Modelling reproductive effort in sub-and maritime Antarctic mosses

A comparison is made of the reproductive effort (RE), considered as the investment in sporophyte relative to gametophyte biomass, of eight species of moss occurring at sub-and maritime Antarctic sites. Six of the species showed smaller sporophytes and game-tophytes at the climatically more extreme maritime Antaretic sites and one species showed no size difference between regions. The remaining species, although showing no regional difference, showed some evidence of a reverse pattern, with higher altitude samples having greater biomass than lower altitude samples. Spore counts indicated a measure of compensation in maritime Antarctic samples, with no significant decrease in spore output in several species despite smaller sporophyte biomass. The relationship between sporophyte (S) and gametophyte (G) biomass within samples was described by an allometric curve (S=aG ^b ) which gave a better fit than a straight line for six species. This form of model allows comparisons of patterns of RE to be made between samples with non-or partially overlapping size distributions, even when the relationship involves size-dependence. An allometric curve was not appropriate for describing samples of one species (Andreaea regularis), and insufficient data were available to identify any relationship in Polytrichum alpinum. The exponent (b) differed between species, but there were no statistically significant differences between exponents from samples of the same species. Samples of two species could further be described by the same coefficient (a), indicating that they lie on the same curve. However, samples of three species from sub-Antarctic South Georgia gave significantly higher coefficients, indicating increased RE relative to maritime Antarctic populations.     

Taxonomic affinities of dark septate root endophytes of Colobanthus quitensis and Deschampsia antarctica,the two native Antarctic vascular plant species

Although dark septate fungal endophytes (DSE) occur widely in association with plant roots in cold-stressed habitats, little is known of the taxonomic status of DSE in Antarctica. Here we investigate the phylogenetic affinities of DSE colonising the roots of Colobanthus quitensis and Deschampsia antarctica, the two maritime Antarctic vascular plant species. Two hundred and forty-three DSE were isolated from roots collected from 17 sites across a 1 470 km transect through maritime and sub-Antarctica. The ITS1-5.8S-ITS2 nuclear ribosomal gene cluster of representative isolates was sequenced, and the sequences were recovered in 10 sequence groups and sub-groups. Nine of the sequence groupings could be placed in the Helotiales and the remaining one showed high homology to a large number of currently unassigned anamorphic ascomycete sequences. Of the Helotiales, Leptodontidium orchidicola, Rhizoscyphus ericae and species of Tapesia and Mollisia could be confidently identified. This study demonstrates that members of the Helotiales, including several widely-recognised DSE genera, commonly colonise the roots of C. quitensis and D. antarctica in the Antarctic.     

Polychaeta “Sedentaria” of Terra Nova Bay (Ross Sea,Antarctica): Capitellidae to Serpulidae

Capitellidae, Maldanidae, Trichobranchidae, Terebllidae, Sabellidae, Serpulidae and Spirorbidae found in Terra Nova Bay (Ross Sea, Antarctica) have been studied. Thirty-one species have been identified, one of which, Praxillella praetermissa antantica is a new subspecies. About 75% of the species are endemic to Antarctica or sub-Antarctica. The geographic distribution of some species has been extended.     

Invited review: climate change impacts in polar regions: lessons from Antarctic moss bank archives

Mosses are the dominant plants in polar and boreal regions, areas which are experiencing rapid impacts of regional warming. Long‐term monitoring programmes provide some records of the rate of recent climate change, but moss peat banks contain an unrivalled temporal record of past climate change on terrestrial plant Antarctic systems. We summarise the current understanding of climatic proxies and determinants of moss growth for contrasting continental and maritime Antarctic regions, as informed by 13C and 18O signals in organic material. Rates of moss accumulation are more than three times higher in the maritime Antarctic than continental Antarctica with growing season length being a critical determinant of growth rate, and high carbon isotope discrimination values reflecting optimal hydration conditions. Correlation plots of 13C and 18O values show that species (Chorisodontium aciphyllum / Polytrichum strictum) and growth form (hummock / bank) are the major determinants of measured isotope ratios. The interplay between moss growth form, photosynthetic physiology, water status and isotope composition are compared with developments of secondary proxies, such as chlorophyll fluorescence. These approaches provide a framework to consider the potential impact of climate change on terrestrial Antarctic habitats as well as having implications for future studies of temperate, boreal and Arctic peatlands. There are many urgent ecological and environmental problems in the Arctic related to mosses in a changing climate, but the geographical ranges of species and life‐forms are difficult to track individually. Our goal was to translate what we have learned from the more simple systems in Antarctica, for application to Arctic habitats.     

