A research agenda for nonvascular photoautotrophs under climate change.

Nonvascular photoautotrophs (NVP), including bryophytes, lichens, terrestrial algae, and cyanobacteria, are increasingly recognized as being essential to ecosystem functioning in many regions of the world. Current research suggests that climate change may pose a substantial threat to NVP, but the extent to which this will affect the associated ecosystem functions and services is highly uncertain. Here, we propose a research agenda to address this urgent question, focusing on physiological and ecological processes that link NVP to ecosystem functions while also taking into account the substantial taxonomic diversity across multiple ecosystem types. Accordingly, we developed a new categorization scheme, based on microclimatic gradients, which simplifies the high physiological and morphological diversity of NVP and world-wide distribution with respect to several broad habitat types. We found that habitat-specific ecosystem functions of NVP will likely be substantially affected by climate change, and more quantitative process understanding is required on: (1) potential for acclimation; (2) response to elevated CO2 ; (3) role of the microbiome; and (4) feedback to (micro)climate. We suggest an integrative approach of innovative, multimethod laboratory and field experiments and ecophysiological modelling, for which sustained scientific collaboration on NVP research will be essential.

Genetic diversity of soil invertebrates corroborates timing estimates for past collapses of the West Antarctic Ice Sheet.

During austral summer field seasons between 1999 and 2018, we sampled at 91 locations throughout southern Victoria Land and along the Transantarctic Mountains for six species of endemic microarthropods (Collembola), covering a latitudinal range from 76.0°S to 87.3°S. We assembled individual mitochondrial cytochrome c oxidase subunit 1 (COI) sequences (n = 866) and found high levels of sequence divergence at both small (<10 km) and large (>600 km) spatial scales for four of the six Collembola species. We applied molecular clock estimates and assessed genetic divergences relative to the timing of past glacial cycles, including collapses of the West Antarctic Ice Sheet (WAIS). We found that genetically distinct lineages within three species have likely been isolated for at least 5.54 My to 3.52 My, while the other three species diverged more recently (<2 My). We suggest that Collembola had greater dispersal opportunities under past warmer climates, via flotation along coastal margins. Similarly increased opportunities for dispersal may occur under contemporary climate warming scenarios, which could influence the genetic structure of extant populations. As Collembola are a living record of past landscape evolution within Antarctica, these findings provide biological evidence to support geological and glaciological estimates of historical WAIS dynamics over the last ca 5 My.

Assessing the conservation values and tourism threats in Barrientos Island, Antarctic Peninsula.

Antarctica has been witnessing continued growth of tourism, both in the overall visitation and in the diversity of itineraries and visitor activities. Expanding tourism presents unique business and educational opportunities, but it is also putting immense pressure on Antarctica's natural, and for the most parts, pristine environment. Understanding the effectiveness of different tourism management strategies and instruments, like the Visitor Site Guidelines adopted by the Antarctic Treaty, is fundamental to the sustainable management of Antarctic tourism. The purpose of this study was to assess the effectiveness of Visitor Site Guidelines and other tourism management actions in reducing impacts to the natural environment and for this, we used Barrientos Island as our case study as this is one of the most popular sites for tourism activities in the Antarctic Peninsula Region. First, we conducted a literature review and biological inventories to enable a thorough description of Barrientos Island's ecological values. The results show that Barrientos Island occupies the third highest biological richness among the top 15 most visited sites in the Antarctic Peninsula Region. We then assessed how tourism use on Barrientos Island affected biodiversity and the environment, and how Visitor Site Guidelines and other management measures helped alleviate these impacts. As intended, these instruments has been positive and valuable by providing operational guidance. However, they may lack significant information for tourism decision-making processes. To this end, we propose an alternative adaptive management approach that can more efficiently conserve biodiversity and environmental values while allowing the development of sustainable tourism activities in Antarctica.

High nitrogen contribution by Gunnera magellanica and nitrogen transfer by mycorrhizas drive an extraordinarily fast primary succession in sub-Antarctic Chile.

