Search

Biocrusts represent associations of lichens, green algae, cyanobacteria, fungi and other microorganisms, colonizing soils in varying proportions of principally arid biomes. The so-called grit crust represents a recently discovered type of biocrust situated in the Coastal Range of the Atacama Desert (Chile) made of microorganisms growing on and in granitoid pebbles, resulting in a checkerboard pattern visible to the naked eye on the landscape scale. This specific microbiome fulfills a broad range of ecosystem services, all probably driven by fog and dew-induced photosynthetic activity of mainly micro-lichens. To understand its biodiversity and impact, we applied a polyphasic approach on the phototrophic microbiome of this biocrust, combining isolation and characterization of the lichen photobionts, multi-gene phylogeny of the photobionts and mycobionts based on a direct sequencing and microphotography approach, metabarcoding and determination of chlorophyll a+b contents. Metabarcoding showed that yet undescribed lichens within the Caliciaceae dominated the biocrust together with Trebouxia as the most abundant eukaryote in all plots. Together with high mean chlorophyll a+b contents exceeding 410 mg m -2 , this distinguished the symbiotic algae Trebouxia as the main driver of the grit crust ecosystem. The trebouxioid photobionts could be assigned to the I (T. impressa/gelatinosa) and A (T. arboricola) clades and represented several lineages containing five potential species candidates, which were identified based on the unique phylogenetic position, morphological features, and developmental cycles of the corresponding isolates. These results designate the grit crust as the only known coherent soil layer with significant landscape covering impact of at least 440 km 2 , predominantly ruled by a single symbiotic algal genus., (© 2024. The Author(s).)

Lichens are remarkable and classic examples of symbiotic organisms that have fascinated scientists for centuries. Yet, it has only been for a couple of decades that significant advances have focused on the diversity of their green algal and/or cyanobacterial photobionts. Cyanolichens, which contain cyanobacteria as their photosynthetic partner, include up to 10% of all known lichens and, as such, studies on their cyanobionts are much rarer compared to their green algal counterparts. For the unicellular cyanobionts, i.e. cyanobacteria that do not form filaments, these studies are even scarcer. Nonetheless, these currently include at least 10 different genera in the cosmopolitan lichen order Lichinales. An international consortium (International Network of CyanoBionts; INCb) will tackle this lack of knowledge. In this article, we discuss the status of current unicellular cyanobiont research, compare the taxonomic resolution of photobionts from cyanolichens with those of green algal lichens (chlorolichens), and give a roadmap of research on how to recondition the underestimated fraction of symbiotic unicellular cyanobacteria in lichens., Competing Interests: None declared., (© The Author(s) 2024. Published by Oxford University Press on behalf of the International Society for Microbial Ecology.)

Over the decades our understanding of lichens has shifted to the fact that they are multiorganismic, symbiotic microecosystems, with their complex interactions coming to the fore due to recent advances in microbiomics. Here, we present a mutualistic-parasitic continuum dynamics scenario between an orange lichen and a lichenicolous fungus from the Atacama Desert leading to the decay of the lichen's photobiont and leaving behind a black lichen thallus. Based on isolation, sequencing, and ecophysiological approaches including metabolic screenings of the symbionts, we depict consequences upon infection with the lichenicolous fungus. This spans from a loss of the lichen's photosynthetic activity and an increased roughness of its surface to an inhibition of the parietin synthesis as a shared pathway between the photobiont and the mycobiont, including a shift of secondary metabolism products. This degree of relations has rarely been documented before, although lichenicolous fungi have been studied for over 200 years, adding an additional level to the view of interactions within lichens.

Deserts represent an extreme challenge for photosynthetic life. Despite their aridity, they are often inhabited by diverse microscopic communities of cyanobacteria. These organisms are commonly found in lithic habitats, where they are partially sheltered from extremes of temperature and UV radiation. However, living under the rock surface imposes additional constraints, such as limited light availability, and enrichment of longer wavelengths than are typically usable for oxygenic photosynthesis. Some cyanobacteria from the genus Chroococcidiopsis can use this light to photosynthesize, in a process known as far-red light photoacclimation, or FaRLiP. This genus has commonly been reported from both hot and cold deserts. However, not all Chroococcidiopsis strains carry FaRLiP genes, thus motivating our study into the interplay between FaRLiP and extreme lithic environments. The abundance of sequence data and strains provided the necessary material for an in-depth phylogenetic study, involving spectroscopy, microscopy, and determination of pigment composition, as well as gene and genome analyses. Pigment analyses revealed the presence of red-shifted chlorophylls d and f in all FaRLiP strains tested. In addition, eight genus-level taxa were defined within the encompassing Chroococcidiopsidales, clarifying the phylogeny of this long-standing polyphyletic order. FaRLiP is near universally present in a generalist genus identified in a wide variety of environments, Chroococcidiopsis sensu stricto, while it is rare or absent in closely related, extremophile taxa, including those preferentially inhabiting deserts. This likely reflects the evolutionary process of gene loss in specialist lineages., (© 2023. ISME Publications B.V.)

Under continuous human disturbance, regeneration is the basis for biodiversity persistence and ecosystem service provision. In tropical dry forests, edaphic ecosystem engineering by biological soil crusts (biocrusts) could impact regeneration by influencing erosion control and soil water and nutrient fluxes, which impact landscape hydrology, geomorphology, and ecosystem functioning. This study investigated the effect of cyanobacteria-dominated biocrusts on water infiltration and aggregate stability in a human-modified landscape of the Caatinga dry forest (NE Brazil), a system characterized by high levels of forest degradation and increasing aridity. By trapping dust and swelling of cyanobacterial filaments, biocrusts can seal soil surfaces and slow down infiltration, which potentially induces erosion. To quantify hydraulic properties and erosion control, we used minidisc-infiltrometry, raindrop-simulation, and wet sieving at two sites with contrasting disturbance levels: an active cashew plantation and an abandoned field experiencing forest regeneration, both characterized by sandy soils. Under disturbance, biocrusts had a stronger negative impact on infiltration (reduction by 42% vs. 37% during regeneration), although biocrusts under regenerating conditions had the lowest absolute sorptivity (0.042 ± 0.02 cm s -1/2 ) and unsaturated hydraulic conductivity (0.0015 ± 0.0008 cm s -1 ), with a doubled water repellency. Biocrusts provided high soil aggregate stability although stability increased considerably with progression of biocrust succession (raindrop simulation disturbed: 0.19 ± 0.22 J vs. regenerating: 0.54 ± 0.22 J). The formation of stable aggregates by early successional biocrusts on sandy soils suggests protection of dry forest soils even on the worst land use/soil degradation scenario with a high soil erosion risk. Our results confirm that biocrusts covering bare interspaces between vascular plants in human-modified landscapes play an important role in surface water availability and erosion control. Biocrusts have the potential to reduce land degradation, but their associated ecosystem services like erosion protection, can be impaired by disturbance. Considering an average biocrust coverage of 8.1% of the Caatinga landscapes, further research should aim to quantify the contribution of biocrusts to forest recovery to fully understand the role they play in the functioning of this poorly explored ecosystem., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2023 Szyja, Felde, Lückel, Tabarelli, Leal, Büdel and Wirth.)

