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.     

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.     

Proceedings of the SMBE Tri-National Young Investigators' Workshop 2005. Southern hemisphere springtails: could any have survived glaciation of Antarctica?

Throughout the Southern Hemisphere many terrestrial taxa have circum-Antarctic distributions. This pattern is generally attributed to ongoing dispersal (by wind, water, or migrating birds) or relict Gondwanan distributions. Few of these terrestrial taxa have extant representatives in Antarctica, but such taxa would contribute to our understanding of the evolutionary origins of the continental Antarctic fauna. Either these taxa have survived the harsh climate cooling in Antarctica over the last 23 Myr (Gondwanan/vicariance origin) or they have dispersed there more recently (<2 MYA). In this context, we examined mtDNA (COI) sequence variation in Cryptopygus and related extant Antarctic and subantarctic terrestrial springtails (Collembola). Sequence divergence was estimated under a maximum likelihood model (general time reversible+I+Gamma) between individuals from subantarctic islands, Australia, New Zealand, Patagonia, Antarctic Peninsula, and continental Antarctica. Recent dispersal/colonization (<2 MYA) of Cryptopygus species was inferred between some subantarctic islands, and there was a close association between estimated times of divergences based on a molecular clock and proposed geological ages of islands. Most lineages generally grouped according to geographic proximity or by inferred dispersal/colonization pathways. In contrast, the deep divergences found for the four endemic Antarctic species indicate that they represent a continuous chain of descent dating from the break up of Gondwana to the present. We suggest that the diversification of these springtail species (21-11 MYA) in ice-free glacial refugia throughout the Trans-Antarctic Mountains was caused by the glaciation of the Antarctic continent during the middle to late Miocene.     

Biological invasions in the Antarctic: extent, impacts and implications

Alien microbes, fungi, plants and animals occur on most of the sub-Antarctic islands and some parts of the Antarctic continent. These have arrived over approximately the last two centuries, coincident with human activity in the region. Introduction routes have varied, but are largely associated with movement of people and cargo in connection with industrial, national scientific program and tourist operations. The large majority of aliens are European in origin. They have both direct and indirect impacts on the functioning of species-poor Antarctic ecosystems, in particular including substantial loss of local biodiversity and changes to ecosystem processes. With rapid climate change occurring in some parts of Antarctica, elevated numbers of introductions and enhanced success of colonization by aliens are likely, with consequent increases in impacts on ecosystems. Mitigation measures that will substantially reduce the risk of introductions to Antarctica and the sub-Antarctic must focus on reducing propagule loads on humans, and their food, cargo, and transport vessels.     

Biodiversity and biogeography of Southern Ocean pycnogonids

The pycnogonids of the Southern Ocean have been studied for almost two centuries and have played a key role in shaping previous biogeographic regions for the Antarctic benthos. The aim of this study was to assess the biogeographic patterns derived from the most current sample records of pycnogonids from the Southern Ocean and neighbouring areas. 332 species of pycnogonids from 1837 sample locations were analysed using 279 3° by 3° grid cells. We investigated richness patterns and the effect of sampling intensity at both local and regional scales, and used multivariate analysis of distribution patterns and species assemblages to define biogeographic trends. These analyses identified a distinct and isolated Antarctic pycnogonid shelf fauna which was different to that of the deep‐sea around Antarctica, the Sub‐Antarctic islands, South America or New Zealand. Within the Antarctic, we found the South Shetland Islands to be the most speciose region and a probable center of radiation for the pycnogonids. No latitudinal gradients in species richness were detected. We note that the distribution patterns observed are based upon classical taxonomy and discuss the potential for changes to these patterns with new insights from molecular techniques. We conclude that, even with the potential for cryptic species, the large‐scale biogeographic trends observed in the pycnogonids should hold true.     

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.     

