Molecular analysis of geographic patterns of eukaryotic diversity in Antarctic soils

We describe the application of molecular biological techniques to estimate eukaryotic diversity (primarily fungi, algae, and protists) in Antarctic soils across a latitudinal and environmental gradient between approximately 60 and 87 degrees S. The data were used to (i) test the hypothesis that diversity would decrease with increasing southerly latitude and environmental severity, as is generally claimed for "higher" faunal and plant groups, and (ii) investigate the level of endemicity displayed in different taxonomic groups. Only limited support was obtained for a systematic decrease in diversity with latitude, and then only at the level of a gross comparison between maritime (Antarctic Peninsula/Scotia Arc) and continental Antarctic sites. While the most southerly continental Antarctic site was three to four times less diverse than all maritime sites, there was no evidence for a trend of decreasing diversity across the entire range of the maritime Antarctic (60 to 72 degrees S). Rather, we found the reverse pattern, with highest diversity at sites on Alexander Island (ca. 72 degrees S), at the southern limit of the maritime Antarctic. The very limited overlap found between the eukaryotic biota of the different study sites, combined with their generally low relatedness to existing sequence databases, indicates a high level of Antarctic site isolation and possibly endemicity, a pattern not consistent with similar studies on other continents.     

Molecular Analysis of Geographic Patterns of Eukaryotic Diversity in Antarctic Soils

We describe the application of molecular biological techniques to estimate eukaryotic diversity (primarily fungi, algae, and protists) in Antarctic soils across a latitudinal and environmental gradient between approximately 60 and 87°S. The data were used to (i) test the hypothesis that diversity would decrease with increasing southerly latitude and environmental severity, as is generally claimed for “higher” faunal and plant groups, and (ii) investigate the level of endemicity displayed in different taxonomic groups. Only limited support was obtained for a systematic decrease in diversity with latitude, and then only at the level of a gross comparison between maritime (Antarctic Peninsula/Scotia Arc) and continental Antarctic sites. While the most southerly continental Antarctic site was three to four times less diverse than all maritime sites, there was no evidence for a trend of decreasing diversity across the entire range of the maritime Antarctic (60 to 72°S). Rather, we found the reverse pattern, with highest diversity at sites on Alexander Island (ca. 72°S), at the southern limit of the maritime Antarctic. The very limited overlap found between the eukaryotic biota of the different study sites, combined with their generally low relatedness to existing sequence databases, indicates a high level of Antarctic site isolation and possibly endemicity, a pattern not consistent with similar studies on other continents.     

Soil fungal community composition does not alter along a latitudinal gradient through the maritime and sub-Antarctic

We investigated the relationships between fungal community composition, latitude and a range of physicochemical parameters in 58 soils sampled from a 2370 km latitudinal gradient between South Georgia (54°S, 38°W) in the sub-Antarctic and Mars Oasis (72°S, 68°W) on Alexander Island in the southern maritime Antarctic. Our study, which is based on approximately ten times the number of samples used in previous similar studies, indicates that latitude and its associated environmental parameters are not related to fungal community composition. Significant changes in the composition of soil fungal communities were observed in relation to gradients of the ratio of total organic carbon to nitrogen, and, to a lesser extent, soil pH.     

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.     

Ciliate biogeography in Antarctic and Arctic freshwater ecosystems: endemism or global distribution of species?

Ciliate diversity was investigated in situ in freshwater ecosystems of the maritime (South Shetland Islands, mainly Livingston Island, 63 degrees S) and continental Antarctic (Victoria Land, 75 degrees S), and the High Arctic (Svalbard, 79 degrees N). In total, 334 species from 117 genera were identified in both polar regions, i.e. 210 spp. (98 genera) in the Arctic, 120 spp. (73 genera) in the maritime and 59 spp. (41 genera) in the continental Antarctic. Forty-four species (13% of all species) were common to both Arctic and Antarctic freshwater bodies and 19 spp. to both Antarctic areas (12% of all species). Many taxa are cosmopolitans but some, e.g. Stentor and Metopus spp., are not, and over 20% of the taxa found in any one of the three areas are new to science. Cluster analysis revealed that species similarity between different biotopes (soil, moss) within a study area was higher than between similar biotopes in different regions. Distinct differences in the species composition of freshwater and terrestrial communities indicate that most limnetic ciliates are not ubiquitously distributed. These observations and the low congruence in species composition between both polar areas, within Antarctica and between high- and temperate-latitude water bodies, respectively, suggest that long-distance dispersal of limnetic ciliates is restricted and that some species have a limited geographical distribution.     