Studies of biological soil crusts (biocrusts) have proliferated over the last few decades. The biocrust literature has broadened, with more studies assessing and describing the function of a variety of biocrust communities in a broad range of biomes and habitats and across a large spectrum of disciplines, and also by the incorporation of biocrusts into global perspectives and biogeochemical models. As the number of biocrust researchers increases, along with the scope of soil communities defined as 'biocrust', it is worth asking whether we all share a clear, universal, and fully articulated definition of what constitutes a biocrust. In this review, we synthesize the literature with the views of new and experienced biocrust researchers, to provide a refined and fully elaborated definition of biocrusts. In doing so, we illustrate the ecological relevance and ecosystem services provided by them. We demonstrate that biocrusts are defined by four distinct elements: physical structure, functional characteristics, habitat, and taxonomic composition. We describe outgroups, which have some, but not all, of the characteristics necessary to be fully consistent with our definition and thus would not be considered biocrusts. We also summarize the wide variety of different types of communities that fall under our definition of biocrusts, in the process of highlighting their global distribution. Finally, we suggest the universal use of the Belnap, Büdel & Lange definition, with minor modifications: Biological soil crusts (biocrusts) result from an intimate association between soil particles and differing proportions of photoautotrophic (e.g. cyanobacteria, algae, lichens, bryophytes) and heterotrophic (e.g. bacteria, fungi, archaea) organisms, which live within, or immediately on top of, the uppermost millimetres of soil. Soil particles are aggregated through the presence and activity of these often extremotolerant biota that desiccate regularly, and the resultant living crust covers the surface of the ground as a coherent layer. With this detailed definition of biocrusts, illustrating their ecological functions and widespread distribution, we hope to stimulate interest in biocrust research and inform various stakeholders (e.g. land managers, land users) on their overall importance to ecosystem and Earth system functioning., (© 2022 The Authors. Biological Reviews published by John Wiley & Sons Ltd on behalf of Cambridge Philosophical Society.)

Plants and other organisms have evolved structures and mechanisms for colonizing land since the Early Ordovician. In this context, their surfaces, the crucial physical interface with the environment, are mainly considered barriers against water loss. It is suggested that extreme water repellency (superhydrophobicity) was an additional key innovation for the transition of algae from water to land some 400 mya. Superhydrophobicity enhances gas exchange on land and excludes aquatic competitors in water films. In a different context, in material science and surface technology, superhydrophobicity has also become one of the most important bioinspired innovations enabling the avoidance of water films and contamination. Here, we present data for an extremely water-repellent cyanobacterial biofilm of the desiccation tolerant Hassallia byssoidea providing evidence for a much earlier prokaryotic Precambrian (ca. 1-2 bya) origin of superhydrophobicity and chemical heterogeneities associated with land transition. The multicellular cyanobacterium is functionally differentiated in a submerged basal hydrophilic absorbing portion like a "rhizoid" and an upright emersed superhydrophobic "phyllocauloid" filament for assimilation, nitrogen fixation, and splash dispersed diaspores. Additional data are provided for superhydrophobic surfaces in terrestrial green algae and in virtually all ancestral land plants (Bryophytes, ferns and allies, Amborella , Nelumbo ), slime molds, and fungi. Rethinking of superhydrophobicity as an essential first step for life in terrestrial environments is suggested., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Barthlott, Büdel, Mail, Neumann, Bartels and Fischer.)

The last decades of research led to a change in understanding of lichens that are now seen as self-sustaining micro-ecosystems, harboring diverse microbial organisms in tight but yet not fully understood relationships. Among the diverse interdependencies, the relationship between the myco- and photobiont is the most crucial, determining the shape, and ecophysiological properties of the symbiotic consortium. Roughly 10% of lichens associate with cyanobacteria as their primary photobiont, termed cyanolichens. Up to now, the diversity of cyanobionts of bipartite lichens resolved by modern phylogenetic approaches is restricted to the filamentous and heterocytous genera of the order Nostocales. Unicellular photobionts were placed in the orders Chroococcales, Pleurocapsales, and Chroococcidiopsidales. However, especially the phylogeny and taxonomy of the Chroococcidiopsidales genera remained rather unclear. Here we present new data on the identity and phylogeny of photobionts from cyanolichens of the genera Gonohymenia , Lichinella , Peccania , and Peltula from a broad geographical range. A polyphasic approach was used, combining morphological and cultivation-depending characteristics (microscopy, staining techniques, life cycle observation, baeocyte motility, and nitrogen fixation test) with phylogenetic analyses of the 16S rRNA and 16S-23S ITS gene region. We found an unexpectedly high cyanobiont diversity in the cyanobacterial lichens of the order Lichinales, including two new genera and seven new species, all of which were not previously perceived as lichen symbionts. As a result, we describe the novel unicellular Chroococcidiopsidales genera Pseudocyanosarcina gen. nov. with the species Pseudocyanosarcina phycocyania sp. nov. (from Peltula clavata , Australia) and Compactococcus gen. nov. with the species Compactococcus sarcinoides sp. nov. (from Gonohymenia sp., Australia) and the new Chroococcidiopsidales species Aliterella compacta sp. nov. (from Peltula clavata , Australia), Aliterella gigantea sp. nov. (from Peltula capensis ; South Africa), Sinocapsa ellipsoidea sp. nov. (from Peccania cerebriformis , Austria), as well as the two new Nostocales species Komarekiella gloeocapsoidea sp. nov. (from Gonohymenia sp., Czechia) and Komarekiella globosa sp. nov. (from Lichinella cribellifera , Canary Islands, Spain). Our study highlights the role of cyanolichens acting as a key in untangling cyanobacterial taxonomy and diversity. With this study, we hope to stimulate further research on photobionts, especially of rare cyanolichens., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Jung, Brust, Schultz, Büdel, Donner and Lakatos.)

Unicellular cyanobacteria inhabit a wide range of ecosytems and can be found throughout the phylum offering space for taxonomic confusion. One example is strain PCC 6712 that was described as Chlorogloea sp. (Nostocales) and later assigned to the genus Chroococcidiopsis (Chroococcidiopsidales). We now show that this strain belongs to the order Pleurocapsales and term it Hyella disjuncta based on morphology, genome analyses and 16S-23S ITS rRNA phylogeny. Genomic analysis indicated that H. disjuncta PCC 6712 shared about 44.7% orthologue genes with its closest relative H. patelloides . Furthermore, 12 cryptic biosynthetic gene clusters (BGCs) with potential bioactivity, such as a mycosporine-like amino acid BGC, were detected. Interestingly, the full set of nitrogen fixation genes was found in H. disjuncta PCC 6712 despite its inability to grow on nitrogen-free medium. A comparison of genes responsible for multicellularity was performed, indicating that most of these genes were present and related to those found in other cyanobacterial orders. This is in contrast to the formation of pseudofilaments-a main feature of the genus Hyella -which is weakly expressed in H. disjuncta PCC 6712 but prominent in Hyella patelloides LEGE 07179. Thus, our study pinpoints crucial but hidden aspects of polyphasic cyanobacterial taxonomy.