Brittle stars from Southern Ocean (Echinodermata: Ophiuroidea)

The present biogeographic study on the ophiuroid fauna from the Southern Ocean (SO) contains an updated checklist, based on a compilation of all the published information provided for the Antarctic and sub-Antarctic regions as well as the information available in SCAR-MarBIN database. Faunal composition and geographical and bathymetric distribution are included. So far, 219 species have been recorded, of which 126 are endemic to the SO, 76 are exclusive to Antarctic waters, and 30 are exclusive to sub-Antarctic waters. This study corroborated the circumpolar and eurybathic character of the ophiuroid fauna of the SO, but some differences are discussed when considering shelf and deep-sea fauna in the whole SO, or in the Antarctic and sub-Antarctic regions separately. The biogeographic affinities of 17 areas considered in the SO are revised, based on a presence/absence datamatrix of the 219 species. This similarity analysis shows three main groups, two of them including sub-Antarctic areas and one for Antarctic areas. The faunal movement patterns between the main geographical connections have been based on historical site records of each species. These movements have a level of faunal exchange that exceeds that of other Antarctic benthic groups. Such movements are mainly from Antarctic and sub-Antarctic regions to the subtropical waters of South America, and from New Zealand and southern Australian waters to sub-Antarctic areas. In this context, the origin of the ophiuroid Antarctic fauna is discussed.     

The significance of the sub-Antarctic Kerguelen Islands for the assessment of the vulnerability of native communities to climate change,alien insect invasions and plant viruses

The suite of environments and anthropogenic modifications of sub-Antarctic islands provide key opportunities to improve our understanding of the potential consequences of climate change and biological species invasions on terrestrial ecosystems. The profound impact of human introduced invasive species on indigenous biota, and the facilitation of establishment as a result of changing thermal conditions, has been well documented on the French sub-Antarctic Kerguelen Islands (South Indian Ocean). The present study provides an overview of the vulnerability of sub-Antarctic terrestrial communities with respect to two interacting factors, namely climate change and alien insects. We present datasets assimilated by our teams on the Kerguelen Islands since 1974, coupled with a review of the literature, to evaluate the mechanism and impact of biological invasions in this region. First, we consider recent climatic trends of the Antarctic region, and its potential influence on the establishment, distribution and abundance of alien insects, using as examples one fly and one beetle species. Second, we consider to what extent limited gene pools may restrict alien species’ colonisations. Finally, we consider the vulnerability of native communities to aliens using the examples of one beetle, one fly, and five aphid species taking into consideration their additional impact as plant virus vectors. We conclude that the evidence assimilated from the sub-Antarctic islands can be applied to more complex temperate continental systems as well as further developing international guidelines to minimise the impact of alien species.     

Evolutionary and ecological aspects of some Antarctic and sub-Antarctic penguin distributions

Penguins probably originated in the core of Gondwanaland when South America, Africa, and Antarctica were just beginning to separate. As the continents drifted apart, the division filled with what became the southern ocean. One of the remaining land masses moved south and was caught at the pole by the Earth's rotation. It became incrusted with ice and is now known as East Antarctica. Linking it to South America was a series of submerged mountain ranges that formed a necklace of islands. The northern portion of the necklace, called the Scotia Arc, is now the "fertile crescent" of the Southern Ocean. The greatest numbers and biomass of penguins are found here as well as that of krill, the primary prey species of most penguins, and many other marine predators. Today penguins are found throughout the sub-Antarctic islands and around the entire Antarctic continent. Using satellite transmitters and time-depth recorders, while taking advantage of the parental dedication of breeding birds, numerous investigators have described foraging habits of several species of penguins. The information obtained is labor intensive and costly so that studies are restricted to certain species, areas and seasons. Here I review the patterns evident among six of the most abundant and completely studied of the penguins. The variation in behavior is considerable from those species that seldom dive deeper than 20 m in search of prey to those that will dive to depths >500 m to catch mesopelagic fish and squid. Foraging trips from breeding colonies vary among species and with the season. Often the birds travel no more than 30 km and at other times the trips may exceed 600 km. Sub-Antarctic species often reach more productive waters near or within the Antarctic Polar Front zone, where the mixing of Antarctic and sub-Antarctic waters provide rich resources for their prey. Antarctic species usually remain close to shore, along the continental slope, or near the sea ice edge. Less is known about penguins during the pelagic phase between breeding cycles. What we do know is surprising in regard to their dispersal, which ranges from hundreds to thousands of kilometers from the breeding colonies.     