The impact of different soil parameters on the community structure of dominant bacteria from nine different soils located on Livingston Island, South Shetland Archipelago, Antarctica

Microorganisms inhabit very different soil habitats in the ice-free areas of Antarctica, playing a major role in nutrient cycling in cold environments. We studied the soil characteristics and the dominant bacterial composition from nine different soil profiles located on Livingston Island (maritime Antarctica). The total carbon (TC) and total nitrogen (TN) values were high for the vegetated soils, decreasing with depth, whereas the values for the mineral soils were generally low. Soil pH was more acidic for moss-covered soils and neutral to alkaline for mineral soils. Numbers of culturable heterotrophic bacteria were higher at vegetated sites, but significant numbers were also detectable in carbon-depleted soils. Patterns of denaturing gradient gel electrophoresis (DGGE) revealed a highly heterogeneous picture throughout the soil profiles. Subsequent sequencing of DGGE bands revealed in total 252 sequences that could be assigned to 114 operational taxonomic units, showing the dominance of members of the Bacteroidetes and Acidobacteria. The results of phospholipid fatty acid analysis showed a lack of unsaturated fatty acids for most of the samples. Samples with a prevalence of unsaturated over saturated fatty acids were restricted to several surface samples. Statistical analysis showed that the dominant soil bacterial community composition is most affected by TC and TN contents and soil physical factors such as grain size and moisture, but not pH.     

Soil Microbial Abundance and Diversity Along a Low Precipitation Gradient

The exploration of spatial patterns of abundance and diversity patterns along precipitation gradients has focused for centuries on plants and animals; microbial profiles along such gradients are largely unknown. We studied the effects of soil pH, nutrient concentration, salinity, and water content on bacterial abundance and diversity in soils collected from Mediterranean, semi-arid, and arid sites receiving approximately 400, 300, and 100 mm annual precipitation, respectively. Bacterial diversity was evaluated by terminal restriction fragment length polymorphism and clone library analyses and the patterns obtained varied with the climatic regions. Over 75% of the sequenced clones were unique to their environment, while ∼2% were shared by all sites, yet, the Mediterranean and semi-arid sites had more common clones (∼9%) than either had with the arid site (4.7% and 6%, respectively). The microbial abundance, estimated by phospholipid fatty acids and real-time quantitative PCR assays, was significantly lower in the arid region. Our results indicate that although soil bacterial abundance decreases with precipitation, bacterial diversity is independent of precipitation gradient. Furthermore, community composition was found to be unique to each ecosystem.     

Precipitation nitrogen at maritime Signy Island and continental Cape Bird,Antarctica

Summary The mineral nitrogen (NH₄-N + NO₃-N) in precipitation occurring at continental and maritime Antarctic sites has been determined. Precipitation at sites remote from animal activity contained much less mineral N than that occurring at sites influenced by such activity. Estimates for nitrogen input at two contrasting fellfield sites, one at continental Cape Bird (dry site), the other at maritime Signy Island (wet site) are presented. At both sites precipitation N represented the major N input to fellfield biota than did wind blown particulate matter containing organic nitrogen or ammonia volatilized from adjacent guano soils and becoming absorbed by moist artificial soils traps.     

Enhanced sexual reproduction in bryophytes at high latitudes in the maritime Antarctic

Abstract

Contrary to the generally accepted hypothesis that bryophyte fertility decreases with increasing latitude and therefore climatic severity, a detailed study of bryophyte reproductive strategies at sites in the southern maritime Antarctic (68–72°S) has revealed that an unexpectedly high proportion of species is capable of producing sporophytes. Of the regional bryoflora, 43% (19 species; 17 mosses, 2 liverworts) in Marguerite Bay and 47% (17 species; 16 mosses, 1 liverwort) in Alexander Island are known to produce sporophytes, although the number fruiting at comparable latitudes on the colder and more arid Antarctic continent is less (33%). These numbers and proportions are much greater than were previously known at such relatively high southern polar latitudes. Sporophytes of several species are recorded for the first time within the Antarctic biome, while those of two liverworts (Lophozia excisa and Cephaloziella varians) are reported for the first time south of 62°S. High incidence of sporophyte production is attributed to locally favourable microclimatic conditions producing small-scale 'oases'. The large majority of fertile mosses are monoecious short acrocarps growing on rather calcareous soils. Spore production and size data for a number of species are compared with previously published studies of the same or closely related species from the northern maritime Antarctic and sub-Antarctic, but no consistent trends are revealed between species over the latitudinal gradient.     