The taxonomy of coccoid cyanobacteria, such as Chroococcidiopsis , Pleurocapsa , Chroococcus , Gloeothece , Gloeocapsa , Gloeocapsopsis , and the related recent genera Sinocapsa and Aliterella , can easily be intermixed when solely compared on a morphological basis. There is still little support on the taxonomic position of some of the addressed genera, as genetic information is available only for a fraction of species that have been described solely on morphology. Modern polyphasic approaches that combine classic morphological investigations with DNA-based molecular analyses and the evaluation of ecological properties can disentangle these easily confusable unicellular genera. By using such an approach, we present here the formal description of two novel unicellular cyanobacterial species that inhabit the Coastal Range of the Atacama Desert, Gloeocapsopsis dulcis (first reported as Gloeocapsopsis AAB1) and Gloeocapsopsis diffluens . Both species could be clearly separated from previously reported species by 16S rRNA and 16S-23S ITS gene sequencing, the resulting secondary structures, p-distance analyses of the 16S-23S ITS, and morphology. For avoiding further confusions emendation of the genus Gloeocapsopsis as well as epitypification of the type species Gloeocapsopsis crepidinum based on the strain LEGE06123 were conducted., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Jung, Azua-Bustos, Gonzalez-Silva, Mikhailyuk, Zabicki, Holzinger, Lakatos and Büdel.)

Since 1965 a cyanobacterial strain termed ' Fischerella ambigua 108b' was the object of several studies investigating its potential as a resource for new bioactive compounds in several European institutes. Over decades these investigations uncovered several unique small molecules and their respective biosynthetic pathways, including the polychlorinated triphenyls of the ambigol family and the tjipanazoles. However, the true taxonomic character of the producing strain remained concealed until now. Applying a polyphasic approach considering the phylogenetic position based on the 16S rRNA and the protein coding gene rbcLX , secondary structures and morphological features, we present the strain ' Fischerella ambigua 108b' as Symphyonema bifilamentata sp. nov. 97.28. Although there is the type species (holotype) S. sinense C.-C. Jao 1944 there is no authentic living strain or material for genetic analyses for the genus Symphyonema available. Thus we suggest and provide an epitypification of S. bifilamentata sp. nov. 97.28 as a valid reference for the genus Symphyonema . Its affiliation to the family Symphyonemataceae sheds not only new light on this rare taxon but also on the classes of bioactive metabolites of these heterocytous and true-branching cyanobacteria which we report here. We show conclusively that the literature on the isolation of bioactive products from this organism provides further support for a clear distinction between the secondary metabolism of Symphyonema bifilamentata sp. nov. 97.28 compared to related and other taxa, pointing to the assignment of this organism into a separate genus.

Bioweathering mediated by microorganisms plays a significant role in biogeochemical cycles on global scales over geological timescales. Single processes induced by specific taxa have been described but could rarely be demonstrated for complex communities that dominate whole landscapes. The recently discovered grit crust of the coastal Atacama Desert, which is a transitional community between a cryptogamic ground cover and a rock-bound lithic assemblage, offers the unique chance to elucidate various bioweathering processes that occur simultaneously. Here, we present a bioweathering scenario of this biocenosis including processes such as penetration of the lithomatrix, microbial responses to wet-dry cycles, alkalinolysis, enzyme activity, and mineral re-localization. Frequently occurring fog, for example, led to a volume increase of microorganisms and the lithomatrix. This, together with pH shifts and dust accumulation, consequently results in biophysical breakdown and the formation of a terrestrial protopedon , an initial stage of pedogenesis fueled by the grit crust ., Competing Interests: The authors declare no competing interests., (© 2020 The Author(s).)

The polyphasic approach has been widely applied in cyanobacterial taxonomy, which frequently led to additions to the species inventory. Increasing our knowledge about species and the habitats they were isolated from enables new insights into the ecology of newly established genera and species allowing speculations about the ecological niche of taxa. Here, we are describing three new species belonging to three genera that broadens the ecological amplitude and the geographical range of each of the three genera. Cyanocohniella crotaloides sp. nov. is described from sandy beach mats of the temperate island Schiermonnikoog, Netherlands, Oculatella crustae-formantes sp. nov. was isolated from biological soil crusts of the Arctic Spitsbergen, Norway, and Aliterella chasmolithica originated from granitic stones of the arid Atacama Desert, Chile. All three species could be separated from related species using molecular sequencing of the 16S rRNA gene and 16S-23S ITS gene region, the resulting secondary structures as well as p-distance analyses of the 16S-23S ITS and various microscopic techniques. The novel taxa described in this study contribute to a better understanding of the diversity of the genera Cyanocohniella, Oculatella, and Aliterella in different habitats., (© 2020 The Authors. Journal of Phycology published by Wiley Periodicals LLC on behalf of Phycological Society of America.)

Biocrusts are associations of various prokaryotic and eukaryotic microorganisms in the top millimeters of soil, which can be found in every climate zone on Earth. They stabilize soils and introduce carbon and nitrogen into this compartment. The worldwide occurrence of biocrusts was proven by numerous studies in Europe, Africa, Asia and North America, leaving South America understudied. Using an integrative approach, which combines morphological and molecular characters (small subunit rRNA and ITS region), we examined the diversity of key biocrust photosynthetic organisms at four sites along the latitudinal climate gradient in Chile. The most northern study site was located in the Atacama Desert (arid climate), followed by open shrubland (semiarid climate), a dry forest region (Mediterranean climate) and a mixed broad leaved-coniferous forest (temperate climate) in the south. The lowest species richness was recorded in the desert (18 species), whereas the highest species richness was observed in the Mediterranean zone with (40 species). Desert biocrusts were composed exclusively of single-celled Chlorophyta algae, followed by cyanobacteria. Chlorophyta, Streptophyta and cyanobacteria dominated semiarid biocrusts, whereas Mediterranean and temperate Chilean biocrusts were composed mostly of Chlorophyta, Streptophyta and Ochrophyta. Our investigation of Chilean biocrust suggests high biodiversity of South American biocrust phototrophs.

The Atacama Desert is the driest non-polar desert on Earth, presenting precarious conditions for biological activity. In the arid coastal belt, life is restricted to areas with fog events that cause almost daily wet-dry cycles. In such an area, we discovered a hitherto unknown and unique ground covering biocenosis dominated by lichens, fungi, and algae attached to grit-sized (~6 mm) quartz and granitoid stones. Comparable biocenosis forming a kind of a layer on top of soil and rock surfaces in general is summarized as cryptogamic ground covers (CGC) in literature. In contrast to known CGC from arid environments to which frequent cyclic wetting events are lethal, in the Atacama Desert every fog event is answered by photosynthetic activity of the soil community and thus considered as the desert's breath. Photosynthesis of the new CGC type is activated by the lowest amount of water known for such a community worldwide thus enabling the unique biocenosis to fulfill a variety of ecosystem services. In a considerable portion of the coastal Atacama Desert, it protects the soil from sporadically occurring splash erosion and contributes to the accumulation of soil carbon and nitrogen as well as soil formation through bio-weathering. The structure and function of the new CGC type are discussed, and we suggest the name grit-crust. We conclude that this type of CGC can be expected in all non-polar fog deserts of the world and may resemble the cryptogam communities that shaped ancient Earth. It may thus represent a relevant player in current and ancient biogeochemical cycling., (© 2019 The Authors. Geobiology published by John Wiley & Sons Ltd.)

