The biology, life cycle and ecophysiology of the Antarctic mite Alaskozetes antarcticus

This paper is dedicated to the late Nigel Bonner, who as Head of the Life Sciences Division at British Antarctic Survey, encouraged and supported this research with his characteristic enthusiasm.
The cryptostigmatid mite Alaskozetes antarcticus (Michael) is a dominant member of many terrestrial communities in the maritime Antarctic, where it survives extreme temperatures, short cold summers, numerous freeze-thaw cycles, desiccating conditions and a limited season for growth and reproduction. However, examination of features of its biology, from morphology, through life-history strategy to physiology, indicate very little specialization to the Antarctic environment. Alaskozetes antarcticus is a herbivore/detritivore, typical of the Cryptostigmata in general, with low feeding and growth rates, long life span and low reproductive output. Physiological specializations exist in the form of low enzyme activation energies and elevated metabolic rates at low temperatures when compared with temperate species, and associated low optimum temperatures for activity, feeding and growth. Growth rates comparable with temperate species are achieved in the field, with an extended life cycle of five years or more as a result of the short growing season, and the ability of all life stages to overwinter equally successfully. Overwintering survival, involving supercooling enhanced by the use of antifreezes such as glycerol, although initially described in Antarctic species, is now known to be characteristic of many temperate relatives, so it is not a specific adaptation to the polar environment. The obvious success of A. antarcticus in maritime Antarctic terrestrial environments must be attributed to a combination of several features characteristic of the Cryptostigmata in general, rather than to specific polar adaptations.     

Aerobiology and colonization in Antarctica — the BIOTAS Programme

Little is known about the aerial transport of microbes, plants and animals into and between Antarctic terrestrial and freshwater habitats. Isolation by the circumpolar Southern Ocean restricts propagules to three main groups: 1) those capable of prolonged survival in the air, 2) those carried by animal vectors, especially Man, and 3) those of marine origin. Diverse ice-free areas available for colonization include: isolated islands with receding ice sheets (e.g., Signy Island), maritime geothermal areas (e.g., Deception Island), and high altitude gcothermal areas on continental volcanoes (e.g., Mt. Erebus). Propagule abundance and diversity are low in Antarctica. Chance affects colonization success because the potential and viability of a propagule must match a favourable habitat for settlement, with adequate time for establishment before conditions become unfavourable.

Aerobiological studies of the whole Antarctic region require international co-operation. The Scientific Committee on Antarctic Research (SCAR) Biological Investigations of Terrestrial Antarctic Systems (BIOTAS) research network has identified aerobiology as a major component of its International Research Programme.

Aerobiology will be ground-, ship- and aircraft-based. To meet the requirements of an intrinsically sparse, diverse Antarctic aerobiota, the British Antarctic Survey is developing a new particle sampler for remote field use.     

Environmental constraints on life histories in Antarctic ecosystems: tempos, timings and predictability

Knowledge of Antarctic biotas and environments has increased dramatically in recent years. There has also been a rapid increase in the use of novel technologies. Despite this, some fundamental aspects of environmental control that structure physiological, ecological and life-history traits in Antarctic organisms have received little attention. Possibly the most important of these is the timing and availability of resources, and the way in which this dictates the tempo or pace of life. The clearest view of this effect comes from comparisons of species living in different habitats. Here, we (i) show that the timing and extent of resource availability, from nutrients to colonisable space, differ across Antarctic marine, intertidal and terrestrial habitats, and (ii) illustrate that these differences affect the rate at which organisms function. Consequently, there are many dramatic biological differences between organisms that live as little as 10 m apart, but have gaping voids between them ecologically.Identifying the effects of environmental timing and predictability requires detailed analysis in a wide context, where Antarctic terrestrial and marine ecosystems are at one extreme of the continuum of available environments for many characteristics including temperature, ice cover and seasonality. Anthropocentrically, Antarctica is harsh and as might be expected terrestrial animal and plant diversity and biomass are restricted. By contrast, Antarctic marine biotas are rich and diverse, and several phyla are represented at levels greater than global averages. There has been much debate on the relative importance of various physical factors that structure the characteristics of Antarctic biotas. This is especially so for temperature and seasonality, and their effects on physiology, life history and biodiversity. More recently, habitat age and persistence through previous ice maxima have been identified as key factors dictating biodiversity and endemism. Modern molecular methods have also recently been incorporated into many traditional areas of polar biology. Environmental predictability dictates many of the biological characters seen in all of these areas of Antarctic research.     