Detecting glacial refugia in the Southern Ocean

Throughout the Quaternary, the continental-based Antarctic ice sheets expanded and contracted repeatedly. Evidence suggests that during glacial maxima, grounded ice eliminated most benthic (bottom-dwelling) fauna across the Antarctic continental shelf. However, paleontological and molecular evidence indicates most extant Antarctica benthic taxa have persisted in situ throughout the Quaternary. Where and how the Antarctic benthic fauna survived throughout repeated glacial maxima remain mostly hypothesised. If understood, this would provide valuable insights into the ecology and evolution of Southern Ocean biota over geological timescales. Here we synthesised and appraised recent studies and presented an approach to demonstrate how genetic data can be effective in identifying where and how Antarctic benthic fauna survived glacial periods. We first examined the geological and ecological evidence for how glacial periods influenced past species demography in order to provide testable frameworks for future studies. We outlined past ice-free areas from Antarctic ice sheet reconstructions that could serve as glacial refugia and discussed how benthic fauna with pelagic or non-pelagic dispersal strategies moved into and out of glacial refugia. We also reviewed current molecular studies and collated proposed locations of Southern Ocean glacial refugia on the continental shelf around Antarctica, in the deep sea, and around sub-Antarctic islands. Interestingly, the proposed glacial refugia based on molecular data generally do not correspond to the ice-free areas identified by Antarctic ice sheet reconstructions. The potential biases in sampling and in the choice of molecular markers in current literature are discussed, along with the future directions for employing testable frameworks and genomic methods in Southern Ocean molecular studies. Continued data syntheses will elucidate greater understanding of where and how Southern Ocean benthic fauna persisted throughout glacial periods and provide insights into their resilience against climate changes in the future.     

Conservation biogeography of the Antarctic

Aim

To present a synthesis of past biogeographic analyses and a new approach based on spatially explicit biodiversity information for the Antarctic region to identify biologically distinct areas in need of representation in a protected area network.

Location

Antarctica and the sub‐Antarctic.

Methods

We reviewed and summarized published biogeographic studies of the Antarctic. We then developed a biogeographic classification for terrestrial conservation planning in Antarctica by combining the most comprehensive source of Antarctic biodiversity data available with three spatial frameworks: (1) a 200‐km grid, (2) a set of areas based on physical parameters known as the environmental domains of Antarctica and (3) expert‐defined bioregions. We used these frameworks, or combinations thereof, together with multivariate techniques to identify biologically distinct areas.

Results

Early studies of continental Antarctica typically described broad bioregions, with the Antarctic Peninsula usually identified as biologically distinct from continental Antarctica; later studies suggested a more complex biogeography. Increasing complexity also characterizes the sub‐Antarctic and marine realms, with differences among studies often attributable to the focal taxa. Using the most comprehensive terrestrial data available and by combining the groups formed by the environmental domains and expert‐defined bioregions, we were able to identify 15 biologically distinct, ice‐free, Antarctic Conservation Biogeographic Regions (ACBRs), encompassing the continent and close lying islands.

Main conclusions

Ice‐free terrestrial Antarctica comprises several distinct bioregions that are not fully represented in the current Antarctic Specially Protected Area network. Biosecurity measures between these ACBRs should also be developed to prevent biotic homogenization in the region.     

The Acari of fresh- and brackish water habitats in the Antarctic and sub-Antarctic regions

Neither the benthic Halacaroidea nor nektonic Hydrachnidia, characteristic of temperate and tropical freshwater mite faunas are represented among the twenty four species and sub-species of Acari recorded from the Antarctic and sub-Antarctic. The mites collected from the streams, lakes or brackish/freshwater pools of the region are either terrestrial species blown by wind into freshwater or interlopers from the marine littoral which have invaded coastal brackish pools.The present study reports on new collections of mites obtained from lakes and pools at a number of locations from both Continental and Maritime Antarctica, as well as from several sub-Antarctic islands. It reviews all published records for the region and the combined fauna is then compared with that of temperate and tropical regions.     