In the Atacama Desert, cyanobacteria grow on various substrates such as soils (edaphic) and quartz or granitoid stones (lithic). Both edaphic and lithic cyanobacterial communities have been described but no comparison between both communities of the same locality has yet been undertaken. In the present study, we compared both cyanobacterial communities along a precipitation gradient ranging from the arid National Park Pan de Azúcar (PA), which resembles a large fog oasis in the Atacama Desert extending to the semiarid Santa Gracia Natural Reserve (SG) further south, as well as along a precipitation gradient within PA. Various microscopic techniques, as well as culturing and partial 16S rRNA sequencing, were applied to identify 21 cyanobacterial species; the diversity was found to decline as precipitation levels decreased. Additionally, under increasing xeric stress, lithic community species composition showed higher divergence from the surrounding edaphic community, resulting in indigenous hypolithic and chasmoendolithic cyanobacterial communities. We conclude that rain and fog water, respectively, cause contrasting trends regarding cyanobacterial species richness in the edaphic and lithic microhabitats., (© 2019 The Authors. Journal of Phycology published by Wiley Periodicals, Inc. on behalf of Phycological Society of America.)

The Atacama Desert is one of the driest and probably oldest deserts on Earth where only a few extremophile organisms are able to survive. This study investigated two terricolous and two epiphytic lichens from the fog oasis "Las Lomitas" within the National Park Pan de Azúcar which represents a refugium for a few vascular desert plants and many lichens that can thrive on fog and dew alone. Ecophysiological measurements and climate records were combined with molecular data of the mycobiont, their green algal photobionts and lichenicolous fungi to gain information about the ecology of lichens within the fog oasis. Phylogenetic and morphological investigations led to the identification and description of the new lichen species Acarospora conafii sp. nov. as well as the lichenicolous fungi that accompanied them and revealed the trebouxioid character of all lichen photobionts. Their photosynthetic responses were compared during natural scenarios such as reactivation by high air humidity and in situ fog events to elucidate the activation strategies of this lichen community. Epiphytic lichens showed photosynthetic activity that was rapidly induced by fog and high relative air humidity whereas terricolous lichens were only activated by fog., (© 2019 The Authors. MicrobiologyOpen published by John Wiley & Sons Ltd.)

Cyanobacteria of biological soil crusts (BSCs) represent an important part of circumpolar and Alpine ecosystems, serve as indicators for ecological condition and climate change, and function as ecosystem engineers by soil stabilization or carbon and nitrogen input. The characterization of cyanobacteria from both polar regions remains extremely important to understand geographic distribution patterns and community compositions. This study is the first of its kind revealing the efficiency of combining denaturing gradient gel electrophoresis (DGGE), light microscopy and culture-based 16S rRNA gene sequencing, applied to polar and Alpine cyanobacteria dominated BSCs. This study aimed to show the living proportion of cyanobacteria as an extension to previously published meta-transcriptome data of the same study sites. Molecular fingerprints showed a distinct clustering of cyanobacterial communities with a close relationship between Arctic and Alpine populations, which differed from those found in Antarctica. Species richness and diversity supported these results, which were also confirmed by microscopic investigations of living cyanobacteria from the BSCs. Isolate-based sequencing corroborated these trends as cold biome clades were assigned, which included a potentially new Arctic clade of Oculatella. Thus, our results contribute to the debate regarding biogeography of cyanobacteria of cold biomes., (© 2018 The Authors. MicrobiologyOpen published by John Wiley & Sons Ltd.)

Seven new species and two varieties of Klebsormidium were described using an integrative approach on the base of 28 strains from the poorly studied phylogenetic superclade G. These strains originated from the unusual and exotic habitats (semi-deserts, semi-arid shrublands, Mediterranean shrub and deciduous vegetation, temperate Araucaria forests, peat bogs, dumps after coal mining, maritime sand dunes etc.) of four continents (Africa, South and North America, and Europe). Molecular phylogenies based on ITS-1,2, rbcL gene and concatenated dataset of ITS-1,2-rbcL, secondary structure of ITS-2, morphology, ecology and biogeography, micrographs and drawings of the investigated strains were assessed. Additionally, phylogeny and morphology of 18 Klebsormidium strains from other lineages isolated from the same localities (different vegetation types of Chile and maritime sand dunes of Germany) were investigated for the comparison with representatives of clade G. Clade G Klebsormidium is characterized by distant phylogenetic position from the other Klebsormidium lineages and prominent morphology: four-lobed chloroplasts and mostly short swollen cells in young culture, compact small pyrenoids, curved or disintegrated filaments, unusual elongation of cells in old culture, formation of specific cluster- and knot-like colonies on agar surface, especially prominent in strains isolated from desert regions, from which the group probably originated. Comparison of Klebsormidium diversity from different biogeographic regions showed that the representatives of clade G are common algae in regions of the southern hemisphere (South Africa and Chile) and rare representatives in terrestrial ecosystems of the northern hemisphere. Further investigation of mostly unstudied territories of the southern hemisphere could bring many surprises and discoveries, leading to a change of the present concept that Klebsormidium is cosmopolitan in distribution., (Copyright © 2018 Elsevier Inc. All rights reserved.)

The Antarctic Peninsula, a tundra biome dominated by lichens and bryophytes, is an ecozone undergoing rapid temperature shifts. Such changes may demand a high physiological plasticity of the local lichen species to maintain their role as key drivers in this pristine habitat. This study examines the response of net photosynthesis and respiration to increasing temperatures for three Antarctic lichen species with different ecological response amplitudes. We hypothesize that negative effects caused by increased temperatures can be mitigated by thermal acclimation of respiration and/or photosynthesis. The fully controlled growth chamber experiment simulated intermediate and extreme temperature increases over the time course of 6 weeks. Results showed that, in contrast to our hypothesis, none of the species was able to down-regulate temperature-driven respiratory losses through thermal acclimation of respiration. Instead, severe effects on photobiont vitality demonstrated that temperatures around 15°C mark the upper limit for the two species restricted to the Antarctic, and when mycobiont demands exceeded the photobiont capacity they could not survive within the lichen thallus. In contrast, the widespread lichen species was able to recover its homoeostasis by rapidly increasing net photosynthesis. We conclude that to understand the complete lichen response, acclimation processes of both symbionts, the photo- and the mycobiont, have to be evaluated separately. As a result, we postulate that any acclimation processes in lichen are species-specific. This, together with the high degree of response variability and sensitivity to temperature in different species that co-occur spatially close, complicates any predictions regarding future community composition in the Antarctic. Nevertheless, our results suggest that species with a broad ecological amplitude may be favoured with on-going changes in temperature., (© 2017 John Wiley & Sons Ltd.)