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.     

An insect introduction to the maritime Antarctic

Although several invertebrates have been introduced by Man into the Antarctic, no holometabolous insects have survived to colonize terrestrial habitats successfully. Data are presented on the survival of populations of a chironomid midge, together with an enchytraeid worm, for 17 years in a maritime Antarctic site at Signy Island, South Orkney Islands. Both species are thought to have been introduced on plant material transplanted from either South Georgia or the Falkland Islands or both in 1967. Population densities average 25718m^-2 for the dipteran larvae and 3243 m^-2 for the worms. Successful completion of the midge's life cycle was indicated by emergence of brachypterous adults and oviposition (the population is parthenogenetic with only females present). Although both taxa are capable of supercooling to between -13 and -26°C, this capacity may not be sufficient in a severe winter to avoid lethal freezing. Four potential cryoprotectants were found in insect extracts, but in concentrations (< 1 % fresh weight) unlikely to influence cold hardiness. Both invertebrates appear to be pre-adapted for survival in much harsher conditions than they normally experience, by the extension of existing physiological mechanisms. It is concluded that the main limitations to invertebrate colonization of suitable Antarctic land areas by soil-dwelling species are geographical.     

Microbial diversity and function in Antarctic freshwater ecosystems

Freshwater lakes occur through much of Antarctica and are characterized by short food chains dominated by microbes. Comparatively few studies have been made of continental freshwater lakes until recently, with the main emphasis being on the less extreme maritime Antarctic lakes. The wide range of trophic status seen at the northern extremes of the maritime Antarctic reduces markedly further south, but a wide range of micro-organisms occur throughout the latitudinal range. Information on seasonal and spatial patterns of microbial activity for freshwater lakes demonstrate rapid changes in community composition at certain times of year despite constant low temperatures. Benthic communities of cyanobacteria and bacteria are a feature of most lakes and are involved in a wide range of geochemical cycling. There is a need for more detailed taxonomic information on most groups and considerable potential for molecular studies.     

A Note on the United States and the Law of the Sea: Looking Back and Moving Forward

Since the Antarctic Treaty was negotiated in 1959, there have been substantial developments in the law of the sea. One of the most significant developments has been the recognition granted to coastal state entitlements to claim a range of offshore maritime areas. Yet, one of the principal aims of the Antarctic Treaty was to eliminate sovereignty disputes between territorial claimants, and the treaty placed limitations on the assertion of new claims. Nevertheless, most Antarctic territorial claimants have asserted some form of Antarctic maritime claim. This article particularly considers Australia's position toward maritime claims offshore the Australian Antarctic Territory (AAT). It reviews the limitations imposed by the Antarctic Treaty, the difficulties in determining baselines in Antarctica, Australia's practice in declaring Antarctic maritime claims, and the potential range of maritime boundaries that Australia may one day have to delimit with neighboring states in the Southern Ocean.     

An examination of species dispersion patterns along the intertidal gradient on Macquarie Island (sub-Antarctica) using a restricted occupancy model

It has been widely held that intertidal zonation boundaries and the tidal emersion levels are causally related: the critical tidal level hypothesis. In order to evaluate this hypothesis, the dispersion patterns of species boundaries on the intertidal rocky shores of Macquarie Island (sub-Antarctica) were examined using a restricted occupancy model to test the null hypothesis that the observed species were randomly dispersed along the tidal emersion gradient. Subsequently this investigation demonstrated that the intertidal species boundaries on Macquarie Island shores were randomly dispersed with respect to the tidal emersion gradient. Hence no prima facie evidence was found to support the critical tidal level hypothesis. This suggests that tidal emersion is not a significant factor structuring intertidal communities on Macquarie Island.     