Seaweed biodiversity in the south-western Antarctic Peninsula: surveying macroalgal community composition in the Adelaide Island/Marguerite Bay region over a 35-year time span

The diversity of seaweed species of the south-western Antarctic Peninsula region is poorly studied, contrasting with the substantial knowledge available for the northern parts of the Peninsula. However, this is a key region affected by contemporary climate change. Significant consequences of this change include sea ice recession, increased iceberg scouring and increased inputs of glacial melt water, all of which can have major impacts on benthic communities. We present a baseline seaweed species checklist for the southern Adelaide Island and northern Marguerite Bay region, combining data obtained during a small number of surveys completed in 1973–1975 and a 6-week intensive diving-based field campaign in 2010–2011. Overall, with a total of 41 macroalgal species recorded (7 brown, 27 red, 6 green, 1 chrysophyte), the region is species-poor compared to the north of the Antarctic Peninsula, and even more so in comparison with the sub-Antarctic. The key canopy-forming species is Desmarestia menziesii, which is abundant in Antarctic Peninsula waters, but lacking in the sub-Antarctic. Himantothallus grandifolius, which is a common species further north in the Antarctic phytobenthos, was absent in our recent collections. This paper also reports the first record of Aplanochytrium sp. (Labyrinthulomycetes) from this part of Antarctica and in association with Elachista sp.     

Large‐scale oceanographic fluctuations drive Antarctic petrel survival and reproduction

Polar Regions are experiencing environmental changes at unprecedented rates. These changes can spread throughout entire food webs from lower trophic levels to apex predators. As many top predators forage over large areas, these indirect effects may be associated with large‐scale patterns of climate variability. Using global climate indices that are known to impact the Southern Ocean ecosystem (the El Niño Southern Oscillation and Antarctic Oscillation Indices) we assessed their efficacy to predict variation in the demographic parameters of Antarctic seabirds. First, we used a long‐term dataset on adult survival (estimated from capture–mark–recapture data) and reproduction of Antarctic petrel Thalassoica antarctica, from the largest known breeding colony (Svarthamaren, Dronning Maud Land) and examined whether large‐scale oceanographic fluctuations impact survival and reproduction. Second, we conducted an exhaustive literature review to determine whether the effects of large‐scale environmental variability on Antarctic seabirds have a coherent fingerprint across the Antarctic continent and nearby islands. We found that most of the variation in both reproductive success, timing of hatching, and survival of Antarctic petrels can be accurately modeled using the two modes of large‐scale climate variability in Antarctica. The literature review, combined with the results from our field study, suggests that while the anticipated trends in the global patterns of climatic variability will generally have detrimental effects on populations of top predators in the Southwest Atlantic, these conclusions cannot be extrapolated to all seabird populations in Antarctica without additional data.     

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.     

Mite dispersal among the Southern Ocean Islands and Antarctica before the last glacial maximum

It has long been maintained that the majority of terrestrial Antarctic species are relatively recent, post last glacial maximum, arrivals with perhaps a few microbial or protozoan taxa being substantially older. Recent studies have questioned this 'recolonization hypothesis', though the range of taxa examined has been limited. Here, we present the first large-scale study for mites, one of two dominant terrestrial arthropod groups in the region. Specifically, we provide a broad-scale molecular phylogeny of a biologically significant group of ameronothroid mites from across the maritime and sub-Antarctic regions. Applying different dating approaches, we show that divergences among the ameronothroid mite genera Podacarus, Alaskozetes and Halozetes significantly predate the Pleistocene and provide evidence of independent dispersals across the Antarctic Polar Front. Our data add to a growing body of evidence demonstrating that many taxa have survived glaciation of the Antarctic continent and the sub-Antarctic islands. Moreover, they also provide evidence of a relatively uncommon trend of dispersals from islands to continental mainlands. Within the ameronothroid mites, two distinct clades with specific habitat preferences (marine intertidal versus terrestrial/supralittoral) exist, supporting a model of within-habitat speciation rather than colonization from marine refugia to terrestrial habitats. The present results provide additional impetus for a search for terrestrial refugia in an area previously thought to have lacked ice-free ground during glacial maxima.     

