Martin Grube

Karl-Franzens-Universität Graz | KFU Graz

Exposing their vegetative bodies to the light, lichens are outstanding amongst other fungal symbioses. Not requiring a pre‐established host, ‘lichenized fungi’ build an entirely new structure together with microbial photosynthetic partners that neither can form alone. The signals involved in the transition of a fungus and a compatible photosyntheti...

Lichens have been reappraised as self-sustaining and long-living ecosystems in which a multiplicity of microorganisms are housed, in addition to the main symbiotic partners. Lichen-associated microfungi can frequently occur cryptically, and their species diversity has recently been more fully elucidated by DNA metabarcoding studies and culture isol...

Taxonomists consider species as discrete units of biological organization, which are subject to a continuous process of evolutionary change and are connected through their shared ancestry. However, the continuous nature of evolutionary change is difficult to reconcile with the discrete outcome of speciation, especially where species boundaries are...

Lichens are well-known examples of complex symbiotic associations between organisms from different Kingdoms. Microfungi in particular, establish diverse associations with the hosting lichen thallus, as species-specific parasites or transient co-inhabitants. The whole community of lichen-associated fungi constitute the ‘lichen mycobiome’ and compris...

Seven Fusarium species complexes are treated, namely F. aywerte species complex (FASC) (two species), F. buharicum species complex (FBSC) (five species), F. burgessii species complex (FBURSC) (three species), F. camptoceras species complex (FCAMSC) (three species), F. chlamydosporum species complex (FCSC) (eight species), F. citricola species compl...

Lichen collected worldwide for centuries have resulted in millions of specimens deposited in herbaria that offer the potential to assess species boundaries, phenotypic diversification, ecology, and distribution. The application of molecular approaches to historical collections has been limited due to DNA fragmentation, but high-throughput sequencin...

Fungal–algal relationships—both across evolutionary and ecological scales—are finely modulated by the presence of the symbionts in the environments and by the degree of selectivity and specificity that either symbiont develop reciprocally. In lichens, the green algal genus Trebouxia Puymaly is one of the most frequently recovered chlorobionts. Treb...

Recent publications have argued that there are potentially serious consequences for researchers in recognising distinct genera in the terminal fusarioid clade of the family Nectriaceae. Thus, an alternate hypothesis, namely a very broad concept of the genus Fusarium was proposed. In doing so, however, a significant body of data that supports distin...

Lichen thalli provide a long-lived and stable habitat for colonization by a wide range of microorganisms. Increased interest in these lichen-associated microbial communities has revealed an impressive diversity of fungi, including several novel lineages which still await formal taxonomic recognition. Among these, members of the Eurotiomycetes and D...

Lichens host highly diverse microbial communities, with bacteria being one of the most explored groups in terms of their diversity and functioning. These bacteria could partly originate from symbiotic propagules developed by many lichens and, perhaps more commonly and depending on environmental conditions, from different sources of the surroundings...

Lichens represent self-supporting symbioses, which occur in a wide range of terrestrial habitats and which contribute significantly to mineral cycling and energy flow at a global scale. Lichens usually grow much slower than higher plants. Nevertheless, lichens can contribute substantially to biomass production. This review focuses on the lichen sym...

Background Antimicrobial resistance (AMR) is a major threat to public health. Microorganisms equipped with AMR genes are suggested to have partially emerged from natural habitats; however, this hypothesis remains inconclusive so far. To understand the consequences of the introduction of exogenic antimicrobials into natural environments, we exposed...

The polyphyletic group of black fungi within the Ascomycota (Arthoniomycetes, Dothideomycetes, and Eurotiomycetes) is ubiquitous in natural and anthropogenic habitats. Partly because of their dark, melanin-based pigmentation, black fungi are resistant to stresses including UV-and ionizing-radiation, heat and desiccation, toxic metals, and organic p...

Assessing the ecological impacts of environmental change on biological communities requires knowledge of the factors driving the spatial patterns of the three diversity facets along extensive environmental gradients. We quantified the taxonomic (TD), functional (FD), and phylogenetic diversity (PD) of lichen epiphytic communities in 23 beech forest...