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.     

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.     

Molecular and morphological characterization of the Achnanthidium minutissimum complex (Bacillariophyta) in Petuniabukta (Spitsbergen,High Arctic) including the description of A. digitatum sp. nov.

Recent morphology-based investigations of freshwater Arctic diatoms suggest that many species remain to be discovered, reflecting a unique polar flora. During a survey of the freshwater diatom flora of northern Billefjorden, including the Petuniabukta fjord region (Spitsbergen), several morphodemes belonging to the Achnanthidium minutissimum species complex were recorded. Molecular phylogenies based on rbcL, 28S and 18S sequences, including single cells from Canada and strains from Marion Island (sub-Antarctica) and GenBank revealed the presence of 12 distinct A. minutissimum complex lineages, of which three contained strains from Spitsbergen. One Arctic lineage is described as a new species. Achnanthidium digitatum sp. nov. is morphologically characterized by narrow, linear to only slightly lanceolate valves and usually two areolae per stria. The two remaining Arctic lineages are in need of a more complete morphological and molecular comparison with other representatives of the A. minutissimum complex to clarify their taxonomic identity. It is argued that implementation of molecular data in the taxonomy of Achnanthidium will be essential to solve the taxonomic problems associated with this group, eventually resulting in a better understanding of the biogeography and niche differentiation of different species belonging to the A. minutissimum complex. Ideally, this should be based on more variable genes than the currently widely used 18S, which does not have a species level resolution in the A. minutissimum complex.     

Isolation and molecular identification of free-living amoebae of the genus Naegleria from Arctic and sub-Antarctic regions

Twenty-three freshwater samples with sediment taken from two regions in the Arctic, Spitzbergen and Greenland, and one region in sub-Antarctica, Ile de la Possession, were cultured for amoebae at 37 degrees C and room temperature (RT). Only two samples yielded amoebae at 37 degrees C and the two isolates were identified from their morphological features to belong to the genus Acanthamoeba. Vahlkampfiid amoebae were isolated from 11 samples at RT. Morphological analysis of the cysts identified all 11 isolates as belonging to the genus Naegleria, although only about half of them (45%) transformed into flagellates. Ribosomal DNA sequence analysis demonstrated that these isolates represent novel species and that N. antarctica, N. dobsoni and N. chilensis are their closest relatives. Not surprisingly, these three species also grow at lower temperatures (<37 degrees C) than the majority of described Naegleria spp. Two of the eight new species were found in both Arctic and sub-Antarctic regions, and other new species from the Arctic are closely related to new species from the sub-Antarctic. Therefore, it seems the Naegleria gene pool present in the polar regions is different from that found in temperate and tropical regions.     

Preface

Abstract

Major naval weapons and weapons systems are examined as they relate to naval strategy and the law of the sea. Although there are some legal limitations on the development and use of nuclear and other mass destruction weapons on the seabed, and in nuclear free zones such as Latin America, and on naval weaponry in Antarctica, the use of naval weapons systems is largely unrestricted by international law and the law of the sea. Weapons systems from mines to large submarines and carriers are related to areas, to categories of states, and to the main sub‐fields of international relations. The conclusion to be drawn is that a third Hague Conference on the laws of war should be called and arms control and disarmament issues in naval weaponry should be addressed urgently. Detailed data on naval weapons, missile systems, and maritime zones are presented in three appendixes. Glossary of Abbreviations for Naval Weaponry immediately precedes the appendices.     

