T. G. Allan Green

The spatial structure of Antarctic biodiversity

Patterns of environmental spatial structure lie at the heart of the most fundamental and familiar patterns of diversity on Earth. Antarctica contains some of the strongest environmental gradients on the planet and therefore provides an ideal study ground to test hypotheses on the relevance of environmental variability for biodiversity. To answer the pivotal question, “How does spatial variation in physical and biological environmental properties across the Antarctic drive biodiversity?” we have synthesized current knowledge on environmental variability across terrestrial, freshwater, and marine Antarctic biomes and related this to the observed biotic patterns. The most important physical driver of Antarctic terrestrial communities is the availability of liquid water, itself driven by solar irradiance intensity. Patterns of biota distribution are further strongly influenced by the historical development of any given location or region, and by geographical barriers. In freshwater ecosystems, free water is also crucial, with further important influences from salinity, nutrient availability, oxygenation, and characteristics of ice cover and extent. In the marine biome there does not appear to be one major driving force, with the exception of the oceanographic boundary of the Polar Front. At smaller spatial scales, ice cover, ice scour, and salinity gradients are clearly important determinants of diversity at habitat and community level. Stochastic and extreme events remain an important driving force in all environments, particularly in the context of local extinction and colonization or recolonization, as well as that of temporal environmental variability. Our synthesis demonstrates that the Antarctic continent and surrounding oceans provide an ideal study ground to develop new biogeographical models, including life history and physiological traits, and to address questions regarding biological responses to environmental variability and change.

Spatial modelling of wetness for the Antarctic Dry Valleys

This paper describes a method used to model relative wetness for part of the Antarctic Dry Valleys using Geographic Information Systems (GIS) and remote sensing. The model produces a relative index of liquid water availability using variables that influence the volume and distribution of water. Remote sensing using Moderate Resolution Imaging Spectroradiometer (MODIS) images collected over four years is used to calculate an average index of snow cover and this is combined with other water sources such as glaciers and lakes. This water source model is then used to weight a hydrological flow accumulation model that uses slope derived from Light Detection and Ranging (LIDAR) elevation data. The resulting wetness index is validated using three-dimensional visualization and a comparison with a high-resolution Advanced Land Observing Satellite image that shows drainage channels. This research demonstrates that it is possible to produce a wetness model of Antarctica using data that are becoming widely available.

Are lichens active under snow in continental Antarctica

Photosynthetic activity, detected as chlorophyll a fluorescence, was measured for lichens under undisturbed snow in continental Antarctica using fibre optics. The fibre optics had been buried by winter snowfall after being put in place the previous year under snow-free conditions. The fibre optics were fixed in place using specially designed holding devices so that the fibre ends were in close proximity to selected lichens. Several temperature and PPFD (photosynthetic photon flux density) sensors were also installed in or close to the lichens. By attaching a chlorophyll a fluorometer to the previously placed fibre optics it proved possible to measure in vivo potential photosynthetic activity of continental Antarctic lichens under undisturbed snow. The snow cover proved to be a very good insulator for the mosses and lichens but, in contrast to the situation reported for the maritime Antarctic, it retained the severe cold of the winter and prevented early warming. Therefore, the lichens and mosses under snow were kept inactive at subzero temperatures for a prolonged time, even though the external ambient air temperatures would have allowed metabolic activity. The results suggest that the major activity period of the lichens was at the time of final disappearance of the snow and lasted about 10–14 days. The activation of lichens under snow by high air humidity appeared to be very variable and species specific. Xanthoria mawsonii was activated at temperatures below −10°C through absorption of water from high air humidity. Physcia dubia showed some activation at temperatures around –5°C but only became fully activated at thallus temperatures of 0°C through liquid water. Candelariella flava stayed inactive until thallus temperatures close to zero indicated that liquid water had become available. Although the snow cover represented the major water supply for the lichens, lichens only became active for a brief time at or close to the time the snow disappeared. The snow did not provide a protected environment, as reported for alpine habitats, but appeared to limit lichen activity. This provides at least one explanation for the observed negative effect of extended snow cover on lichen growth.