Lichenized fungi usually develop complex, stratified morphologies through an intricately balanced living together with their algal partners, but several species are known to form only more or less loose associations with algae. These borderline lichens are still little explored although they could inform us about early stages of lichen evolution. W...

Bacteria have recently emerged as important constituents of lichen ho-lobionts. Here, 29 bacterial metagenome-assembled genome (MAG) sequences were reconstructed from lichen metagenomes and taxonomically classified in four phyla. These results provide a pivotal resource for further exploration of the ecological roles played by bacterial symbionts i...

Abstract Lichens provide valuable systems for studying symbiotic interactions. In lichens, these interactions are frequently described in terms of availability, selectivity and specificity of the mycobionts and photobionts towards one another. The lichen-forming, green algal genus Trebouxia Puymaly is among the most widespread photobiont, associati...

Characterizing functional trait variation and covariation, and its drivers, is critical to understand the response of species to changing environmental conditions. Evolutionary and environmental factors determine how traits vary among and within species at multiple scales. However, disentangling their relative contribution is challenging and a comp...

We have studied the highly oceanic genus Coniocarpon in Norway. Our aim has been to delimit species of Coniocarpon in Norway based on an integrative taxonomic approach. The material studied comprises 120 specimens of Coniocarpon , obtained through recent collecting efforts (2017 and 2018) or received from major fungaria in Denmark, Finland, Norway...

Fungi with dark-coloured cells and mycelia—also known as black fungi—form a ubiquitous fraction of microbial communities on rock surfaces and often occur on rock-inhabiting lichens as well. The diversity and lifestyles of these fungi are still insufficiently known. In this chapter we review the current state of knowledge about the diversity and bio...

Climate change and the anthropic emission of pollutants are likely to have an accelerated impact in high-elevation mountain areas. This phenomenon could have negative consequences on alpine habitats and for species of conservation in relative proximity to dense human populations. This premise implies that the crucial task is in the early detection...

Rock-inhabiting fungi (RIF) are adapted to thrive in oligotrophic environments and to survive under conditions of abiotic stress. Under these circumstances, they form biocoenoses with other tolerant organisms, such as lichens, or with less specific phototrophic consortia of aerial algae or cyanobacteria. RIF are phylogenetically diverse, and their...

Background Arugula is a traditional medicinal plant and popular leafy green today. It is mainly consumed raw in the Western cuisine and known to contain various bioactive secondary metabolites. However, arugula has been also associated with high-profile outbreaks causing severe food-borne human diseases. A multiphasic approach integrating data from...

Borderline lichens are simple mutualistic symbioses between fungi and algae, where the fungi form loose mycelia interweaving algal cells, instead of forming a lichen thallus. Schizoxylon albescens shows two nutritional modes: it can either live as a borderline lichen on Populus tremula bark or as a saprotroph on Populus wood. This enables us to inv...

Host-associated microbiota play an important role in the health and persistence of more complex organisms. In this study, metagenomic analyses were used to reveal microbial community adaptations in three lichen samples as a response to different arsenic concentrations at the sampling sites. Elevated arsenic concentrations at a former mining site ex...

The role of host-associated microbiota in enduring dehydration and drought is largely unknown. We have used lichens to study this increasingly important problem because they are the organisms that are optimally adapted to reoccurring hydration/dehydration cycles, and they host a defined and persistent bacterial community. The analysis of metatransc...

The microbial diversity and function of terrestrial lichens have been well studied, but knowledge about the non-photosynthetic bacteria associated with marine lichens is still scarce. 16S rRNA gene Illumina sequencing was used to assess the culture-independent bacterial diversity in the strictly marine cyanolichen species Lichina pygmaea and Lichin...

Sample location and environmental conditions List of lichens sampled with details of the different clusters, their geographical location and the temperature at the time of sampling.

Closest relatives of lichen marker gene sequences as assessed by blastn Results of the blastn analyses of the sequenced marker genes used to check the morphological lichen identifications.

Closest relatives assessed by blastn of the top marine and terrestrial OTUs associated with the lichen species List of the 30 most abundant OTUs for either the marine lichens or the maritime-inland terrestrial lichen group, presented together with their GreenGenes taxonomic identity and the closest relative obtained by blastn analysis.