Polar Marine Communities

SYNOPSIS. This paper offers a sweeping but very superficialreview of the marine biology of polar seas. The marine systemsin the Arctic and Antarctic have in common polar positions andcold temperatures, otherwise they are strikingly different.The Arctic has broad shallow continental shelves with seasonallyfluctuating physical conditions and a massive fresh water impactin the northern coastal zones. However, it has a low seasonalityof pack ice and little vertical mixing. In contrast, the Antarctichas over twice the oceanic surface area, deep narrow shelves,and, except for ice cover, a relatively stable physical environmentwith very little terrestrial input. The Antarctic has greatpack ice seasonality and much vertical mixing. Primary productivityin the polar areas tends to be strongly pulsed with the zooplanktonlagging behind; however there are many exceptions to such generalizations.Most recent research has focused on specific patterns and processesresulting in biological hot spots such as predictable leadsin the ice, polynyas, oceanographic fronts, areas of intensemixing, and the marginal ice zone. This review attempts to weavethese recent oceanographic studies into the geological historyof each habitat in an effort to develop a holistic understandingof the biological processes     

Temporal metabolic rate variation in a continental Antarctic springtail

Terrestrial systems in Antarctica are characterized by substantial spatial and temporal variation. However, few studies have addressed the paucity of data on metabolic responses to the unpredictable Antarctic environment, particularly with regard to terrestrial biota. This study measured metabolic rate variation for individual springtails at a continental Antarctic site using a fiber-optic closed respirometry system incorporating a custom-made respiration chamber. Concurrent measures of (behavioural) activity were made via daily pitfall counts.Metabolic rate of Gomphiocephalus hodgsoni measured at constant temperature varied systematically with progression through the austral summer, and was greatest mid-season. This finding of clear intra-seasonal and temperature-independent variation in mass-specific metabolic rate in G. hodgsoni is one of very few such reports for a terrestrial invertebrate (and the only such study for Antarctica), and parallels physiological studies in the Antarctic marine environment linking metabolic rate elevation with biological function rather than temperature adaptation per se. However, response to temperature at relatively short time-scales is also likely to be an important part of the life history strategy of Antarctic terrestrial invertebrates such as G. hodgsoni, which appears capable of both physiologically and behaviourally ‘tuning’ in to short-term thermal variability to respond appropriately to the local unpredictable Antarctic habitat.     

Problems of Pollen Zone Correlation in Southeastern Canada

A brief account is given documenting the development of aerobiological research in the Antarctic. The results of the British Antarctic Survey's contribution to an international programme on long-distance dispersal of aeroplankton over the Southern Ocean are presented. This was achieved by collecting airspora deposited in Tauber traps and in surface snow at sites on South Georgia (sub-Antarctic) and Signy Island (maritime Antarctic). Although only a small number of the samples were analysed, the results provided ample evidence of a continuous immigration of exotic sporomorpha of southern South American provenance. The cause of this rain of biological material is attributed to the not infrequent easterly tracking storm events generated over the south-east Pacific Ocean. As they gain momentum over southern South America they become seeded with pollen and spores, and possibly by larger organelles such as invertebrates and seeds. These high winds may be deflected southwards by a blocking anticyclone over the South Atlantic Ocean, allowing a proportion of the sporomorpha to be deposited over land far to the south. The occurrence of such exotic sporomorpha in these remote and environmentally hostile regions is used here as evidence to support the hypothesis that there is a continuous input into the Antarctic biome of viable propagules from more northerly landmasses. While no exotic bryophyte or lichen spores have yet been detected in trapping experiments, the extremely rare occurrence of certain bryophytes associated only with geothermal sites in the Antarctic and in laboratory-cultured soils from barren ice-free terrain indicates that a pool of viable but dormant propagules is probably widespread in Antarctic soils and ice. However, germination and development in situ are possible only under exceptional environmental circumstances. An international programme is being planned to detect the main trajectories of long-distance transport of propagules into the Antarctic and to test their viability.     