Chronosequences at the forefront of retreating glaciers provide information about colonization rates of bare surfaces. In the northern hemisphere, forest development can take centuries, with rates often limited by low nutrient availability. By contrast, in front of the retreating Pia Glacier (Tierra del Fuego, Chile), a Nothofagus forest is in place after only 34 yr of development, while total soil nitrogen (N) increased from near zero to 1.5%, suggesting a strong input of this nutrient. We measured N-fixation rates, carbon fluxes, leaf N and phosphorus contents and leaf δ15 N in the dominant plants, including the herb Gunnera magellanica, which is endosymbiotically associated with a cyanobacterium, in order to investigate the role of N-fixing and mycorrhizal symbionts in N-budgets during successional transition. G. magellanica presented some of the highest nitrogenase activities yet reported (potential maximal contribution of 300 kg N ha-1  yr-1 ). Foliar δ15 N results support the framework of a highly efficient N-uptake and transfer system based on mycorrhizas, with c. 80% of N taken up by the mycorrhizas potentially transferred to the host plant. Our results suggest the symbiosis of G. magellanica with cyanobacteria, and trees and shrubs with mycorrhizas, to be the key processes driving this rapid succession.

Desiccation Tolerance of Epiphytic Macrolichens in an Evergreen Temperate Rain Forest (Alerce Costero National Park, Chile).

The Valdivian region has a temperate rainy climate with differences in rainfall throughout the year. This heterogeneity results in periods of summer drought that expose the poikilohydric epiphytes to desiccation. With this research, we aim to answer different research questions related to phorophyte preference, response to desiccation, and response to radiation. How does the diversity of macrolichens vary at a local and microclimate scale in three tree species within an evergreen forest? What is the tolerance limit of macrolichens against prolonged desiccation, according to evaluation of the maximum efficiency of PSII (Fv/Fm) and pigment concentration? What is the tolerance limit against a potential increase in radiation? We found that macrolichen communities are determined by tree species, which regulate the suitability of the substrate by modifying the temperature and humidity conditions. In addition, our results show a rapid photosynthetic alteration in temporal exposure to desiccation, measured through Fv/Fm and pigment concentration. Our results showed that the most sensitive lichens to radiation and desiccation are not coincident. We confirm the low tolerance of macrolichen species to high radiation, reflected in the saturation profile obtained for the set studied. The lichen community in the evergreen forest showed high complexity and vulnerability, pointing to the importance of more research.

TerrANTALife 1.0 Biodiversity data checklist of known Antarctic terrestrial and freshwater life forms.

Background: Incomplete species inventories for Antarctica represent a key challenge for comprehensive ecological research and conservation in the region. Additionally, data required to understand population dynamics, rates of evolution, spatial ranges, functional traits, physiological tolerances and species interactions, all of which are fundamental to disentangle the different functional elements of Antarctic biodiversity, are mostly missing. However, much of the fauna, flora and microbiota in the emerged ice-free land of the continent have an uncertain presence and/or unresolved status, with entire biodiversity compendia of prokaryotic groups (e.g. bacteria) being missing. All the available biodiversity information requires consolidation, cross-validation, re-assessment and steady systematic inclusion in order to create a robust catalogue of biodiversity for the continent. New information: We compiled, completed and revised eukaryotic species inventories present in terrestrial and freshwater ecosystems in Antarctica in a new living database: terrANTALife (version 1.0). The database includes the first integration in a compendium for many groups of eukaryotic microorganisms. We also introduce a first catalogue of amplicon sequence variants (ASVs) of prokaryotic biodiversity. Available compendia and literature to date were searched for Antarctic terrestrial and freshwater species, integrated, taxonomically harmonised and curated by experts to create comprehensive checklists of Antarctic organisms. The final inventories comprises 470 animal species (including vertebrates, free-living invertebrates and parasites), 306 plants (including all Viridiplantae: embryophytes and green algae), 997 fungal species and 434 protists (sensu lato). We also provide a first account for many groups of microorganisms, including non-lichenised fungi and multiple groups of eukaryotic unicellular species (Stramenophila, Alveolata and Rhizaria (SAR), Chromists and Amoeba), jointly referred to as "protists". In addition, we identify 1753 bacterial (obtained from 348117 ASVs) and 34 archaeal genera (from 1848 ASVs), as well as, at least, 14 virus families. We formulate a basic tree of life in Antarctica with the main lineages listed in the region and their "known-accepted-species" numbers.