Supplementary figures and data analysis pipeline Supplementary information comprising supplementary figures and details of the data analysis pipeline. Figure S1. Relative abundance (%) of bacterial and cyanobacterial sequences recovered from each lichen species. Xanthoria sp. Atl; from Brittany, and Xanthoria sp. Med. from the Mediterranean coast....

Bacterial isolates and lichen bacterial OTUs showing >97% 16S rRNA gene sequence identity List of lichen bacterial 16S rRNA OTUs and their taxonomy that showed >97% 16S rRNA gene sequence identity to bacterial isolates cultured from L. pygmaea, L. confinis and L. auriforme.

Barcodes and primers for each sample List of sample IDs for each lichen sample and their corresponding barcode used to tag the forward primer 341F.

The clade of Myxogastria, commonly described as true or plasmodial slime molds, contains more than 1000 species. During their life cycle many of these slime molds develop extended networks of connected veins, known as unicellular (phanero)plasmodia. Among those, Physarum polycephalum gathered by far the most attention of biologists and physicists....

The microbial diversity and function of terrestrial lichens has been well studied, but knowledge about the non-photosynthetic bacteria associated with marine lichens is still scarce. 16S rRNA gene Illumina sequencing was used to assess the culture-independent bacterial diversity in the strictly marine cyanolichen species Lichina pygmaea and Lichina...

Arthonia sanguinaria is described as new to science. The East Asian Arthonia lopingensis and the widely distributed Arthonia picea are reported as new to Japan from Honshu and the Ogasawara Islands, respectively. The phylogenetic position of A. picea, A. sanguinaria and Coniocarpon cinnabarinum from the Ogasawara Islands is shown by RAxML and Bayes...

Nomenclatural type definitions are one of the most important concepts in biological nomenclature. Being physical objects that can be re-studied by other researchers, types permanently link taxonomy (an artificial agreement to classify biological diversity) with nomenclature (an artificial agreement to name biological diversity). Two proposals to am...

Lichen symbioses develop long-living thallus structures even in the harshest environments on Earth. These structures are also habitats for many other microscopic organisms, including other fungi, which vary in their specificity and interaction with the whole symbiotic system. This contribution reviews the recent progress regarding the understanding...

The thallus structure of the lichen symbiosis provides a fungal shelter for the growth of algal partners. The long-living thallus also provides a habitat for other fungi, but experimental studies, which could inform us about the details of their interactions have hardly been conducted. We present a new approach by embedding axenically cultured stra...

Biological soil crusts (biocrusts) cover about 12% of the Earth's land masses, thereby providing ecosystem services and affecting biogeochemical fluxes on a global scale. They comprise photoautotrophic cyanobacteria, algae, lichens and mosses, which grow together with heterotrophic microorganisms, forming a model system to study facilitative intera...

Alphaproteobacterium strain MOLA1416, related to Mycoplana ramosa DSM 7292 and Chelativorans intermedius CC-MHSW-5 (93.6% 16S rRNA sequence identity) was isolated from the marine lichen, Lichina pygmaea and its chemical composition was characterized by a metabolomic network analysis using LC-MS/MS data. Twenty-five putative different compounds were...

Biological soil crusts are ecologically important communities in areas where vascular plant coverage is low, and their presence is often vital in prevention of soil erosion. Despite recurrent threats to biological soil crusts across different environments, their recovery after disturbance has been little studied. We therefore established experiment...

As self-supporting and long-living symbiotic structures, lichens provide a habitat for many other organisms beside the traditionally considered lichen symbionts – the myco- and the photobionts. The lichen-inhabiting fungi either develop diagnostic phenotypes or occur asymptomatically. Because the degree of specificity toward the lichen host is poor...

Background Recent evidence of specific bacterial communities extended the traditional concept of fungal-algal lichen symbioses by a further organismal kingdom. Although functional roles were already assigned to dominant members of the highly diversified microbiota, a substantial fraction of the ubiquitous colonizers remained unexplored. We employed...

Microscopic and molecular studies suggest that lichen symbioses contain a plethora of associated fungi. These are potential producers of novel bioactive compounds, but strains isolated on standard media usually represent only a minor subset of these fungi. By using various in vitro growth conditions we are able to modulate and extend the fraction o...