Characterization of Bacterial,Archaeal and Eukaryote Symbionts from Antarctic Sponges Reveals a High Diversity at a Three-Domain Level and a Particular Signature for This Ecosystem

Sponge-associated microbial communities include members from the three domains of life. In the case of bacteria, they are diverse, host specific and different from the surrounding seawater. However, little is known about the diversity and specificity of Eukarya and Archaea living in association with marine sponges. This knowledge gap is even greater regarding sponges from regions other than temperate and tropical environments. In Antarctica, marine sponges are abundant and important members of the benthos, structuring the Antarctic marine ecosystem. In this study, we used high throughput ribosomal gene sequencing to investigate the three-domain diversity and community composition from eight different Antarctic sponges. Taxonomic identification reveals that they belong to families Acarnidae, Chalinidae, Hymedesmiidae, Hymeniacidonidae, Leucettidae, Microcionidae, and Myxillidae. Our study indicates that there are different diversity and similarity patterns between bacterial/archaeal and eukaryote microbial symbionts from these Antarctic marine sponges, indicating inherent differences in how organisms from different domains establish symbiotic relationships. In general, when considering diversity indices and number of phyla detected, sponge-associated communities are more diverse than the planktonic communities. We conclude that three-domain microbial communities from Antarctic sponges are different from surrounding planktonic communities, expanding previous observations for Bacteria and including the Antarctic environment. Furthermore, we reveal differences in the composition of the sponge associated bacterial assemblages between Antarctic and tropical-temperate environments and the presence of a highly complex microbial eukaryote community, suggesting a particular signature for Antarctic sponges, different to that reported from other ecosystems.     

The terrestrial arthropod fauna of the Byers Peninsula,Livingston Island,South Shetland Islands

We report the first detailed study of the terrestrial invertebrate fauna of the Byers Peninsula SSSI, Livingston Island, South Shetland Islands. Fourteen micro-arthropod taxa (10 Acari, four Collembola) and two Diptera are recorded, including the first record of the mite Edwardzetes dentifer from the maritime Antarctic. The first record of the midge Belgica antarctica from neighbouring Snow Island is also given. Population composition and density were described in samples from a wide range of terrestrial and freshwater habitats. There was no strong relationship between habitat and microarthropod species occurrence, although comparison of completely vegetated and more stony sites revealed greater population densities at the vegetated sites, and different species proportions at each. Some individual samples contained a wide range of species with none achieving numerical dominance, whilst others from superficially similar sites were dominated by one species. Dipterans were limited to a small number of lakes, streams and seepage areas, where they were sometimes abundant. Population density data and species occurrence are compared with previously published studies from the maritime Antarctic and elsewhere.     

Temperature sensitivity of Antarctic soil-humic substance degradation by cold-adapted bacteria

Heteropolymer humic substances (HS) are the largest constituents of soil organic matter and are key components that affect plant and microbial growth in maritime Antarctic tundra. We investigated HS decomposition in Antarctic tundra soils from distinct sites by incubating samples at 5°C or 8°C (within a natural soil thawing temperature range of −3.8°C to 9.6°C) for 90 days (average Antarctic summer period). This continuous 3-month artificial incubation maintained a higher total soil temperature than that in natural conditions. The long-term warming effects rapidly decreased HS content during the initial incubation, with no significant difference between 5°C and 8°C. In the presence of Antarctic tundra soil heterogeneity, the relative abundance of Proteobacteria (one of the major bacterial phyla in cold soil environments) increased during HS decomposition, which was more significant at 8°C than at 5°C. Contrasting this, the relative abundance of Actinobacteria (another major group) did not exhibit any significant variation. This microcosm study indicates that higher temperatures or prolonged thawing periods affect the relative abundance of cold-adapted bacterial communities, thereby promoting the rate of microbial HS decomposition. The resulting increase in HS-derived small metabolites will possibly accelerate warming-induced changes in the Antarctic tundra ecosystem.     