Macrolichen Communities Depend on Phorophyte in Conguillío National Park, Chile.

The community composition of epiphytic macrolichens from two tree species (Araucaria araucana and Nothofagus antarctica) was conducted in temperate forests in the Conguillío National Park, Chile. The composition of lichen biota is influenced by phorophyte species, bark pH, and microclimatic conditions. A total of 31 species of macrolichens were found on A. araucana and N. antarctica. Most of the species showed phorophyte preference, with nine being exclusive to A. araucana and 10 to N. antarctica. The detrended correspondence analysis (DCA) indicated the formation of three communities: one representing macrolichens growing on N. antarctica and two others growing chiefly on A. araucana, either with north or south exposure. More work is needed to study the lichen biota of the forests of the Chilean Andes, which are under multiple threats, including clearing and climate change. In order to counteract such risks to native forests and to the biodiversity of the associated epiphytic lichens, conservation plans should be established that consider the factors that influence the composition of the lichen community.

Floristic Similarities between the Lichen Flora of Both Sides of the Drake Passage: A Biogeographical Approach.

This paper analyses the lichen flora of Navarino Island (Tierra del Fuego, Cape Horn Region, Chile), identifying species shared with the South Shetland Islands (Antarctic Peninsula). In this common flora, species are grouped by their biogeographic origin (Antarctic-subantarctic endemic, austral, bipolar, and cosmopolitan), their habitat on Navarino Island (coastal, forest, and alpine), their morphotype (crustaceous, foliaceous, fruticulose, and cladonioid), and the substrate from which they were collected (epiphytic, terricolous and humicolous, and saxicolous). A total of 124 species have been recognised as common on both sides of the Drake Passage, predominantly bipolar, crustaceous, and saxicolous species, and with an alpine distribution on Navarino Island. The most interesting fact is that more than 30% of the flora is shared between the southern tip of South America and the western Antarctic Peninsula, which is an indication of the existence of a meridian flow of propagules capable of crossing the Antarctic polar front.

Physiological Plasticity as a Strategy to Cope with Harsh Climatic Conditions: Ecophysiological Meta-Analysis of the Cosmopolitan Moss Ceratodon purpureus in the Southern Hemisphere.

Determining the physiological tolerance ranges of species is necessary to comprehend the limits of their responsiveness under strong abiotic pressures. For this purpose, the cosmopolitan moss Ceratodon purpureus (Hedw.) Brid. is a good model due to its wide geographical distribution throughout different biomes and habitats. In order to disentangle how this species copes with stresses such as extreme temperatures and high radiation, we designed a meta-analysis by including the main photosynthetic traits obtained by gas exchange measurements in three contrasting habitats from the Southern Hemisphere. Our findings highlight that traits such as respiration homeostasis, modulation of the photosynthetic efficiency, adjustment of the optimal temperature, and switching between shade and sun-adapted forms, which are crucial in determining the responsiveness of this species. In fact, these ecophysiological traits are in concordance with the climatic particularities of each habitat. Furthermore, the photosynthetic trends found in our study point out how different Livingston Island (Maritime Antarctica) and Granite Harbour (Continental Antarctica) are for plant life, while the population from the Succulent Karoo Desert (South Africa) shares traits with both Antarctic regions. Altogether, the study highlights the high resilience of C. purpureus under abrupt climate changes and opens new perspectives about the wide spectrum of physiological responses of cryptogams to cope with climate change scenarios.

Continuous monitoring of chlorophyll a fluorescence and microclimatic conditions reveals warming-induced physiological damage in biocrust-forming lichens.