Lichens are recognized by macroscopic structures formed by a heterotrophic fungus, the mycobiont, which hosts internal autotrophic photosynthetic algal and/or cyanobacterial partners, referred to as the photobiont. We analyzed structure and functionality of the entire lung lichen Lobaria pulmonaria L. Hoffm. collected from two different sites by st...

Fungi are more important to our lives than is assumed by the general public. They can comprise both devastating pathogens and plant-associated mutualists in nature, and several species have also become important workhorses of biotechnology. Fungal diversity research has in a short time transcended from a low-tech research area to a method-intensive...

Knowledge of bacterial community host-specificity has increased greatly in recent years. However, the inter-microbiome relationships of unrelated but spatially close organisms remain little understood. Trunks of trees covered by epiphytes represent complex habitats with a mosaic of ecological niches. In this context, we investigated the structure,...

Trentepohliaceae are a distinct lineage of green algae frequently found in association with lichen-forming fungi. We review the current knowledge about the phenotypic diversity of this group of algae, which is complemented by currently available molecular data. These data do not confirm the traditional classification of the genera based on morpholo...

Objectives Despite the input of microbiome research, a group of 20 bacteria continues to be the focus of periodontal diagnostics and therapy. The aim of this study was to compare three commercial kits and laboratory-developed primer pairs for effectiveness in detecting such periodontopathogens. Materials and methods Fourteen bacterial mock communit...

Two new compounds, cyaneodimycin (I) and cyaneomycin bearing a rare methacrylate residue, are isolated from cultures of the title bacterium associated with the marine lichen L.

Arthonia parietinaria is described as new to science. Host of the type and at the same time the only confirmed host species is the foliose macrolichen Xanthoria parietina. Sequence data of nucLSU rRNA genes reveal a close relationship to Arthonia molendoi. A. parietinaria is recorded for many countries in Europe, western Asia, and northern Africa.

Grube, M., Obermayer, W., Mayrhofer, H. & Spribille, T. 2016. Josef Hafellner – a life amongst lichens and their parasites. – Herzogia 29: 213 –234.

Lichenized Fungi and the Evolution of Symbiotic Organization, Page 1 of 2 Abstract Lichen symbioses comprise a fascinating relationship between algae and fungi. The lichen symbiotic lifestyle evolved early in the evolution of ascomycetes and is also known from a few basidiomycetes. The ascomycete lineages have diversified in the lichenized stage...

Self-sustaining lichen symbioses potentially can become very old, sometimes even thousands of years in nature. In the joint structures, algal partners are sheltered between fungal structures that are externally colonized by bacterial communities. With this arrangement lichens survive long periods of drought, and lichen thalli can be revitalized eve...

Evolution of the Life/Dead Ratio over time. Each individual plot displays the evolution of the fractions of living and dead bacteria over time. Each plot belongs to one subject and the running number (SXX) is given on the top left corner. Within an individual plot, each data point represents the mean over all fractions of living (top plot) and dead...

Statistical analysis on class level. A p < 0.05 was considered significant after Bonferroni correction.

Analysis of the recorded biofilm and exclusion of suspected yeast cells. The movie shows an entire confocal stack of a life/dead stained biofilm and its corresponding analysis. (A) Living bacteria stained in green. (B) Dead bacteria and suspected yeast cells stained in red. The boundary of the detected foreground of (A,B) is displayed as a green li...

Statistical analysis on order level. A p < 0.05 was considered significant after Bonferroni correction.

Statistical analysis on genus level. A p < 0.05 was considered significant after Bonferroni correction.

Structure of the biofilm. Exemplary images of life/dead stained biofilms with the different observed structures. Each image is the maximum projection of the respective recorded confocal stack. Often observed structures are cocci filaments (A,C) and cocci staples (A,B).

Statistical analysis on family level. A p < 0.05 was considered significant after Bonferroni correction.

Experiments to resynthesize lichens so far focused on co-cultures of fungal and algal partners. However, recent studies have revealed that bacterial communities colonize lichens in a stable and host-specific manner. We were therefore interested in testing how lichenized fungi and algae interact with selected bacteria in an experimental setup. We se...