Antarctic terrestrial life--challenging the history of the frozen continent?

Antarctica is a continent locked in ice, with almost 99.7% of current terrain covered by permanent ice and snow, and clear evidence that, as recently as the Last Glacial Maximum (LGM), ice sheets were both thicker and much more extensive than they are now. Ice sheet modelling of both the LGM and estimated previous ice maxima across the continent give broad support to the concept that most if not all currently ice-free ground would have been overridden during previous glaciations. This has given rise to a widely held perception that all Mesozoic (pre-glacial) terrestrial life of Antarctica was wiped out by successive and deepening glacial events. The implicit conclusion of such destruction is that most, possibly all, contemporary terrestrial life has colonised the continent during subsequent periods of glacial retreat. However, several recently emerged and complementary strands of biological and geological research cannot be reconciled comfortably with the current reconstruction of Antarctic glacial history, and therefore provide a fundamental challenge to the existing paradigms. Here, we summarise and synthesise evidence across these lines of research. The emerging fundamental insights corroborate substantial elements of the contemporary Antarctic terrestrial biota being continuously isolated in situ on a multi-million year, even pre-Gondwana break-up timescale. This new and complex terrestrial Antarctic biogeography parallels recent work suggesting greater regionalisation and evolutionary isolation than previously suspected in the circum-Antarctic marine fauna. These findings both require the adoption of a new biological paradigm within Antarctica and challenge current understanding of Antarctic glacial history. This has major implications for our understanding of the key role of Antarctica in the Earth System.     

Antarctic terrestrial biodiversity in a changing world

Recent analyses of Antarctic terrestrial biodiversity data, in combination with molecular biological studies, have created a new paradigm that long-term persistence and regional isolation are general features of most of the major groups of Antarctic terrestrial biota, overturning the previously widely assumed view of a generally recent colonisation history. This paradigm, as well as incorporating a new and much longer timescale in which to consider the evolution and adaptation of Antarctic terrestrial biota, opens important new cross-disciplinary linkages with geologists and glaciologists seeking to unravel the history of the continent itself. This unique biota now faces the twin challenges of responding to the complex processes of climate change facing some parts of the continent, and the direct impacts associated with human occupation and activity. In many instances, this biota is likely to benefit, initially at least, from the current environmental changes, and there is an expectation of increased production, biomass, population size, community complexity, and colonisation. However, the impacts of climate change may themselves be outweighed by other, direct, impacts of human activities, and in particular, the introduction of non-indigenous organisms from which until recently the terrestrial ecosystems of the continent have been protected. The Antarctic research community and those responsible for governance under the Antarctic treaty system are faced with the pressing challenges of (1) ensuring there is sufficient baseline monitoring and survey activity to enable identification of these changes, however caused and (2) ensuring that effective operational management and biosecurity procedures are in place to minimise the threat from anthropogenic activities.     

Survival strategies in polar terrestrial arthropods

Three components of the survival strategy of a terrestrial Antarctic mite, Alaskozetes antarcticus (Acari: Cryptostigmata) are considered: overwintering survival, energetics and life history. Supercooling is an important feature of its cold tolerance, whilst elevation of standard metabolism allows activity at low temperatures, both of which contribute tcTa long development and maximum survival of individuals in the population. These are facets of the overall survival strategy evolved by such a species in response to the Antarctic terrestrial environment, but which may be widespread in polar invertebrates.     