Purpose: Biocrust communities, which are important regulators of multiple ecosystem functions in drylands, are highly sensitive to climate change. There is growing evidence of the negative impacts of warming on the performance of biocrust constituents like lichens in the field. Here, we aim to understand the physiological basis behind this pattern. Methods: Using a unique manipulative climate change experiment, we monitored every 30 minutes and for 9 months the chlorophyll a fluorescence and microclimatic conditions (lichen surface temperature, relative moisture and photosynthetically active radiation) of Psora decipiens, a key biocrust constituent in drylands worldwide. This long-term monitoring resulted in 11,847 records at the thallus-level, which allowed us to evaluate the impacts of ~2.3 °C simulated warming treatment on the physiology of Psora at an unprecedented level of detail. Results: Simulated warming and the associated decrease in relative moisture promoted by this treatment negatively impacted the physiology of Psora, especially during the diurnal period of the spring, when conditions are warmer and drier. These impacts were driven by a mechanism based on the reduction of the length of the periods allowing net photosynthesis, and by declines in Yield and Fv/Fm under simulated warming. Conclusion: Our study reveals the physiological basis explaining observed negative impacts of ongoing global warming on biocrust-forming lichens in the field. The functional response observed could limit the growth and cover of biocrust-forming lichens in drylands in the long-term, negatively impacting in key soil attributes such as biogeochemical cycles, water balance, biological activity and ability of controlling erosion. Supplementary Information: The online version contains supplementary material available at 10.1007/s11104-022-05686-w.

Freeze Substitution Accelerated via Agitation: New Prospects for Ultrastructural Studies of Lichen Symbionts and Their Extracellular Matrix.

(1) Background: Lichens, as an important part of the terrestrial ecosystem, attract the attention of various research disciplines. To elucidate their ultrastructure, transmission electron microscopy of resin-embedded samples is indispensable. Since most observations of lichen samples are generated via chemical fixation and processing at room temperature, they lack the rapid immobilization of live processes and are prone to preparation artefacts. To improve their preservation, cryoprocessing was tested in the past, but never widely implemented, not least because of an extremely lengthy protocol. (2) Methods: Here, we introduce an accelerated automated freeze substitution protocol with continuous agitation. Using the example of three lichen species, we demonstrate the preservation of the native state of algal photobionts and mycobionts in association with their extracellular matrix. (3) Results: We bring to attention the extent and the structural variability of the hyphae, the extracellular matrix and numerous crystallized metabolites. Our findings will encourage studies on transformation processes related to the compartmentation of lichen thalli. They include cryopreserved aspects of algal photobionts and observations of putative physiological relevance, such as the arrangement of numerous mitochondria within chloroplast pockets. (4) Conclusions: In summary, we present accelerated freeze substitution as a very useful tool for systematic studies of lichen ultrastructures.

Extreme phenotypic variation in Cetraria aculeata (lichenized Ascomycota): adaptation or incidental modification?

BACKGROUND AND AIMS: Phenotypic variability is a successful strategy in lichens for colonizing different habitats. Vagrancy has been reported as a specific adaptation for lichens living in steppe habitats around the world. Among the facultatively vagrant species, the cosmopolitan Cetraria aculeata apparently forms extremely modified vagrant thalli in steppe habitats of Central Spain. The aim of this study was to investigate whether these changes are phenotypic plasticity (a single genotype producing different phenotypes), by characterizing the anatomical and ultrastructural changes observed in vagrant morphs, and measuring differences in ecophysiological performance. METHODS: Specimens of vagrant and attached populations of C. aculeata were collected on the steppes of Central Spain. The fungal internal transcribed spacer (ITS), glyceraldehyde-3-phosphate dehydrogenase (GPD) and the large sub-unit of the mitochondrial ribosomal DNA (mtLSUm), and the algal ITS and actin were studied within a population genetics framework. Semi-thin and ultrathin sections were analysed by means of optical, scanning electron and transmission electron microscopy. Gas exchange and chlorophyll fluorescence were used to compare the physiological performance of both morphs. KEY RESULTS AND CONCLUSIONS: Vagrant and attached morphs share multilocus haplotypes which may indicate that they belong to the same species in spite of their completely different anatomy. However, differentiation tests suggested that vagrant specimens do not represent a random sub-set of the surrounding population. The morphological differences were related to anatomical and ultrastructural differences. Large intercalary growth rates of thalli after the loss of the basal-apical thallus polarity may be the cause of the increased growth shown by vagrant specimens. The anatomical and morphological changes lead to greater duration of ecophysiological activity in vagrant specimens. Although the anatomical and physiological changes could be chance effects, the genetic differentiation between vagrant and attached sub-populations and the higher biomass of the former show fitness effects and adaptation to dry environmental conditions in steppe habitats.