Temporal resolution of cold acclimation and de-acclimation in the Antarctic collembolan, Cryptopygus antarcticus

Abstract.  The Antarctic collembolan, Cryptopygus antarcticus (Willem), can switch its supercooling point (SCP) between 'winter' and 'summer' modes of cold hardiness over a matter of hours. High resolution temporal scaling of the acquisition and loss of cold hardiness is undertaken by assaying changes in the proportion of animals freezing below −15 °C in response to cooling rate, acclimation temperature, and access to food and moisture. Rapid de-acclimation to the 'summer' modal state is readily achieved after 1–6 h in response to warming and access to food; however, rapid acclimation to the 'winter' modal state is only evident in response to slow cooling and narrow ranges of temperature (0–5 °C). The rapid loss of cold tolerance at higher temperatures with access to food, in particular, emphasizes this species' opportunistic responses to resource availability in the short polar summers. Cold hardiness is apparently more readily traded off against nutrient acquisition than vice versa in this maritime Antarctic species.     

Feeding repellence of Antarctic and sub-Antarctic benthic invertebrates against the omnivorous sea star Odontaster validus

Antarctic and sub-Antarctic benthic invertebrates are subjected to intense predation by mobile macroinvertebrates. Accordingly, chemical protection as well as other defensive mechanisms are expected to be common in organisms inhabiting these ecosystems. In order to evaluate anti-predation activities and allocation of chemical defenses within the anatomy of marine benthic Antarctic and sub-Antarctic invertebrates, 55 species were tested for feeding repellence against the sea star Odontaster validus, a common eurybathic sympatric predator. The invertebrates tested were collected from the deep waters of two poorly surveyed areas in terms of chemical ecology studies: the eastern Weddell Sea (Antarctica) and the vicinities of Bouvet Island (sub-Antarctica). Experiments were conducted at the Spanish Antarctic Base in Deception Island. In the feeding deterrence experiments, shrimp pieces were treated with crude lipophilic fractions obtained from each species, and were offered to the sea stars. A total of 29 species (53 %) from 7 different phyla (Porifera, Cnidaria, Chordata, Bryozoa, Echinodermata, Mollusca, and Annelida) showed feeding repellence against O. validus, and are therefore chemically protected against this keystone predator. Furthermore, 25 species were dissected into parts to investigate the possible allocation of defensive compounds. Some of the results obtained from these analyses support the prediction that the most exposed/vulnerable tissues concentrate chemical defenses to avoid predation against the sea stars. In summary, the results obtained in our survey support the hypothesis that deep-water Antarctic and sub-Antarctic benthic invertebrates are well protected chemically against sympatric predators, similarly to what has been reported in previous studies investigating shallow-water Antarctic species.     

Early life history strategies of notothenioids at South Georgia

Antarctic notothenioid early life history strategies are examined in general and then for common species at South Georgia. Channichthyids, bathydraconids, artedidraconids and some nototheniids have large eggs 3·0–4·9 mm whereas other nototheniids and arpagiferids have smaller eggs 1·6–2·7 mm. At South Georgia the larvae of species with large eggs hatched between August (late winter) and late November (late spring) at 11–16 mm standard length ( L _s). Larvae of species with small eggs hatched mainly during October and December at 4.5–9 mm L _s. Most of the larvae of all species attain urostyle flexion between October and January, and develop to the end of the larval stage between November and May. The duration of the larval stage varies from 2 months in species with smaller larvae to 6 months for some of the species with larger larvae. Two nototheniid species develop to the early juvenile stage before a channichthyid and a bathydraconid that hatch around 2 months earlier. During their first winter, the early-juveniles of most species with large eggs are pelagic, whereas those of species with small eggs may be pelagic or demersal. Four groups of strategies are proposed based on egg size and the winter ecotype of the early-juvenile stage.     