Response of pioneer soil microalgal colonists to environmental change in Antarctica

There is increasing evidence of climate change in Antarctica, especially elevated temperature and ultraviolet B (UVB) flux within the ozone hole. Its origins are debatable, but the effects on ice recession, water availability, and summer growth conditions are demonstrable. Light-dependent, temperature-sensitive, fast-growing organisms respond to these physical and biogeographical changes. Microalgae (cyanobacteria and eukaryotic algae), which are pioneer colonists of Antarctic mineral fellfield soils, are therefore highly suitable biological indicators of such changes. In frost-heaved soil polygons containing naturally sorted fine mineral particles, microalgal growth is restricted to a shallow zone of light penetration. By virtue of this light requirement, microalgae are exposed to extreme seasonal fluctuations in temperature (air and black-body radiation), photosynthetically active radiation, UV radiation, and desiccation. Dominance of conspicuous autofluorescent indicator species with distinctive morphology allowed quantification of responses using epifluorescence microscopy, and image analysis of undisturbed, unstained communities. However, the physical changes in climate, although significant in the long term, are gradual. The changes were therefore amplified experimentally by enclosing the communities at a fellfield site on Signy Island, maritime Antarctica, in cloches (small greenhouses). These were made of polystyrene of either UV transparent or UV opaque acrylic plastic, with or without walls. During a 6-year period, statistically significant changes were observed in microalgal colonization of the soil surface and in the morphology of filamentous populations. Evidence of community succession correlated with measured changes in local environment was found. Results from Signy Island and at continental sites on Alexander Island suggested that rates of microalgal colonization and community development might change significantly during current climate changes in Antarctica. Correspondence to: D.D. Wynn-Williams.     

Soil bacterial diversity in the Arctic is not fundamentally different from that found in other biomes

The severe environmental stresses of the Arctic may have promoted unique soil bacterial communities compared with those found in lower latitude environments. Here, we present a comprehensive analysis of the biogeography of soil bacterial communities in the Arctic using a high resolution bar‐coded pyrosequencing technique. We also compared arctic soils with soils from a wide range of more temperate biomes to characterize variability in soil bacterial communities across the globe. We show that arctic soil bacterial community composition and diversity are structured according to local variation in soil pH rather than geographical proximity to neighboring sites, suggesting that local environmental heterogeneity is far more important than dispersal limitation in determining community‐level differences. Furthermore, bacterial community composition had similar levels of variability, richness and phylogenetic diversity within arctic soils as across soils from a wide range of lower latitudes, strongly suggesting a common diversity structure within soil bacterial communities around the globe. These results contrast with the well‐established latitudinal gradients in animal and plant diversity, suggesting that the controls on bacterial community distributions are fundamentally different from those observed for macro‐organisms and that our biome definitions are not useful for predicting variability in soil bacterial communities across the globe.     

Bacterial diversity in the mountains of South-West China: Climate dominates over soil parameters

Certain patterns in soil bacterial diversity and community composition have become evident from metagenomics studies on a range of scales, from various parts of the world. For example, soil pH has generally been seen as dominating variation in bacterial diversity, above all other soil and climate parameters. It is important however to test the generality of these relationships by studying previously un-sampled areas. We compared soil bacterial diversity and community composition under a wide range of climatic and edaphic conditions in mountainous Yunnan Province, SW China. Soil samples were taken from a range of primary forest types and altitudes, reflecting the great variation of forest environments in this region. From each soil sample, DNA was extracted and pyrosequenced for bacterial 16S rRNA gene identification. In contrast to other recent studies from other parts of the world, pH was a weaker predictor of bacterial community composition and diversity than exchangeable Ca²⁺ concentration, and also the more poorly defined environmental parameter of elevation. Samples from within each forest type clustered strongly, showing the distinctive pattern of their microbial communities on a regional scale. It is clear that on a regional scale in a very heterogeneous environment, additional factors beyond pH can emerge as more important in determining bacterial diversity.     

Diversity of the cyanobacterial microflora of the northern part of James Ross Island,NW Weddell Sea,Antarctica

The diversity and ecological distribution of cyanobacteria in the northern, deglaciated part of James Ross Island were studied during the Antarctic summer season 2005–2006. Seventy-five cyanobacterial morphotypes were observed in various habitats of this area. The identified cyanobacterial taxa belong to the characteristic and dominant types of coastal Antarctica, and majority of them appeared connected to special habitats and formed distinct populations and ecologically delimited communities. The results are compared and discussed with respect to phenotypically characterised cyanobacterial microflora of maritime Antarctica and to recent molecular analyses of cyanobacterial strains from different Antarctic regions. The existence of a specificity and characteristic composition of Antarctic cyanobacterial communities was demonstrated.     

How Does Conversion of Natural Tropical Rainforest Ecosystems Affect Soil Bacterial and Fungal Communities in the Nile River Watershed of Uganda?