Climate change leads to higher NPP at the end of the century in the Antarctic Tundra: Response patterns through the lens of lichens.

Poikilohydric autotrophs are the main colonizers of the permanent ice-free areas in the Antarctic tundra biome. Global climate warming and the small human footprint in this ecosystem make it especially vulnerable to abrupt changes. Elucidating the effects of climate change on the Antarctic ecosystem is challenging because it mainly comprises poikilohydric species, which are greatly influenced by microtopographic factors. In the present study, we investigated the potential effects of climate change on the metabolic activity and net primary photosynthesis (NPP) in the widespread lichen species Usnea aurantiaco-atra. Long-term monitoring of chlorophyll a fluorescence in the field was combined with photosynthetic performance measurements in laboratory experiments in order to establish the daily response patterns under biotic and abiotic factors at micro- and macro-scales. Our findings suggest that macroclimate is a poor predictor of NPP, thereby indicating that microclimate is the main driver due to the strong effects of microtopographic factors on cryptogams. Metabolic activity is also crucial for estimating the NPP, which is highly dependent on the type, distribution, and duration of the hydration sources available throughout the year. Under RCP 4.5 and RCP 8.5, metabolic activity will increase slightly compared with that at present due to the increased precipitation events predicted in MIROC5. Temperature is highlighted as the main driver for NPP projections, and thus climate warming will lead to an average increase in NPP of 167-171% at the end of the century. However, small changes in other drivers such as light and relative humidity may strongly modify the metabolic activity patterns of poikilohydric autotrophs, and thus their NPP. Species with similar physiological response ranges to the species investigated in the present study are expected to behave in a similar manner provided that liquid water is available.

Macroclimatic conditions as main drivers for symbiotic association patterns in lecideoid lichens along the Transantarctic Mountains, Ross Sea region, Antarctica.

Lecideoid lichens as dominant vegetation-forming organisms in the climatically harsh areas of the southern part of continental Antarctica show clear preferences in relation to environmental conditions (i.e. macroclimate). 306 lichen samples were included in the study, collected along the Ross Sea coast (78°S-85.5°S) at six climatically different sites. The species compositions as well as the associations of their two dominant symbiotic partners (myco- and photobiont) were set in context with environmental conditions along the latitudinal gradient. Diversity values were nonlinear with respect to latitude, with the highest alpha diversity in the milder areas of the McMurdo Dry Valleys (78°S) and the most southern areas (Durham Point, 85.5°S; Garden Spur, 84.5°S), and lowest in the especially arid and cold Darwin Area (~ 79.8°S). Furthermore, the specificity of mycobiont species towards their photobionts decreased under more severe climate conditions. The generalist lichen species Lecanora fuscobrunnea and Lecidea cancriformis were present in almost all habitats, but were dominant in climatically extreme areas. Carbonea vorticosa, Lecidella greenii and Rhizoplaca macleanii were confined to milder areas. In summary, the macroclimate is considered to be the main driver of species distribution, making certain species useful as bioindicators of climate conditions and, consequently, for assessing the consequences of climate change.

Lichen Vitality After a Space Flight on Board the EXPOSE-R2 Facility Outside the International Space Station: Results of the Biology and Mars Experiment.

As part of the Biology and Mars Experiment (BIOMEX; ILSRA 2009-0834), samples of the lichen Circinaria gyrosa were placed on the exposure platform EXPOSE-R2, on the International Space Station (ISS) and exposed to space and to a Mars-simulated environment for 18 months (2014-2016) to study: (1) resistance to space and Mars-like conditions and (2) biomarkers for use in future space missions (Exo-Mars). When the experiment returned (June 2016), initial analysis showed rapid recovery of photosystem II activity in the samples exposed exclusively to space vacuum and a Mars-like atmosphere. Significantly reduced recovery levels were observed in Sun-exposed samples, and electron and fluorescence microscopy (transmission electron microscope and field emission scanning electron microscope) data indicated that this was attributable to the combined effects of space radiation and space vacuum, as unirradiated samples exhibited less marked morphological changes compared with Sun-exposed samples. Polymerase chain reaction analyses confirmed that there was DNA damage in lichen exposed to harsh space and Mars-like environmental conditions, with ultraviolet radiation combined with space vacuum causing the most damage. These findings contribute to the characterization of space- and Mars-resistant organisms that are relevant to Mars habitability.