Ecology of endolithic lichens colonizing granite in continental Antarctica

In this study, the symbiont cells of several endolithic lichens colonizing granite in continental Antarctica and the relationships they have with the abiotic environment were analyzed in situ, in order to characterize the microecosystems integrating these lichens, from a microecological perspective. Mycobiont and photobiont cells, the majority classified as living by fluoresecent vitality testing, were observed distributed through the fissures of the granite. The fact that extracellular polymeric substances were commonly observed close to these cells and the features of these compounds, suggest a certain protective role for these substances against the harsh conditions of the environment. Different chemical, physical and biological relationships take place within the endo- lithic biofilms where the lichens are found, possibly affecting also the survival and distribution of these organisms. The alteration of bedrock minerals and synthesis of biominerals in the proximity of these lichens gives rise to different chemical microenvironments and suggests their participation in mineral nutrient cycling.

High thallus water content severely limits photosynthetic carbon gain of central European epilithic lichens under natural conditions

Experiments under controlled conditions have shown that net photosynthesis (NP) of many lichens is depressed when their thalli are highly hydrated. In this study we characterise the light and water content (WC) dependency of CO2 exchange for selected epilithic lichens in the laboratory and match this against samples monitored in their natural habitat by a novel, fully automatic cuvette. Laboratory measurements showed that, at a photosynthetic photon flux density (PPFD) of 1500 μmol m-2 s-1, NP of the epilithic foliose lichen Xanthoria calcicola was reduced by about 85% (compared to NP at optimal water content) when the thallus was suprasaturated (maximal hydration was defined as WC after spraying, submerging and subsequent removal of adhering water droplets by shaking). Only after loss of about 80% of its maximal WC were the highest rates of NP possible. This depression was still substantial at 50 μmol m-2 s-1 PPFD. Responses were similar for the crustose epilithic species Lecanora muralis. CO2 exchange of both lichens was monitored under natural conditions by means of the cuvette built into a man-made wall-a common habitat of the species-in the Botanical Garden, Würzburg. For both species, rates of NP were low during and after heavy rain even if incident PPFD and temperature were favourable. This situation occurred frequently and could last through all daylight hours, resulting in a negative carbon balance when nocturnal rates of respiration were high. Often, after rainfall, there was a brief, high peak of NP when optimal WC was transiently attained before metabolic activity finally ceased through desiccation. Other periods with profitable rates of NP occurred after moderate moistening of the lichens by dew, fog or light rain. The lichens were found to perform identically in the field and laboratory. When the two data sets were compared it was clear that the full range of WC produced in the laboratory also occurred in nature and that the productivity of the epilithic lichens was regularly and severely limited by high WC. It is concluded that blockage of diffusive pathways for CO2 in the thallus through high water contents is an important ecological factor for productivity of these central European epilithic lichens.