Ecological contrasts across an Antarctic land–sea interface

Abstract We report the composition of terrestrial, intertidal and shallow sublittoral faunal communities at sites around Rothera Research Station, Adelaide Island, Antarctic Peninsula. We examined primary hypotheses that the marine environment will have considerably higher species richness, biomass and abundance than the terrestrial, and that both will be greater than that found in the intertidal. We also compared ages and sizes of individuals of selected marine taxa between intertidal and subtidal zones to test the hypothesis that animals in a more stressed environment (intertidal) would be smaller and shorter lived. Species richness of intertidal and subtidal communities was found to be similar, with considerable overlap in composition. However, terrestrial communities showed no overlap with the intertidal, differing from previous reports, particularly from further north on the Antarctic Peninsula and Scotia Arc. Faunal biomass was variable but highest in the sublittoral. While terrestrial communities were depauperate with low biomass they displayed the highest overall abundance, with a mean of over 3 × 10⁵ individuals per square metre. No significant differences in ages of intertidal and subtidal individuals of the same species were found, with bryozoan colonies of up to 4 years of age being present in the intertidal. In contrast with expectation and the limited existing literature we conclude that, while the Antarctic intertidal zone is clearly a suboptimal and highly stressful habitat, its faunal community can be well established and relatively diverse, and is not limited to short‐term opportunists or waifs and strays.     

Controls on the distribution of productivity and organic resources in Antarctic Dry Valley soils

The Antarctic Dry Valleys are regarded as one of the harshest terrestrial habitats on Earth because of the extremely cold and dry conditions. Despite the extreme environment and scarcity of conspicuous primary producers, the soils contain organic carbon and heterotrophic micro-organisms and invertebrates. Potential sources of organic compounds to sustain soil organisms include in situ primary production by micro-organisms and mosses, spatial subsidies from lacustrine and marine-derived detritus, and temporal subsidies ('legacies') from ancient lake deposits. The contributions from these sources at different sites are likely to be influenced by local environmental conditions, especially soil moisture content, position in the landscape in relation to lake level oscillations and legacies from previous geomorphic processes. Here we review the abiotic factors that influence biological activity in Dry Valley soils and present a conceptual model that summarizes mechanisms leading to organic resources therein.     

Evolution and biodiversity of Antarctic organisms: a molecular perspective

The Antarctic biota is highly endemic, and the diversity and abundance of taxonomic groups differ from elsewhere in the world. Such characteristics have resulted from evolution in isolation in an increasingly extreme environment over the last 100 Myr. Studies on Antarctic species represent some of the best examples of natural selection at the molecular, structural and physiological levels. Analyses of molecular genetics data are consistent with the diversity and distribution of marine and terrestrial taxa having been strongly influenced by geological and climatic cooling events over the last 70 Myr. Such events have resulted in vicariance driven by continental drift and thermal isolation of the Antarctic, and in pulses of species range contraction into refugia and subsequent expansion and secondary contact of genetically distinct populations or sister species during cycles of glaciation. Limited habitat availability has played a major role in structuring populations of species both in the past and in the present day. For these reasons, despite the apparent simplicity or homogeneity of Antarctic terrestrial and marine environments, populations of species are often geographically structured into genetically distinct lineages. In some cases, genetic studies have revealed that species defined by morphological characters are complexes of cryptic or sibling species. Climate change will cause changes in the distribution of many Antarctic and sub-Antarctic species through affecting population-level processes such as life history and dispersal.     

Living on the edge – plants and global change in continental and maritime Antarctica

Antarctic terrestrial ecosystems experience some of the most extreme growth conditions on Earth and are characterized by extreme aridity and subzero temperatures. Antarctic vegetation is therefore at the physiological limits of survival and, as a consequence, even slight changes to growth conditions are likely to have a large impact, rendering Antarctic terrestrial communities sensitive to climate change. Climate change is predicted to affect the high‐latitude regions first and most severely. In recent decades, the Antarctic has undergone significant environmental change, including the largest increases in ultraviolet‐B (UV‐B; 290–320 nm) radiation levels in the world and, in the maritime region at least, significant temperature increases. This review describes the current evidence for environmental change in Antarctica, and the impacts of this change on the terrestrial vegetation. This is largely restricted to cryptogams, such as bryophytes, lichens and algae; only two vascular plant species occur in the Antarctic, both restricted to the maritime region. We review the range of ecological and physiological consequences of increasing UV‐B radiation levels, and of changes in temperature, water relations and nutrient availability. It is clear that climate change is already affecting the Antarctic terrestrial vegetation, and significant impacts are likely to continue in the future. We conclude that, in order to gain a better understanding of the complex dynamics of this important system, there is a need for more manipulative, long‐term field experiments designed to address the impacts of changes in multiple abiotic factors on the Antarctic flora.     