Planktonic microbial assemblages and the potential effects of metazooplankton predation on the food web of lakes from the maritime Antarctica and sub-Antarctic islands

Antarctica is the continent with the harshest climate on the Earth. Antarctic lakes, however, usually presents liquid water, at least during part of the year or below the ice cover, especially those from the sub-Antarctic islands and the maritime Antarctic region where climatic conditions are less extreme. Planktonic communities in these lakes are mostly dominated by microorganisms, including bacteria and phototrophic and heterotrophic protists, and by metazooplankton, usually represented by rotifers and calanoid copepods, the latter mainly from the genus Boeckella. Here I report and discuss on studies performed during the last decade that show that there is a potential for top–down control of the structure of the planktonic microbial food web in sub-Antarctic and maritime Antarctic lakes. In some of the studied lakes, the effect of copepod grazing on protozoa, either ciliates or flagellates, depending on size of both the predator and the prey, could promote cascade effects that would be transmitted to the bacterioplankton assemblage.     

Biogeographic traits and checklist of Antarctic demosponges

Summary The biogeography of Antarctic demosponges has been studied by dividing Antarctic and circumantarctic areas into geographic entities, and then assigning to these entities all recorded species according to literature reports. Correspondence analysis ordination based on the presence or absence of species shows the existence of a distinct Antarctic Faunistic Complex (AFC), including continental Antarctica, most of the Antarctic and circumantarctic islands and the Magellan area. Such a result has enabled us to drawup a checklist of 352 Antarctic demosponge species. Investigation of within-AFC patterns indicates that the continent is a highly homogeneous area, establishing closer relationships with the Scotia Arc and to a lesser extent with the Magellan region. The AFC has low specific affinities with the other circumantarctic regions (South Africa, temperate Australia and New Zealand), whereas at the generic level relationships appear more pronounced. This biogeographic pattern may lead us to suppose a common Gondwanian origin for Antarctic and circumantarctic sponge faunas, followed by differentiation due to Gondwana fragmentation. Antarctica moved towards polar latitudes and became progressively isolated, only maintaining active interchanges with South America. Climatic changes possibly induced intense processes of speciation in the Antarctic demosponge fauna, thus contributing to its differentiation.     

Conservation of Southern Ocean Islands: invertebrates as exemplars

The Southern Ocean Islands (SOI) have an exceptionally high conservation status, and human activity on the islands is low by comparison with more tropical islands. In consequence, overexploitation, pollution and habitat destruction have had little influence on the invertebrate biotas of the islands, although overexploitation of pelagic species has the potential for an indirect influence via reduction of nutrient inputs to the terrestrial systems. By contrast, invasive alien species, the local effects of global climate change, and interactions between them are having large impacts on invertebrate populations and, as a consequence, on ecosystem functioning. Climate change is not only having direct impacts on indigenous invertebrates, but also seems to be promoting the ease of establishment of new alien invertebrate species. It is also contributing to population increases of invertebrate alien species already on the islands, sometimes with pronounced negative consequences for indigenous species and ecosystem functioning. Moreover, alien plants and mammals are also affecting indigenous invertebrate populations, often with climate change expected to exacerbate the impacts. Although the conservation requirements are reasonably well-understood for terrestrial systems, knowledge of freshwater and marine near-shore systems is inadequate. Nonetheless, what is known for terrestrial, freshwater and marine systems suggests that ongoing conservation of SOI invertebrates requires intervention from the highest political levels internationally, to slow climate change, to local improvements of quarantine measures to reduce the rates and impacts of biological invasions.