Dryland vegetation developed morphological and physiological strategies to cope with drought. However, as aridity increases, vascular plant coverage gets sparse and microbially-dominated surface communities (MSC), comprising cyanobacteria, algae, lichens and bryophytes together with heterotropic bacteria, archaea and fungi, gain relevance. Nevertheless, the relevance of MSC net primary productivity has only rarely been considered in ecosystem scale studies, and detailed information on their contribution to the total photosynthetic biomass reservoir is largely missing. In this study, we mapped the spatial distribution of two different MSC (biological soil crusts and quartz fields hosting hypolithic crusts) at two different sites within the South African Succulent Karoo (Soebatsfontein and Knersvlakte). Then we characterized both types of MSC in terms of chlorophyll content, and combining these data with the biocrust and quartz field maps, we estimated total biomass values of MSCs and their spatial patterns within the two different ecosystems. Our results revealed that MSC are important vegetation components of the South African Karoo biome, revealing clear differences between the two sites. At Soebatsfontein, MSC occurred as biological soil crusts (biocrusts), which covered about one third of the landscape reaching an overall biomass value of ~480gha -1 of chlorophyll a+b at the landscape scale. In the Knersvlakte, which is characterized by harsher environmental conditions (i.e. higher solar radiation and potential evapotranspiration), MSC occurred as biocrusts, but also formed hypolithic crusts growing on the lower soil-immersed parts of translucent quartz pebbles. Whereas chlorophyll concentrations of biocrusts and hypolithic crusts where insignificantly lower in the Knersvlakte, the overall MSC biomass reservoir was by far larger with ~780gha -1 of chlorophyll a+b. Thus, the complementary microbially-dominated surface communities promoted biomass formation within the environmentally harsh Knersvlakte ecosystem., (Copyright © 2017 Elsevier B.V. All rights reserved.)

Background: Many fungal species occur across a variety of habitats. Particularly lichens, fungi forming symbioses with photosynthetic partners, have evolved remarkable tolerances for environmental extremes. Despite their ecological importance and ubiquity, little is known about the genetic basis of adaption in lichen populations. Here we studied patterns of genome-wide differentiation in the lichen-forming fungus Lasallia pustulata along an altitudinal gradient in the Mediterranean region. We resequenced six populations as pools and identified highly differentiated genomic regions. We then detected gene-environment correlations while controlling for shared population history and pooled sequencing bias, and performed ecophysiological experiments to assess fitness differences of individuals from different environments., Results: We detected two strongly differentiated genetic clusters linked to Mediterranean and temperate-oceanic climate, and an admixture zone, which coincided with the transition between the two bioclimates. High altitude individuals showed ecophysiological adaptations to wetter and more shaded conditions. Highly differentiated genome regions contained a number of genes associated with stress response, local environmental adaptation, and sexual reproduction., Conclusions: Taken together our results provide evidence for a complex interplay between demographic history and spatially varying selection acting on a number of key biological processes, suggesting a scenario of ecological speciation.

Unraveling the complex relationship between lichen fungal and algal partners has been crucial in understanding lichen dispersal capacity, evolutionary processes, and responses in the face of environmental change. However, lichen symbiosis remains enigmatic, including the ability of a single fungal partner to associate with various algal partners. Psora decipiens is a characteristic lichen of biological soil crusts (BSCs), across semi-arid, temperate, and alpine biomes, which are particularly susceptible to habitat loss and climate change. The high levels of morphological variation found across the range of Psora decipiens may contribute to its ability to withstand environmental change. To investigate Psora decipiens acclimation potential, individuals were transplanted between four climatically distinct sites across a European latitudinal gradient for 2 years. The effect of treatment was investigated through a morphological examination using light and SEM microscopy; 26S rDNA and rbcL gene analysis assessed site-specific relationships and lichen acclimation through photobiont switching. Initial analysis revealed that many samples had lost their algal layers. Although new growth was often determined, the algae were frequently found to have died without evidence of a new photobiont being incorporated into the thallus. Mycobiont analysis investigated diversity and determined that new growth was a part of the transplant, thus, revealing that four distinct fungal clades, closely linked to site, exist. Additionally, P. decipiens was found to associate with the green algal genus Myrmecia, with only two genetically distinct clades between the four sites. Our investigation has suggested that P. decipiens cannot acclimate to the substantial climatic variability across its environmental range. Additionally, the different geographical areas are home to genetically distinct and unique populations. The variation found within the genotypic and morpho-physiological traits of P. decipiens appears to have a climatic determinant, but this is not always reflected by the algal partner. Although photobiont switching occurs on an evolutionary scale, there is little evidence to suggest an active environmentally induced response. These results suggest that this species, and therefore, other lichen species, and BSC ecosystems themselves may be significantly vulnerable to climate change and habitat loss.

Cyanobacteria associated with biological soil crusts (BSCs) have important attributes, such as nitrogen fixation and soil stabilisation. However, research on these organisms has been minimal, and their diversity and distribution throughout temperate Europe is currently unknown. The SCIN (Soil Crust International) project is a multidisciplinary research initiative that aims to achieve improved understanding of the BSCs of Europe, one facet being an investigation into the cyanobacterial communities of BSCs across a latitudinal gradient. Cyanobacteria assemblages were analysed by both morphological and molecular analysis. Two treatments were applied prior to DNA extraction, continued sample wetting and a dry sample process, and 16S ribosomal RNA (rRNA) amplicons were processed by Illumina MiSeq sequencing. The results reveal high and variable cyanobacterial diversity with each site showing a unique assemblage. Many common cyanobacterial genera, for example Nostoc and Microcoleus, were found in all sites but the abundances of different genera varied considerably. The polyphasic approach was found to be essential in recording the presence of important cyanobacteria that a single method itself did not highlight. The wet and dry treatments showed some differences in diversity, but mainly in abundance, this may suggest how cyanobacterial composition of BSCs changes with seasonal variability., (© FEMS 2016.)

Biological soil crusts (BSC) are the dominant functional vegetation unit in some of the harshest habitats in the world. We assessed BSC response to stress through changes in biotic composition, CO2 gas exchange and carbon allocation in three lichen-dominated BSC from habitats with different stress levels, two more extreme sites in Antarctica and one moderate site in Germany. Maximal net photosynthesis (NP) was identical, whereas the water content to achieve maximal NP was substantially lower in the Antarctic sites, this apparently being achieved by changes in biomass allocation. Optimal NP temperatures reflected local climate. The Antarctic BSC allocated fixed carbon (tracked using (14)CO2) mostly to the alcohol soluble pool (low-molecular weight sugars, sugar alcohols), which has an important role in desiccation and freezing resistance and antioxidant protection. In contrast, BSC at the moderate site showed greater carbon allocation into the polysaccharide pool, indicating a tendency towards growth. The results indicate that the BSC of the more stressed Antarctic sites emphasise survival rather than growth. Changes in BSC are adaptive and at multiple levels and we identify benefits and risks attached to changing life traits, as well as describing the ecophysiological mechanisms that underlie them.

Biological soil crusts (BSCs) are communities of cryptogamic organisms, occurring in arid and semiarid regions all over the world. Based on both morphological identification and genetic analyses, we established a first cyanobacterial inventory using the biphasic approach for BSCs within two major biomes of southern Africa. The samples were collected at two different sites in the Succulent Karoo and one in the Nama Karoo. After cultivation and morphological identification, the 16S rRNA gene was sequenced from the cyanobacterial cultures. From the soil samples, the DNA was extracted, and the 16S rRNA gene sequenced. All the sequences of the clone libraries from soil and cultures were compared with those of the public databases. Forty-five different species were morphologically identified in the samples of the Succulent Karoo (observatories of Soebatsfontein and Goedehoop). Based on the genetic analyses, 60 operational taxonomic units (OTUs) were identified for the Succulent Karoo and 43 for the Nama Karoo (based on 95% sequence similarity). The cloned sequences corresponded well with the morphologically described taxa in cultures and sequences in the public databases. Besides known species of typical crust-forming cyanobacterial genera (Microcoleus, Phormidium, Tolypothrix and Scytonema), we found sequences of so far undescribed species of the genera Leptolyngbya, Pseudanabaena, Phormidium, Oscillatoria, Schizothrix and Microcoleus. Most OTUs were restricted to distinct sites. Grazed soils showed lower taxa numbers than undisturbed soils, implying the presence of early successional crust types and reduced soil surface protection. Our combined approach of morphological identification and genetic analyses allowed both a taxa inventory and the analysis of species occurring under specific habitat conditions.