Life, decay and fossilisation of endolithic microorganisms from the Ross Desert, Antarctica

The adaptation and survival of the endolithic microorganisms that colonise the near-surface layer of porous sandstone rock in the Ross Desert (Antarctica) depend upon a precarious equilibrium of biological, geological and climatic factors. Any unfavourable change in external conditions can result in the death and disappearance of microscopic organisms, and this may be followed by trace microfossil formation. The sequence of events leading to the extinction of life in the Antarctic desert is considered to be a terrestrial analogue of the disappearance of possible life on early Mars. The present paper reviews the current state of knowledge on the endolithic microorganisms of the Ross Desert with particular reference to their decay and fossilisation processes. Ideas for in situ further research on this microbial ecosystem are also proposed, including several new microscopy techniques such as CLSM, LTSEM, SEM-BSE and EDS. Preliminary images are presented and it is proposed that, for the first time, such techniques will permit the in situ study of the ecology of Antarctic lithobiontic microorganisms and the identification and characterisation of fossilised traces of past life.     

Diversity of Antarctic terrestrial protozoa

Heterotrophic protozoa have a global distribution in terrestrial habitats. The functional groups significantly represented are zooflagellates, cillates, gymnamoebae and testate amoebae. Their range extends into the Antarctic zone, but the species richness of the communities is rarely of the same order of magnitude as those in temperate latitudes. Species diversity is usually very low owing to dominance of the communities by single, or a few, species which are best adapted to the Antarctic terrestrial environment. This is characterized by seasonal, diurnal or unpredictable fluctuations in moisture, temperature and bacterial food supply of high amplitude. The fauna shows pauperization with latitude and climatic severity. Nearly all records of species distribution are consistent with the model that community composition is determined by local conditions. An important exception is the distribution of the testate amoeba genus Nebela whose species distribution is influenced by biogeographical factors. Successional changes in community composition in fellfield habitats are characterized by the sequence: pioneer microflagellate colonizers, larger flagellates and small ciliates, and finally testate amoebae. The succession is most closely correlated with the accumulation of organic matter. A model of the strategies of dominant microflagellate species can be constructed by ordinating them on a two-dimensional habitat template of A-r-K selection continuum. The globally ubiquitous microflagellate Heteromita globosa emerges as the most strongly A-selected and K-selected. The occurrence of terrestrial protozoa near their latitudinal limits of distribution can serve as sensitive indicators of the biological effects of climatic change. Having short generation times and effective means of cyst dispersal, changes in the gross distribution can provide rapid warning of critical changes in thermal regimes.     

The cyanobacterial community of polygon soils at an inland Antarctic nunatak

Inland Antarctic terrestrial ecosystems and biodiversity are poorly understood in comparison with Antarctic coastal regions. Microorganisms, as primary colonists, are integral to Antarctic soil ecosystem development, essential for pedogenesis and structuring the soil, and providing the nutrients necessary for the subsequent establishment of macroorganisms. This study analysed the microbial communities present in polygon soils of Coal Nunatak (Alexander Island, at the southern limit of the maritime Antarctic). Soils were analysed across three polygons (centre and margins) and at three depths (0–1, 1–2, 2–5 cm). Cyanobacterial communities were characterised using two complementary molecular biological approaches, temperature gradient gel electrophoresis and clone library analysis. The three polygons exhibited conspicuous differences in community composition, both between different polygons and spatially (horizontally and vertically) within a single polygon. Comparison of our data with that from previous studies using classical culture and morphological identification techniques clearly shows the need for more intensive research on patterns of microbial diversity in terrestrial habitats throughout the Antarctic. The majority of the 17 cyanobacterial genera identified at Coal Nunatak are thought to have ubiquitous distributions, while none are known only from the Antarctic. Three of the genera present are also known to be capable of being lichen photobionts.     

Cyanobacteria in Antarctica: ecology, physiology and cold adaptation.