Eretmoptera murphyi: pre‐adapted to survive a colder climate

During the late 1960s, larvae of the flightless midge Eretmoptera murphyi Schaeffer were accidentally transferred from the sub‐Antarctic island of South Georgia to Signy Island in the maritime Antarctic. Higher insects are rare in the Antarctic and the introduction and establishment of a new species is an unusual event. The fly has overcome the two major barriers to colonization of the Antarctic by new species: the geographical isolation of the region and its severe climate. Larvae of the flightless midge overwinter in the surface layers of soil on Signy Island where the temperature may fall to below ?10 °C, compared with as little as ?1.5 °C on South Georgia. This suggests the possession of a level of pre‐adaption to colder conditions. Summer‐collected larvae have a supercooling point (SCP or whole body freezing point) of approximately ?5.0 °C but survive experimental exposure to ?13 °C, giving them a level of freeze tolerance. After acclimation at ?4 °C for 4 days, the SCP changes little but the temperature at which 50% of the population would die decreases to lower than ?19 °C. Larvae are also resistant to dehydration. Under experimental conditions of 88% relative humidity at 5 °C, larvae lose water linearly (0.42% h^?1) over the first 30 h but resist further water loss once their water content decreases to approximately 1.4 g g^?1 dry weight. All larvae survive these conditions for the duration of the experiment (55 h). Eretmoptera murphyi is well adapted to survive on Signy Island, and these studies suggest that it has the ability to survive at more extreme locations at higher latitudes if it were to be inadvertently transferred to a suitable habitat.     

The “Trojan horse” strategy: Seed fungal endophyte symbiosis helps to explain the invasion success of the grass,Poa annua,in Maritime Antarctica

Aim

Poa annua L. (annual bluegrass) is presently the sole invasive vascular plant species to have successfully established in Maritime Antarctica, where it poses a significant conservation threat to native plant species. However, the reasons for its success in the region have yet to be established. Here, we determined whether the invasiveness of P. annua, and its competitiveness with the native Antarctic hairgrass Deschampsia antarctica, is influenced by symbioses formed with seed fungal endophytes, and whether plants derived from seeds from four global regions differ in their performance.

Locations

Four regions (Maritime Antarctica, sub-Antarctica, South America and Europe).

Methods

Endophyte frequency was measured in P. annua seeds collected from the four regions. The germination, survival, biomass accumulation, flowering and competitiveness with D. antarctica of P. annua plants grown from endophyte-uncolonised and uncolonised seeds was determined in the laboratory. The effects of endophytes on P. annua seed germination and survival and seedling osmoprotection were also assessed in the Maritime Antarctic natural environment using locally-sourced seeds.

Results

Endophytes were at least twice as frequent in seeds from Maritime Antarctica than in those from other regions. A higher proportion of endophyte-colonized seeds germinated and survived than did uncolonised seeds, but only when they originated from Maritime Antarctica. Seed endophytes increased the competitiveness of P. annua with D. antarctica, but only for plants grown from Maritime Antarctic seeds. In the field, endophyte-colonized seeds from Maritime Antarctica germinated and survived more frequently than uncolonised seeds, and osmoprotection was higher in seedlings grown from colonized seed.

Main Conclusions

The findings indicate beneficial effects of seed endophytes on invasion-related traits of P. annua, such as survival, germination success and flowering. Together with vegetative and reproductive traits facilitating the colonization process, the seed-fungal endophyte symbiosis can be invoked as an important factor explaining the invasiveness of P. annua in Maritime Antarctica.     

Deschampsia antarctica: genetic and molecular-biological aspects of spreading in Antarctica

The genetical and molecular biological attributes of Deschampsia antarctica are considered in relation to its remarkable success in the colonization of the maritime Antarctic. However, none of the data from several published research studies provide convincing evidence that this species possess any obvious unique or specialised adaptations which might account for this success. While its occurrence appears to have spanned much of the Holocene, there is no evidence that there have ever been any other native vascular plant species in the maritime Antarctic. It is still unclear why no other taxa have succeeded in colonising this region. Further intensive studies of genetics and molecular biology of this species, and of other potential immigrants, may provide a better understanding of their enigmatic success in the maritime Antarctic.