Here we report details of the European research initiative "Soil Crust International" (SCIN) focusing on the biodiversity of biological soil crusts (BSC, composed of bacteria, algae, lichens, and bryophytes) and on functional aspects in their specific environment. Known as the so-called "colored soil lichen community" (Bunte Erdflechtengesellschaft), these BSCs occur all over Europe, extending into subtropical and arid regions. Our goal is to study the uniqueness of these BSCs on the regional scale and investigate how this community can cope with large macroclimatic differences. One of the major aims of this project is to develop biodiversity conservation and sustainable management strategies for European BSCs. To achieve this, we established a latitudinal transect from the Great Alvar of Öland, Sweden in the north over Gössenheim, Central Germany and Hochtor in the Hohe Tauern National Park, Austria down to the badlands of Tabernas, Spain in the south. The transect stretches over 20° latitude and 2,300 m in altitude, including natural (Hochtor, Tabernas) and semi-natural sites that require maintenance such as by grazing activities (Öland, Gössenheim). At all four sites BSC coverage exceeded 30 % of the referring landscape, with the alpine site (Hochtor) reaching the highest cyanobacterial cover and the two semi-natural sites (Öland, Gössenheim) the highest bryophyte cover. Although BSCs of the four European sites share a common set of bacteria, algae (including cyanobacteria) lichens and bryophytes, first results indicate not only climate specific additions of species, but also genetic/phenotypic uniqueness of species between the four sites. While macroclimatic conditions are rather different, microclimatic conditions and partly soil properties seem fairly homogeneous between the four sites, with the exception of water availability. Continuous activity monitoring of photosystem II revealed the BSCs of the Spanish site as the least active in terms of photosynthetic active periods.

Facilitative effects and plant-plant interactions are well known for higher plants, but there is a lack of information about their relevance in cryptogams. Additional information about facilitative effects between bryophytes and lichens would be an important contribution to recent research on positive plant-plant interactions, as these can have striking influences not only on the organisation of early successional terrestrial communities but also on succession dynamics by kick-starting ecosystem development through the import of key nutrients. We investigated and quantified these mechanisms between Peltigera rufescens and its associated mosses. Moss-associated thalli had a different morphology that led to several benefits from the association. They had 66% higher net photosynthetic rate and, because the majority of the gas exchange of lichen thalli took place through the lower surface, there was a further increase as the CO(2) concentration was >25% higher beneath moss-associated thalli. Microclimatic measurements showed that mean light levels were substantially lower and temperature extremes slightly ameliorated for moss-associated thalli. As a consequence, desiccation was slower which is, together with an increase in thallus thickness and water storage, the reason for extended periods of optimal net photosynthesis for the moss-associated thalli. All these benefits combined to produce a growth rate of the moss-associated thalli which was significantly higher, twice that of non-associated thalli [0.75 ± 0.4 vs. 0.30 ± 0.1 mm/month (mean ± SD)]. This appears to be the first demonstration of a strong mechanistic basis for facilitative effects between lichens and bryophytes.

• Additional water supplied by dew formation is an important resource for microbes, plants and animals in precipitation-limited habitats, but has received little attention in tropical forests until now. • We evaluated the micro-environmental conditions of tree stem surfaces and their epiphytic organisms in a neotropical forest, and present evidence for a novel mechanism of diurnal dew formation on these surfaces until midday that has physiological implications for corticolous epiphytes such as lichens. • In the understorey of a lowland forest in French Guiana, heat storage of stems during the day and delayed radiative loss during the night decreased stem surface temperatures by 6°C in comparison to the dew-point temperature of ambient air. This measured phenomenon induced modelled totals of diurnal dew formation between 0.29 and 0.69 mm d⁻¹ on the surface of the bark and the lichens until early afternoon. • Crustose lichens substantially benefit from this dew formation, because it prolongs photosynthetic activity. This previously unrecognized mechanism of midday dew formation contributes to the water supply of most corticolous organisms, and may be a general feature in forest habitats world-wide., (© 2012 The Authors. New Phytologist © 2012 New Phytologist Trust.)

In the animal kingdom, carotenoids are usually absorbed from dietary sources and transported to target tissues. Despite their general importance, the uptake mechanism is still poorly understood. Here we report the "red crop" phenomenon, an accumulation of α- and β-carotene in crystalline inclusions in the enlarged foregut of the polyphagous Spodoptera larvae feeding on some potentially toxic plant leaves. The carotene crystals give the insect foregut a distinctive orange-red color. The crystals are embedded in a homogenous lawn of the bacterium Enterococcus casseliflavus, but the carotene seems to be selectively taken from the food plant. Caterpillars which fail to develop these carotene crystals exhibit a high mortality or fail to develop to adulthood. The crystallization of carotene and the enlargement of the foregut thus appears to manifest a multiple-step physiological adaptation of the insects to toxic food plants., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

The two endolithic lichen species Hymenelia prevostii and Hymenelia coerulea were investigated with regard to their thallus morphology and their effects on the surrounding substrate. The physiological processes responsible for the observed alterations of the rock were identified. Whereas the thallus surface of H. coerulea was level, H. prevostii formed small depressions that were deepest in the thallus center. In a cross-section, both species revealed an algal zone consisting of algal cavities parallel to the substrate surface and a fungal zone below. However, H. prevostii revealed significantly larger cavities with more than twice the cell number and a denser pattern of cavities than H. coerulea, resulting in a biomass per surface area being more than twice as large. Below H. prevostii the layer of macroscopically visibly altered rock material was about twice as deep and within this layer, the depletion of calcium and manganese was considerably higher. In simultaneous measurements of the oxygen uptake/oxygen release and pH shift, the isolated algal strains of both lichens revealed respiration-induced acidification of the medium in the dark. At higher light intensities, H. coerulea and to a lesser extent also H. prevostii alkalized the medium which may lessen the acidification effect somewhat under natural conditions. In a long-term growth experiment, the isolated algal strains of both lichens revealed acidification of the medium to a similar extent. Neither acidic lichen substances nor oxalic acid was identified. The significant differences between the weathering patterns of both species are based on the same respiration-induced acidification mechanism, with H. prevostii having a greater effect due to its higher biomass per area., (© 2010 Blackwell Publishing Ltd.)