Ecophysiology of Desiccation/Rehydration Cycles in Mosses and Lichens

Although both lichens and bryophytes are all poikilohydric the groups seem to behave very differently. Bryophytes also show a clear preference for wetter areas and this seems to be a result of the different structures of the organisms. A lichen is algae (or cyanobacteria) suspended in a mycobiont with excess water often having a negative effect on photosynthesis. Bryophytes, in contrast, are true multicellular plants and can construct photosynthetic tissues that can effectively separate their photosynthetic and water storage functions. Under dry atmospheric conditions lichens and bryophytes will desiccate to low water contents and they become dormant. Ability to tolerate desiccation varies considerably both between and within the groups. Somewhat surprisingly, lichens appear to show less ability to tolerate long periods of desiccation than bryophytes, and even some vascular plants. Actual mechanisms of desiccation have been best studied in bryophytes and appear to be constitutive, no protein synthesis is required on rehydration to enable the commencement of metabolism and the necessary protection appears to be always present. Consistently high sucrose levels, for instance are reported from bryophytes. Cellular structure is often maintained when desiccated. Recovery from dryness also differs between the groups with bryophytes generally hydrating more slowly but there are large species differences. In general, rate of recovery may be related to the length of the hydrated activity period, species that hydrate and then dry rapidly, as on rock surfaces, recover rapidly. Species in habitats that remain wet for long periods once hydrated appear to recover more slowly from dryness. In addition to a photosynthetic response to light and temperature, the poikilohydric lichens and bryophytes also have a photosynthetic response to thallus water content. Starting with a dry thallus, addition of water will both increase the thallus water content and also allow photosynthesis and respiration to commence. Both processes increase almost linearly with further hydration at low water contents. Photosynthesis reaches a maximum at an optimal thallus water content (WCopt) that is strongly species dependant. In both groups this photosynthetic optimum represents full cellular turgor. At water contents above this optimum surface or external water can interfere with carbon dioxide uptake and can severely limit photosynthetic rates, especially in lichens. When thallus water contents are normalised to WCopt = 1, then the net photosynthesis (NP) response curves at water contents below WCopt are very similar for liverworts, mosses and higher plants, suggesting a common mechanism in controlling NP. It is suggested that this might be an inhibitor acting on Rubisco activity. In contrast to vascular plants both groups can carry out photosynthesis at lower, suboptimal thallus water contents and very low water potentials but the contribution that this makes to total carbon budget appears to be a major difference between the groups. Bryophytes seem to pass rapidly through this water content range when both drying and hydrating for tens of minutes are often enough. In contrast, it is now apparent that lichens are often active at low thallus water contents. They can not only hydrate from humid air alone, or from dew and fog, but can use these water sources very effectively, often achieving a major part of their annual carbon gain. Information on when the lichens and bryophytes are actually active is only recently starting to appear but, again, the groups seem to differ. Bryophytes strongly prefer wetter habitats and can be active and fully hydrated for long periods and seem to have excellent capacity to tolerate high light and UV radiation when wet. In contrast many lichens, in particular those with green algal symbionts, rarely seem to be hydrated for long periods, especially in high light conditions, and rapidly dry out. Lichens seem to be active mainly under suboptimal conditions one of which is suboptimal water content.

Photosynthetic responses of three common mosses from continental Antarctica

Predicting the effects of climate change on Antarctic terrestrial vegetation requires a better knowledge of the ecophysiology of common moss species. In this paper we provide a comprehensive matrix for photosynthesis and major environmental parameters for three dominant Antarctic moss species (Bryum subrotundifolium, B. pseudotriquetrum and Ceratodon purpureus). Using locations in southern Victoria Land, (Granite Harbour, 77°S) and northern Victoria Land (Cape Hallett, 72°S) we determined the responses of net photosynthesis and dark respiration to thallus water content, thallus temperature, photosynthetic photon flux densities and CO2 concentration over several summer seasons. The studies also included microclimate recordings at all sites where the research was carried out in field laboratories. Plant temperature was influenced predominantly by the water regime at the site with dry mosses being warmer. Optimal temperatures for net photosynthesis were 13.7°C, 12.0°C and 6.6°C for B. subrotundifolium, B. pseudotriquetrum and C. purpureus, respectively and fall within the known range for Antarctic mosses. Maximal net photosynthesis at 10°C ranked as B. subrotundifolium > B. pseudotriquetrum > C. purpureus. Net photosynthesis was strongly depressed at subzero temperatures but was substantial at 0°C. Net photosynthesis of the mosses was not saturated by light at optimal water content and thallus temperature. Response of net photosynthesis to increase in water content was as expected for mosses although B. subrotundifolium showed a large depression (60%) at the highest hydrations. Net photosynthesis of both B. subrotundifolium and B. pseudotriquetrum showed a large response to increase in CO2 concentration and this rose with increase in temperature; saturation was not reached for B. pseudotriquetrum at 20°C. There was a high level of variability for species at the same sites in different years and between different locations. This was substantial enough to make prediction of the effects of climate change very difficult at the moment.