Cyanobacterial species composition of fresh water and terrestrial ecosystems and chemical environment of water in Schirmacher Oasis in Continental Antarctica was investigated. Over 35 species of cyanobacteria were recorded. Diazotrophic species both heterocystous and unicellular contributed more than half to the count except in lake ecosystem. The species composition varied among the fresh water as well as terrestrial ecosystems. The physico-chemical analyses of water revealed its poor nurient content which might have supported the growth of diazotrophic cyanobacteria in an Antarctic environment. Among the cyanobacteria Oscillatoria, Phormidium and Nostoc commune were the dominant flora in most of the habitats. The physiological characteristics of isolated cyanobacteria strains indicated that N2-fixation, nitrate uptake, nitrate-reduction, ammonium-uptake, GS-transferase activity and photosynthesis was unaffected at low temperature (5 degrees C) which indicated low temperature adaptation for Antarctic cyanobacteria. This phenomenon was not evident in different strains of tropical origin. The temperature optima for N2-fixation for the different Antarctic cyanobacterial strains was in the range of 15-25 degrees C, nearly 10 degrees C lower than their respective reference strains of tropical origin. Similar results were obtained for cyanobacteria-moss association. The low endergonic activation energy exhibited by the above metabolic activities supported the view that cyanobacteria were adapted to Antarctic ecosystem.     

Reconsidering connectivity in the sub‐Antarctic

Extreme and remote environments provide useful settings to test ideas about the ecological and evolutionary drivers of biological diversity. In the sub‐Antarctic, isolation by geographic, geological and glaciological processes has long been thought to underpin patterns in the region's terrestrial and marine diversity. Molecular studies using increasingly high‐resolution data are, however, challenging this perspective, demonstrating that many taxa disperse among distant sub‐Antarctic landmasses. Here, we reconsider connectivity in the sub‐Antarctic region, identifying which taxa are relatively isolated, which are well connected, and the scales across which this connectivity occurs in both terrestrial and marine systems. Although many organisms show evidence of occasional long‐distance, trans‐oceanic dispersal, these events are often insufficient to maintain gene flow across the region. Species that do show evidence of connectivity across large distances include both active dispersers and more sedentary species. Overall, connectivity patterns in the sub‐Antarctic at intra‐ and inter‐island scales are highly complex, influenced by life‐history traits and local dynamics such as relative dispersal capacity and propagule pressure, natal philopatry, feeding associations, the extent of human exploitation, past climate cycles, contemporary climate, and physical barriers to movement. An increasing use of molecular data – particularly genomic data sets that can reveal fine‐scale patterns – and more effective international collaboration and communication that facilitates integration of data from across the sub‐Antarctic, are providing fresh insights into the processes driving patterns of diversity in the region. These insights offer a platform for assessing the ways in which changing dispersal mechanisms, such as through increasing human activity and changes to wind and ocean circulation, may alter sub‐Antarctic biodiversity patterns in the future.     

Low cyanobacterial diversity in biotopes of the Transantarctic Mountains and Shackleton Range (80-82°S), Antarctica

The evolutionary history and geographical isolation of the Antarctic continent have produced a unique environment rich in endemic organisms. In many regions of Antarctica, cyanobacteria are the dominant phototrophs in both aquatic and terrestrial ecosystems. We have used microscopic and molecular approaches to examine the cyanobacterial diversity of biotopes at two inland continental Antarctic sites (80-82°S). These are among the most southerly locations where freshwater-related ecosystems are present. The results showed a low cyanobacterial diversity, with only 3-7 operational taxonomic units (OTUs) per sample obtained by a combination of strain isolations, clone libraries and denaturing gradient gel electrophoresis based on 16S rRNA genes. One OTU was potentially endemic to Antarctica and is present in several regions of the continent. Four OTUs were shared by the samples from Forlidas Pond and the surrounding terrestrial mats. Only one OTU, but no internal transcribed spacer (ITS) sequences, was common to Forlidas Pond and Lundstr?m Lake. The ITS sequences were shown to further discriminate different genotypes within the OTUs. ITS sequences from Antarctic locations appear to be more closely related to each other than to non-Antarctic sequences. Future research in inland continental Antarctica will shed more light on the geographical distribution and evolutionary isolation of cyanobacteria in these extreme habitats.