The rough bark of orchard trees (Malus) around Darmstadt is predominantly covered in red to purple-brown layers (biofilms) of epiphytic terrestrial alga of Trentepohlia umbrina. The smooth bark of forest trees (Fagus sylvatica L. and Acer sp.) in the same area is covered by bright green biofilms composed of the green algae Desmococcus, Apatococcus and Trebouxia, with a few cells of Coccomyxa and 'Chlorella' trebouxioides between them. These algae are desiccation tolerant. After samples of bark with the biofilms were kept in dry air in darkness for various periods of time, potential quantum yield of PSII, F(v)/F(m), recovered during rehydration upon rewetting. The kinetics and degree of recovery depended on the length of time that the algae were kept in dry air in the desiccated state. Recovery was better for green biofilm samples, i.e. quite good even after 80 days of desiccation (F(v)/F(m) = ca. 50% of initial value), than the red samples, where recovery was only adequate up to ca. 30-40 days of desiccation (F(v)/F(m) = ca. 20-55% of initial value). It is concluded that the different bark types constitute different ecophysiological niches that can be occupied by the algae and that can be distinguished by their capacity to recover from desiccation after different times in the dry state.

We present a 6-gene, 420-species maximum-likelihood phylogeny of Ascomycota, the largest phylum of Fungi. This analysis is the most taxonomically complete to date with species sampled from all 15 currently circumscribed classes. A number of superclass-level nodes that have previously evaded resolution and were unnamed in classifications of the Fungi are resolved for the first time. Based on the 6-gene phylogeny we conducted a phylogenetic informativeness analysis of all 6 genes and a series of ancestral character state reconstructions that focused on morphology of sporocarps, ascus dehiscence, and evolution of nutritional modes and ecologies. A gene-by-gene assessment of phylogenetic informativeness yielded higher levels of informativeness for protein genes (RPB1, RPB2, and TEF1) as compared with the ribosomal genes, which have been the standard bearer in fungal systematics. Our reconstruction of sporocarp characters is consistent with 2 origins for multicellular sexual reproductive structures in Ascomycota, once in the common ancestor of Pezizomycotina and once in the common ancestor of Neolectomycetes. This first report of dual origins of ascomycete sporocarps highlights the complicated nature of assessing homology of morphological traits across Fungi. Furthermore, ancestral reconstruction supports an open sporocarp with an exposed hymenium (apothecium) as the primitive morphology for Pezizomycotina with multiple derivations of the partially (perithecia) or completely enclosed (cleistothecia) sporocarps. Ascus dehiscence is most informative at the class level within Pezizomycotina with most superclass nodes reconstructed equivocally. Character-state reconstructions support a terrestrial, saprobic ecology as ancestral. In contrast to previous studies, these analyses support multiple origins of lichenization events with the loss of lichenization as less frequent and limited to terminal, closely related species.

Biological soil crusts (BSCs) are found in all dryland regions of the world, including the polar regions. They are also known to occur in the southern African region. Although there were a number of case studies on BSCs from that region, we did not know if they are a normal part of the vegetation cover or just a phenomenon that occasionally occurs here and there. In order to investigate diversity, distribution patterns, and the driving factors of both, we followed a random sampling system of observatories along a transect, stretching from the Namibian-Angolan border down south to the Cape Peninsula, covering seven different major biomes. Biological soil crusts were found to occur in six out of seven biomes. Despite the fact that soil-dwelling algae occurred in the Fynbos biome, crust formation was not observed for hitherto unknown reasons. Seven BSC types were distinguished on the basis of morphology and taxonomic composition: three of them were cyanobacteria-dominated, one with additional chlorolichens, two with bryophytes, one hypolithic type restricted to quartz gravel pavements, and the unique lichen fields of the Namib Desert. Besides 29 green algal species in 21 genera, one heterokont alga, 12 cyanolichens, 14 chlorolichens, two genera of liverworts, and three genera of mosses, these crusts are positioned among the most diverse BSCs worldwide mainly because of the unusual high cyanobacterial species richness comprising 58 species in 21 genera. They contribute considerably to the biodiversity of arid and semi-arid bioregions. Taxonomic diversity of cyanobacteria was significantly higher in the winter rain zone than in the summer rain zone (54 versus 32 species). The soil photosynthetic biomass (chlorophylla/m2), the carbon content of the soil and the number of BSC types were significantly higher in the winter rain zone (U27, 29=215.0, p=0.004 [chla]; U21, 21=135.0, p=0.031 [C]; U27, 29=261.5, p=0.028 [BSC types]; excluding the fog-dominated Namib biome). The winter rain zone is characterized by a lower precipitation amount, but a higher rain frequency with the number of rainy days more evenly distributed over the year. The dry period is significantly shorter per year in the winter rain zone (U8, 9=5.0, p=0.003). We conclude that rain frequency and duration of dry periods rather than the precipitation amount is the main factor for BSC growth and succession. Nitrogen content of the soils along the transect was generally very low and correlated with soil carbon content. There was a weak trend that an increasing proportion of silt and clay (<0.63 mm) in the soil is associated with higher values of BSC chlorophyll content (Pearson correlation coefficient=0.314, p=0.237). Furthermore, we found a significant positive correlation between silt and clay and the number of BSC types (Pearson correlation coefficient=0.519, p=0.039), suggesting that fine grain-size promotes BSC succession and their biomass content. Lichens and bryophytes occurred in BSCs with lower disturbance frequencies (e.g. trampling) only. Crust thickness and chlorophyll content increased significantly from crusts of the early successional type to the late successional crust types. From our results, we conclude that BSCs are a normal and frequent element of the vegetation in arid and semi-arid southwestern Africa, and that rain frequency and duration of dry periods rather than the precipitation amount are the key factors for the development, differentiation and composition of BSCs.

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., (© 2008 Phycological Society of America.)

In this study we analyzed the genetic population structure of the hygrophilous tall-herb Cicerbita alpina in the western Alps because this group of mountain plants is underrepresented in the biogeographical literature. AFLP (amplified fragment length polymorphism) fingerprints of 40 samples were analyzed from four populations situated in a transect from the southwestern Alps to the eastern part of the western Alps and one population from the Black Forest outside the Alps. Two genetic groups can be distinguished. The first group (A) comprises the populations from the northern and eastern parts of the western Alps, and the second group (B) comprises the populations from the southwestern Alps and the Black Forest. Group A originates most likely from at least one refugium in the southern piedmont regions of the Alps. This result provides molecular evidence for a humid climate at the southern margin of the Alps during the Würm glaciation. Group B originates presumably from western or northern direction and we discuss two possible scenarios for the colonization of the Alps, i.e. (1) long-distance dispersal from southwestern refugia and (2) colonization from nearby refugia in the western and/or northern Alpine forelands. The study demonstrates that the target species harbours considerable genetic diversity, even on a regional scale, and therefore is a suitable model for phylogeographic research.

Pezizomycotina is the largest subphylum of Ascomycota and includes the vast majority of filamentous, ascoma-producing species. Here we report the results from weighted parsimony, maximum likelihood and Bayesian phylogenetic analyses of five nuclear loci (SSU rDNA, LSU rDNA, RPB1, RPB2 and EF-lalpha) from 191 taxa. Nine of the 10 Pezizomycotina classes currently recognized were represented in the sampling. These data strongly supported the monophyly of Pezizomycotina, Arthoniomycetes, Eurotiomycetes, Orbiliomycetes and Sordariomycetes. Pezizomycetes and Dothideomycetes also were resolved as monophyletic but not strongly supported by the data. Lecanoromycetes was resolved as paraphyletic in parsimony analyses but monophyletic in maximum likelihood and Bayesian analyses. Leotiomycetes was polyphyletic due to exclusion of Geoglossaceae. The two most basal classes of Pezizomycotina were Orbiliomycetes and Pezizomycetes, both of which comprise species that produce apothecial ascomata. The seven remaining classes formed a monophyletic group that corresponds to Leotiomyceta. Within Leotiomyceta, the supraclass clades of Leotiomycetes s.s. plus Sordariomycetes and Arthoniomycetes plus Dothideomycetes were resolved with moderate support.

