeframe. We recommend (a) the further development and application of biosecurity measures by all stakeholders active in the APR, including surveillance for species such as those identified during this horizon scanning exercise, and (b) use of this methodology across the other regions of Antarctica. Without the application of appropriate biosecurity measures, rates of introductions and invasions within the APR are likely to increase, resulting in negative consequences for the biodiversity of the whole continent, as introduced species establish and spread further due to climate change and increasing human activity.     

Areas of endemism in the Antarctic – a case study of the benthic hydrozoan genus Oswaldella (Cnidaria,Kirchenpaueriidae)

Aim The aim of this study is to investigate areas of endemism within the distribution of Oswaldella species in the Southern Ocean, thereby testing previous hypotheses and proposing alternative scenarios for Antarctic evolution. Location Southern Ocean, Antarctic and sub-Antarctic waters of southern South America. Methods We prepared a database for the 31 currently known species of the Antarctic genus Oswaldella, which includes geographical locations gathered from published taxonomic studies as well as materials from museums and expeditions. A parsimony analysis of endemicity (PAE) was used to test hypotheses of distribution patterns. Results Four areas of endemism are hypothesized: southern South America, two high Antarctic areas (eastern and western) and a larger area, mainly in western Antarctica at lower latitudes and including insular areas (but not the Balleny Islands). Main conclusions The results support, in part, previous hypotheses for the Southern Ocean region, while providing more detailed resolution. The areas of endemism may reflect both historical and ecological processes that influenced the Antarctic biota. The Magellanic area reflects the well-known affinities of the Antarctic biota with that of South America and may be a consequence of dispersal through deeper (and colder) waters, followed by speciation. The second area, the largest one, encompasses most of the insular faunas and may also be associated with deeper waters formed since 43 Ma. The third area may be explained by the development of seaways in the circum-Antarctic region beginning 50 Ma. Finally, the fourth zone, with a very poor fauna, coincides with the opening of the Tasman Strait and the formation of the Australo-Antarctic Gulf, associated with a minor wind-driven current.     

The polar lichens Caloplaca darbishirei and C. soropelta highlight the direction of bipolar migration

A proper phytogeographic affiliation of Antarctic lichen species has become feasible using molecular phylogeographic methods. Caloplaca citrina is a heterogeneous taxon including several species which occurs in polar regions and is common in Antarctica. Collections of C. citrina from the Antarctic were revised using morphological, anatomical and molecular characters (ITS). They were found to belong to two species: Caloplaca darbishirei (C.W. Dodge & G.E. Baker) Cretz. and C. soropelta (E.S. Hansen, Poelt & S?chting) S?chting. The molecular phylogeny showed them to be sister species, but well separated. Morphological and chemical characters, ecology and distribution of the species are discussed. C. darbishirei is the most common species in the Antarctic, and it is so far known only from Antarctica and Southern South America. C. soropelta, reported here as new to South America, is a bipolar species with all close relatives in the Southern Hemisphere; it is therefore most likely that the species colonized the Arctic from the south. C. citrina s. str. is not confirmed to occur in Antarctica. The study emphasizes the suitability of genotyping for understanding the taxonomy and phylogeography of bipolar lichens.     

Ocean barriers and glaciation: evidence for explosive radiation of mitochondrial lineages in the Antarctic sea slug Doris kerguelenensis (Mollusca, Nudibranchia)

Strong currents and deep passages of water can be barriers for larval dispersal of continental marine animals, but potential effects on direct developers are under-investigated. We examined the genetic structure of Doris kerguelenensis , a directly developing sea slug that occurs across the Drake Passage, the body of water separating Antarctica from South America. We found deep mitochondrial divergences within populations on both sides of the Drake Passage, and South American animals formed multiple sister-group relationships with Antarctic animals. A generalised molecular clock suggested these trans-Drake pairs diverged during the Pliocene–Pleistocene, after the formation of the Drake Passage. Statistical parsimony methods recovered 29 separate haplotype networks (many sympatric) that likely correlate with allopatric events caused by repeated glacial cycles. Data from 16S were congruent but more conserved than COI, and the estimated ancestral 16S haplotype was widespread. The marked difference in the substitution rates between these two mitochondrial genes results in different estimates of connectivity. Demographic analyses on networks revealed some evidence for selection and expanding populations. Contrasting with the Northern Hemisphere, glaciation in Antarctica appears to have increased rather than reduced genetic diversity. This suggests orbitally forced range dynamics based on Northern Hemisphere phylogeography do not hold for Antarctica. The diverse lineages found in D. kerguelenensis point towards a recent, explosive radiation, likely reflecting multiple refuges during glaciation events, combined with limited subsequent dispersal. Whether recognised as cryptic species or not, genetic diversity in Antarctic marine invertebrates appears higher than expected from morphological analyses, and supports the Antarctic biodiversity pump phenomenon.