Functional and spatial pressures on terrestrial vegetation in Antarctica forced by global warming

There is growing interest in what controls the present distribution of terrestrial vegetation in Antarctica because of the potential use of biodiversity as an indicator or predictor of the effects of climate change. Recent advances in knowledge of distribution and ecophysiological performance of terrestrial vegetation means that an initial analysis of the potential influence of temperature is now possible. Regressions of species numbers of lichens, mosses and hepatics on latitude and mean annual temperature (standard macroclimatic data) were carried out, and the terrestrial vegetation in Antarctica could be divided into two zones. The microenvironmental zone lies south of around 72°S, and biodiversity (richness and location) is uncoupled from the macroenvironment and is, instead, determined by the occasional coincidences of warmth, water, light and shelter. The macroenvironmental zone lies north of about 72°S, and biodiversity (richness, cover and growth) is strongly positively linked to mean annual temperature; species numbers increase at about 9–10% per K (24.0, 9.3 and 1.8 species for lichens, mosses and hepatics, respectively) probably due to improved water availability through increased precipitation and longer active period (monthly degree-days also reach zero at about 72°S) allowing greater productivity, completion of metabolic processes and a switch from survival to growth strategies. Cyanobacterial lichens appear to be a special case and may be expanding after being forced into northerly refugia. Warming will cause a southward movement of the boundary between the two zones but distribution in the microenvironmental zone will remain determined by local coincidences of environment and resources.

DEWFALL AS A WATER SOURCE FREQUENTLY ACTIVATES THE ENDOLITHIC CYANOBACTERIAL COMMUNITIES IN THE GRANITES OF TAYLOR VALLEY, ANTARCTICA1

Endolithic photosynthetic microorganisms like cyanobacteria and algae are well known from savannas and deserts of the world, the high Arctic, and also Antarctic habitats like the Dry Valleys in the Ross Dependency. These endolithic microbial communities are thought to be at the limits of life with reported ages in the order of thousands of years. Here we report on an extensive chasmoendolithic cyanobacterial community inside granite rocks of Mt. Falconer in the lower Taylor Valley, Dry Valleys. On average, the cyanobacterial community was 4.49 ± 0.95 mm below the rock surface, where it formed a blue‐green layer. The community was composed mainly of the cyanobacterium Chroococcidiopsis sp., with occasional Cyanothece cf. aeruginosa (Nägeli) Komárek and Nostoc sp. Mean biomass was 168 ± 44 g carbon · m−2, and the mean chl a content was 24.3 ± 34.2 mg · m−2. In situ chl fluorescence measurements—a relative measure of photosynthetic activity—showed that they were active over long periods each day and also showed activity the next day in the absence of any moisture. Radiocarbon dating gave a relatively young age (175–280 years) for the community. Calculations from microclimate data demonstrated that formation of dew or rime was possible and could frequently activate the cyanobacteria and may explain the younger age of microbial communities at Mt. Falconer compared to older and less active endolithic microorganisms reported earlier from Linnaeus Terrace, a higher altitude region that experiences colder, drier conditions.

Lichen myco- and photobiont diversity and their relationships at the edge of life (McMurdo Dry Valleys, Antarctica)

Lichen-forming fungi are among the most diverse group of organisms in Antarctica. Being poikilohydric, lichens are able to cope with harsh environmental conditions that exclude other organisms like vascular plants. The McMurdo Dry Valleys (Victoria Land, Continental Antarctica) are a hyperarid cold desert where macroscopic life is reduced to a few lichens and bryophyte species. We investigated the diversity of lichen-forming fungi and their associated photobionts in three valleys (Garwood, Marshall, and Miers). Correct identification of lichen-forming fungi from extreme ecosystems is complicated by the presence of numerous sterile and extremely modified thalli. To overcome this problem, we used a combined approach for the identification of the species present in the area, the first involving identification by means of standard characters and the second, two DNA-based (ITS region) species delimitation methods (General Mixed Yule-Coalescent model and genetic distances). In addition, we also used ITS sequences for the identification of the photobionts associated with the mycobionts. We studied the relationships between both bionts and assessed the degree of selectivity and specificity found in those associations. We also looked for landscape level spatial patterns in these associations. The two DNA-based methods performed quite differently, but 27 species of lichen-forming fungi and five putative species of photobionts were found in the studied area. Although there was a general trend for low selectivity in the relationships, high specificity was found in some associations and differential selectivity was observed in some lichen-forming fungi. No spatial structure was detected in the distribution of photobionts in the studied area.