The Lecanoromycetes includes most of the lichen-forming fungal species (> 13500) and is therefore one of the most diverse class of all Fungi in terms of phenotypic complexity. We report phylogenetic relationships within the Lecanoromycetes resulting from Bayesian and maximum likelihood analyses with complementary posterior probabilities and bootstrap support values based on three combined multilocus datasets using a supermatrix approach. Nine of 10 orders and 43 of 64 families currently recognized in Eriksson's classification of the Lecanoromycetes (Outline of Ascomycota--2006 Myconet 12:1-82) were represented in this sampling. Our analyses strongly support the Acarosporomycetidae and Ostropomycetidae as monophyletic, whereas the delimitation of the largest subclass, the Lecanoromycetidae, remains uncertain. Independent of future delimitation of the Lecanoromycetidae, the Rhizocarpaceae and Umbilicariaceae should be elevated to the ordinal level. This study shows that recent classifications include several nonmonophyletic taxa at different ranks that need to be recircumscribed. Our phylogenies confirm that ascus morphology cannot be applied consistently to shape the classification of lichen-forming fungi. The increasing amount of missing data associated with the progressive addition of taxa resulted in some cases in the expected loss of support, but we also observed an improvement in statistical support for many internodes. We conclude that a phylogenetic synthesis for a chosen taxonomic group should include a comprehensive assessment of phylogenetic confidence based on multiple estimates using different methods and on a progressive taxon sampling with an increasing number of taxa, even if it involves an increasing amount of missing data.

The ancestors of fungi are believed to be simple aquatic forms with flagellated spores, similar to members of the extant phylum Chytridiomycota (chytrids). Current classifications assume that chytrids form an early-diverging clade within the kingdom Fungi and imply a single loss of the spore flagellum, leading to the diversification of terrestrial fungi. Here we develop phylogenetic hypotheses for Fungi using data from six gene regions and nearly 200 species. Our results indicate that there may have been at least four independent losses of the flagellum in the kingdom Fungi. These losses of swimming spores coincided with the evolution of new mechanisms of spore dispersal, such as aerial dispersal in mycelial groups and polar tube eversion in the microsporidia (unicellular forms that lack mitochondria). The enigmatic microsporidia seem to be derived from an endoparasitic chytrid ancestor similar to Rozella allomycis, on the earliest diverging branch of the fungal phylogenetic tree.

In tropical lowland forests, corticolous crustose green algal lichens are abundant and highly diverse. This may be related to adaptation to prevailing microenvironmental conditions including, for example, high precipitation and low light intensities. In the understory of a tropical lowland rain forest in French Guiana, we studied the morphology of crustose green algal lichens and measured gas exchange and chlorophyll a fluorescence. We found that (i) periods of thallus suprasaturation with water were reduced by the presence of water-repelling surface structures of mycobiont hyphae at the thallus surface and the medulla; (ii) photosynthesis was adapted to the low light intensities present in the understory; and (iii) photosynthesis was rapidly activated in fluctuating light. The combination of these three mechanisms enables corticolous lichens to implement specific morphological and physiological strategies, which may favour growth in the limiting understory habitat of tropical lowland rain forests.

Many filamentous cyanobacteria reduce atmospheric nitrogen in specialized differentiated cells called heterocysts. Here we present evidence that shows that members of the unicellular non-heterocyst-differentiating genus Chroococcidiopsis and the filamentous heterocyst-differentiating cyanobacteria are each other's closest living relatives. Distance, maximum-parsimony, and maximum-likelihood analyses of complete small subunit ribosomal RNA gene sequences yielded highly congruent support for the monophyly of Chroococcidiopsis and the heterocyst-differentiating cyanobacteria. Our results demonstrate that the order Pleurocapsales, which traditionally contains Chroococcidiopsis, is a polyphyletic assemblage with the ability to reproduce by multiple fission having arisen independently at least twice during the cyanobacterial radiation. Our data also reject Myxosarcina as a sister taxon to Chroococcidiopsis, indicating that the numerous presumed shared derived characters thought to unite the two genera evolved independently. The sequence divergence within the Chroococcidiopsis lineage is comparable to and probably exceeds that in the entire heterocyst-differentiating lineage. Chroococcidiopsis forms unique survival cells under nitrogen-limiting conditions, and the sister group relationship with the heterocystous cyanobacteria shown here suggests that differentiation of these cells and heterocysts may be related processes.

The occurrence of ultraviolet (UV)-absorbing substances like scytonemin and mycosporine-glycine is reported for the first time from cyanobacterial lichens of the genera Collema, Gonohymenia and Peltula, all coming from high-light-intensity habitats. Except for Collema with the filamentous Nostoc, all other cyanobionts belong to the unicellular genera Chroococcidiopsis, Cyanosarcina, Gloeocapsa or Myxosarcina. From transmission electron microscope studies it is evident that the pigmentation (scytonemin) is located extracellularly in the sheath of the outer thallus parts. Fluorescence microscopy and microprobe measurements clearly show UV radiation into the lichen thallus and hence the relevance of UV sunscreens for the protection of the organism.

CO 2 exchange rate in relation to thallus water content (WC, % of dry weight) was determined for 22 species of lichens, mainly members of the genera Pseudocyphellaria and Sticta, from a temperate rainforest, Urewere National Park, New Zealand. All data were obtained in the field, either using a standard technique in which the lichens were initially wetted (soaked or sprayed, then shaken) and allowed to slowly dry, or from periodic measurements on samples that were continuously exposed in their natural habitat. A wide range of WC was found, with species varying from 357 to 3360% for maximal WC in the field, and from 86 to 1300% for optimal WC for photosynthesis. Maximal WC for lichens, wetted by the standard technique, were almost always much less than the field maxima, due to the presence of water on the thalli. The relationships between CO 2 exchange rate and WC could be divided into four response types based on the presence, and degree, of depression of photosynthesis at high WC. Type A lichens showed no depression, and Type B only a little at maximal WC. Type C had a very large depression and, at the highest WC, CO 2 release could occur even in the light. Photosynthetic depression commenced soon after optimal WC was reached. Type D lichens showed a similar depression but the response curve had an inflection so that net photosynthesis was low but almost constant, and never negative, at higher WC. There was little apparent relationship between lichen genus or photobiont type and the response type. It was shown that high WC does limit photosynthetic CO 2 uptake under natural conditions. Lichens, taken directly from the field and allowed to dry under controlled conditions, had net photosynthesis rates that were initially strongly inhibited but rose to an optimum, before declining at low WC. The limiting effects of high WC were clearly shown when, under similar light conditions, severe photosynthetic depression followed a brief, midday, rain storm. Over the whole measuring period the lichens were rarely at their optimal WC for photosynthesis, being mostly too wet or, occasionally, too dry. Photosynthetic performance by the lichens exposed in the field was similar to that expected from the relationship between the photosynthetic rate and WC established by the standard procedure.