A multi-proxy approach to Late Holocene fluctuations of Tungnahryggsjökull glaciers in the Tröllaskagi peninsula (northern Iceland).

The Tröllaskagi Peninsula in northern Iceland hosts more than a hundred small glaciers that have left a rich terrestrial record of Holocene climatic fluctuations in their forelands. Traditionally, it has been assumed that most of the Tröllaskagi glaciers reached their Late Holocene maximum extent during the Little Ice Age (LIA). However, there is evidence of slightly more advanced pre-LIA positions. LIA moraines from Iceland have been primary dated mostly through lichenometric dating, but the limitations of this technique do not allow dating of glacial advances prior to the 18th or 19th centuries. The application of 36Cl Cosmic-Ray Exposure (CRE) dating to Tungnahryggsjökull moraine sequences in Vesturdalur and Austurdalur (central Tröllaskagi) has revealed a number of pre-LIA glacial advances at ~400 and ~700 CE, and a number of LIA advances in the 15th and 17th centuries, the earliest LIA advances dated so far in Tröllaskagi. This technique hence shows that the LIA chronology in Tröllaskagi agrees with that of other European areas such as the Alps or the Mediterranean mountains. The combined use of lichenometric dating, aerial photographs, satellite images and fieldwork shows that the regional colonization lag of the commonly used lichen species Rhizocarpon geographicum is longer than previously assumed. For exploratory purposes, an alternative lichen species (Porpidia soredizodes) has been tested for lichenometric dating, estimating a tentative growth rate of 0.737 mm yr-1.

Identity of plant, lichen and moss species connects with microbial abundance and soil functioning in Maritime Antarctica.

BACKGROUND AND AIMS: We lack studies evaluating how the identity of plant, lichen and moss species relates to microbial abundance and soil functioning on Antarctica. If species identity is associated with soil functioning, distributional changes of key species, linked to climate change, could significantly affect Antarctic soil functioning. METHODS: We evaluated how the identity of six Antarctic plant, lichen and moss species relates to a range of soil attributes (C, N and P cycling), microbial abundance and structure in Livingston Island, Maritime Antarctica. We used an effect size metric to predict the association between species (vs. bare soil) and the measured soil attributes. RESULTS: We observed species-specific effects of the plant and biocrust species on soil attributes and microbial abundance. Phenols, phosphatase and ß-D-cellobiosidase activities were the most important attributes characterizing the observed patterns. We found that the evaluated species positively correlated with soil nutrient availability and microbial abundance vs. bare soil. CONCLUSIONS: We provide evidence, from a comparative study, that plant and biocrust identity is associated with different levels of soil functioning and microbial abundance in Maritime Antarctica. Our results suggest that changes in the spatial distribution of these species linked to climate change could potentially entail changes in the functioning of Antarctic terrestrial ecosystems.

Cellular Responses of the Lichen Circinaria gyrosa in Mars-Like Conditions.

Lichens are extremely resistant organisms that colonize harsh climatic areas, some of them defined as "Mars-analog sites." There still remain many unsolved questions as to how lichens survive under such extreme conditions. Several studies have been performed to test the resistance of various lichen species under space and in simulated Mars-like conditions. The results led to the proposal that Circinaria gyrosa (Lecanoromycetes, Ascomycota) is one of the most durable astrobiological model lichens. However, although C. gyrosa has been exposed to Mars-like environmental conditions while in a latent state, it has not been exposed in its physiologically active mode. We hypothesize that the astrobiological test system "Circinaria gyrosa," could be able to be physiologically active and to survive under Mars-like conditions in a simulation chamber, based on previous studies performed at dessicated-dormant stage under simulated Mars-like conditions, that showed a complete recover of the PSII activity (Sánchez et al., 2012). Epifluorescence and confocal laser scanning microscopy (CLSM) showed that living algal cells were more abundant in samples exposed to niche conditions, which simulated the conditions in micro-fissures and micro-caves close to the surface that have limited scattered or time-dependent light exposure, than in samples exposed to full UV radiation. The medulla was not structurally affected, suggesting that the niche exposure conditions did not disturb the lichen thalli structure and morphology as revealed by field emission scanning electron microscopy (FESEM). In addition, changes in the lichen thalli chemical composition were determined by analytical pyrolysis. The chromatograms resulting from analytical pyrolysis at 500°C revealed that lichen samples exposed to niche conditions and full UV radiation consisted primarily of glycosidic compounds, lipids, and sterols, which are typical constituents of the cell walls. However, specific differences could be detected and used as markers of the UV-induced damage to the lichen membranes. Based on its viability responses after rehydration, our study shows that the test lichen survived the 30-day incubation in the Mars chamber particularly under niche conditions. However, the photobiont was not able to photosynthesize under the Mars-like conditions, which indicates that the surface of Mars is not a habitable place for C. gyrosa.