Uganda''s forests are globally important for their conservation values but are under pressure from increasing human population and consumption. In this study, we examine how conversion of natural forest affects soil bacterial and fungal communities. Comparisons in paired natural forest and human-converted sites among four locations indicated that natural forest soils consistently had higher pH, organic carbon, nitrogen, and calcium, although variation among sites was large. Despite these differences, no effect on the diversity of dominant taxa for either bacterial or fungal communities was detected, using polymerase chain reaction-denaturing gradient gel electrophoresis (PCR-DGGE). Composition of fungal communities did generally appear different in converted sites, but surprisingly, we did not observe a consistent pattern among sites. The spatial distribution of some taxa and community composition was associated with soil pH, organic carbon, phosphorus and sodium, suggesting that changes in soil communities were nuanced and require more robust metagenomic methods to understand the various components of the community. Given the close geographic proximity of the paired sampling sites, the similarity between natural and converted sites might be due to continued dispersal between treatments. Fungal communities showed greater environmental differentiation than bacterial communities, particularly according to soil pH. We detected biotic homogenization in converted ecosystems and substantial contribution of β-diversity to total diversity, indicating considerable geographic structure in soil biota in these forest communities. Overall, our results suggest that soil microbial communities are relatively resilient to forest conversion and despite a substantial and consistent change in the soil environment, the effects of conversion differed widely among sites. The substantial difference in soil chemistry, with generally lower nutrient quantity in converted sites, does bring into question, how long this resilience will last.     

Extracellular enzymes produced by microorganisms isolated from maritime Antarctica

Antarctic environments can sustain a great diversity of well-adapted microorganisms known as psychrophiles or psychrotrophs. The potential of these microorganisms as a resource of enzymes able to maintain their activity and stability at low temperature for technological applications has stimulated interest in exploration and isolation of microbes from this extreme environment. Enzymes produced by these organisms have a considerable potential for technological applications because they are known to have higher enzymatic activities at lower temperatures than their mesophilic and thermophilic counterparts. A total of 518 Antarctic microorganisms, were isolated during Antarctic expeditions organized by the Instituto Antártico Uruguayo. Samples of particules suspended in air, ice, sea and freshwater, soil, sediment, bird and marine animal faeces, dead animals, algae, plants, rocks and microbial mats were collected from different sites in maritime Antarctica. We report enzymatic activities present in 161 microorganisms (120 bacteria, 31 yeasts and 10 filamentous fungi) isolated from these locations. Enzymatic performance was evaluated at 4 and 20°C. Most of yeasts and bacteria grew better at 20°C than at 4°C, however the opposite was observed with the fungi. Amylase, lipase and protease activities were frequently found in bacterial strains. Yeasts and fungal isolates typically exhibited lipase, celullase and gelatinase activities. Bacterial isolates with highest enzymatic activities were identified by 16S rDNA sequence analysis as Pseudomonas spp., Psychrobacter sp., Arthrobacter spp., Bacillus sp. and Carnobacterium sp. Yeasts and fungal strains, with multiple enzymatic activities, belonged to Cryptococcus victoriae, Trichosporon pullulans and Geomyces pannorum.     

The Effect of Resource Islands on Abundance and Diversity of Bacteria in Arid Soils

Bacteria and nutrients were determined in upper soil samples collected underneath and between canopies of the dominant perennial in each of three sites along a steep precipitation gradient ranging from the Negev desert in the south of Israel to a Mediterranean forest in the north. Bacterial abundance, monitored by phospholipid fatty acid analysis, was significantly higher under the shrub canopy (compared to barren soils) in the arid and semi-arid sites but not in the Mediterranean soils. Bacterial community composition, determined using terminal restriction fragment length polymorphism and clone libraries, differed according to the sample’s origin. Closer examination revealed that in the arid and semi-arid sites, α-Proteobacteria are more abundant under the shrub canopy, while barren soils are characterized by a higher abundance of Actinobacteria. The bacterial communities in the Mediterranean soils were similar in both patch types. These results correspond to the hypothesis of “resource islands”, suggesting that shrub canopies provide a resource haven in low-resource landscapes. Yet, a survey of the physicochemical parameters of inter- and under-shrub soils could not attribute the changes in bacterial diversity to soil moisture, organic matter, or essential macronutrients. We suggest that in the nutrient-poor soils of the arid and semi-arid sites, bacteria occupying the soil under the shrub canopy may have longer growth periods under favorable conditions, resulting in their increased biomass and altered community composition.     

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.