Ultrastructural and genetic characteristics of endolithic cyanobacterial biofilms colonizing Antarctic granite rocks.

The precise identification of the cyanobacteria that comprise an endolithic biofilm is hindered by difficulties in culturing the organisms found in these biofilms and a lack of previous molecular and ultrastructural data. This study characterizes, both at the ultrastructural and molecular level, two different cyanobacterial biofilms found in fissures of granite from continental Antarctica. Electron microscopy revealed structural differences between the two biofilms. One was only loosely adhered to the substrate, while the other biofilm showed a closer association between cells and rock minerals and was tightly attached to the substrate. Cells from both biofilms where ultrastructurally distinct, displaying, for instance, clear differences in their sheaths. The amounts of EPS and their organization associated with the cyanobacteria may determine the differences in adhesion and effects on the lithic substrate observed in the biofilms. By sequencing part of the 16S rRNA gene, the two cyanobacteria were also genetically characterized. The gene sequence of the cells comprising the biofilm that was tightly attached to the lithic substrate showed most homology with that of an endolithic cyanobacterium from Switzerland (AY153458), and the cyanobacterial type loosely adhered to the rock, clustered with Acaryochloris marina, the only organism unequivocally known to contain chlorophyll d. This study reveals the presence of at least two different types of endolithic biofilm, dominated each by a single type of cyanobacterium, able to withstand the harsh conditions of the Antarctic climate.

Lichens survive in space: results from the 2005 LICHENS experiment.

This experiment was aimed at establishing, for the first time, the survival capability of lichens exposed to space conditions. In particular, the damaging effect of various wavelengths of extraterrestrial solar UV radiation was studied. The lichens used were the bipolar species Rhizocarpon geographicum and Xanthoria elegans, which were collected above 2000 m in the mountains of central Spain and as endolithic communities inhabiting granites in the Antarctic Dry Valleys. Lichens were exposed to space in the BIOPAN-5 facility of the European Space Agency; BIOPAN-5 is located on the outer shell of the Earth-orbiting FOTON-M2 Russian satellite. The lichen samples were launched from Baikonur by a Soyuz rocket on May 31, 2005, and were returned to Earth after 16 days in space, at which time they were tested for survival. Chlorophyll fluorescence was used for the measurement of photosynthetic parameters. Scanning electron microscopy in back-scattered mode, low temperature scanning electron microscopy, and transmission electron microscopy were used to study the organization and composition of both symbionts. Confocal laser scanning microscopy, in combination with the use of specific fluorescent probes, allowed for the assessment of the physiological state of the cells. All exposed lichens, regardless of the optical filters used, showed nearly the same photosynthetic activity after the flight as measured before the flight. Likewise, the multimicroscopy approach revealed no detectable ultrastructural changes in most of the algal and fungal cells of the lichen thalli, though a greater proportion of cells in the flight samples had compromised membranes, as revealed by the LIVE/DEAD BacLight Bacterial Viability Kit. These findings indicate that most lichenized fungal and algal cells can survive in space after full exposure to massive UV and cosmic radiation, conditions proven to be lethal to bacteria and other microorganisms. The lichen upper cortex seems to provide adequate protection against solar radiation. Moreover, after extreme dehydration induced by high vacuum, the lichens proved to be able to recover, in full, their metabolic activity within 24 hours.