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Abstract. From extracellular freezing to cloud glaciation, the crystallization of water is ubiquitous and shapes life as we know it. Efficient biological ice nucleators (INs) are crucial for organism survival in cold environments and, when aerosolized, serve as a significant source of atmospheric ice nuclei. Several lichen species have been identified as potent INs capable of inducing freezing at high subzero temperatures. Despite their importance, the abundance and diversity of lichen INs are still not well understood. Here, we investigate ice nucleation activity in the cyanolichen-forming genus Peltigera from across a range of ecosystems in the Arctic, the northwestern United States, and Central and South America. We find strong IN activity in all tested Peltigera species, with ice nucleation temperatures above −12 °C and 35 % of the samples initiating freezing at temperatures at or above −6.2 °C. The Peltigera INs in aqueous extract appear to be resistant to freeze–thaw cycles, suggesting that they can survive dispersal through the atmosphere and thereby potentially influence precipitation patterns. An axenic fungal culture termed L01-tf-B03, from the lichen Peltigera britannica JNU22, displays an ice nucleation temperature of −5.6 °C at 1 mg mL−1 and retains remarkably high IN activity at concentrations as low as 0.1 ng mL−1. Our analysis suggests that the INs released from this fungus in culture are 1000 times more potent than the most active bacterial INs from Pseudomonas syringae. The global distribution of Peltigera lichens, in combination with the IN activity, emphasizes their potential to act as powerful ice-nucleating agents in the atmosphere. 

Abstract PremiseSouthern Africa is a biodiversity hotspot rich in endemic plants and lichen‐forming fungi. However, species‐level data about lichen photobionts in this region are minimal. We focused onTrebouxia(Chlorophyta), the most common lichen photobiont, to understand how southern African species fit into the global biodiversity of this genus and are distributed across biomes and mycobiont partners. MethodsWe sequencedTrebouxianuclear ribosomal ITS andrbcLof 139 lichen thalli from diverse biomes in South Africa and Namibia. GlobalTrebouxiaphylogenies incorporating these new data were inferred with a maximum likelihood approach.Trebouxiabiodiversity, biogeography, and mycobiont–photobiont associations were assessed in phylogenetic and ecological network frameworks. ResultsAn estimated 43 putativeTrebouxiaspecies were found across the region, including seven potentially endemic species. Only five clades represent formally described species:T. arboricolas.l. (A13),T. cf.cretacea(A01),T. incrustata(A06),T. lynniae(A39), andT. maresiae(A46). Potential endemic species were not significantly associated with the Greater Cape Floristic Region or desert.Trebouxiaspecies occurred frequently across multiple biomes. Annual precipitation, but not precipitation seasonality, was significant in explaining variation inTrebouxiacommunities. Consistent with other studies of lichen photobionts, theTrebouxia–mycobiont network had an anti‐nested structure. ConclusionsDepending on the metric used, ca. 20–30% of globalTrebouxiabiodiversity occurs in southern Africa, including many species yet to be described. With a classification scheme forTrebouxianow well established, tree‐based approaches are preferable over “barcode gap” methods for delimiting new species. 

Abstract Peltigera globulata Miadl. & Magain, a new species in the P. ponojensis/monticola species complex of section Peltigera , is formally described. This clade was previously given the interim designation Peltigera sp. 17. It is found in sun-exposed and xeric habitats at high altitudes in Peru and Ecuador. Peltigera globulata can be easily recognized by its irregularly globulated margins covered mostly by thick, white pruina, somewhat resembling the sorediate thallus margins of P. soredians , another South American species from section Peltigera . The hypervariable region of ITS1 (ITS1-HR), which is in general highly variable among species of section Peltigera , does not have diagnostic value for species identification within the P. ponojensis/monticola complex. Nevertheless, no significant level of gene flow was detected among eight lineages representing a clade of putative species (including P. globulata ) within this complex. ITS sequences from the holotype specimens of P. monticola Vitik. (collected in 1979) and P. soredians Vitik. (collected in 1981) and lectotype specimens of P. antarctica C. W. Dodge (collected in 1941) and P. aubertii C. W. Dodge (collected in 1952) were successfully obtained through Sanger and Illumina metagenomic sequencing. BLAST results of these sequences revealed that the type specimen of P. monticola falls within the P. monticola/ponojensis 7 clade, which represents P. monticola s. str., and confirmed that the type specimen of P. aubertii falls within a clade identified previously as P. aubertii based on morphology. The ITS sequence from the type specimen of P. soredians , which superficially resembles P. globulata , confirms its placement in the P. rufescens clade. Finally, we discovered that the name P. antarctica was erroneously applied to a lineage in the P. ponojensis/monticola clade. The ITS sequence from the type specimen of P. antarctica represents a lineage within the P. rufescens clade, which is sister to the P. ponojensis/monticola clade. 

Abstract Prokaryotic genomes are often considered to be mosaics of genes that do not necessarily share the same evolutionary history due to widespread horizontal gene transfers (HGTs). Consequently, representing evolutionary relationships of prokaryotes as bifurcating trees has long been controversial. However, studies reporting conflicts among gene trees derived from phylogenomic data sets have shown that these conflicts can be the result of artifacts or evolutionary processes other than HGT, such as incomplete lineage sorting, low phylogenetic signal, and systematic errors due to substitution model misspecification. Here, we present the results of an extensive exploration of phylogenetic conflicts in the cyanobacterial order Nostocales, for which previous studies have inferred strongly supported conflicting relationships when using different concatenated phylogenomic data sets. We found that most of these conflicts are concentrated in deep clusters of short internodes of the Nostocales phylogeny, where the great majority of individual genes have low resolving power. We then inferred phylogenetic networks to detect HGT events while also accounting for incomplete lineage sorting. Our results indicate that most conflicts among gene trees are likely due to incomplete lineage sorting linked to an ancient rapid radiation, rather than to HGTs. Moreover, the short internodes of this radiation fit the expectations of the anomaly zone, i.e., a region of the tree parameter space where a species tree is discordant with its most likely gene tree. We demonstrated that concatenation of different sets of loci can recover up to 17 distinct and well-supported relationships within the putative anomaly zone of Nostocales, corresponding to the observed conflicts among well-supported trees based on concatenated data sets from previous studies. Our findings highlight the important role of rapid radiations as a potential cause of strongly conflicting phylogenetic relationships when using phylogenomic data sets of bacteria. We propose that polytomies may be the most appropriate phylogenetic representation of these rapid radiations that are part of anomaly zones, especially when all possible genomic markers have been considered to infer these phylogenies. [Anomaly zone; bacteria; horizontal gene transfer; incomplete lineage sorting; Nostocales; phylogenomic conflict; rapid radiation; Rhizonema.] 

Abstract Lichen thalli are formed through the symbiotic association of a filamentous fungus and photosynthetic green alga and/or cyanobacterium. Recent studies have revealed lichens also host highly diverse communities of secondary fungal and bacterial symbionts, yet few studies have examined the viral component within these complex symbioses. Here, we describe viral biodiversity and functions in cyanolichens collected from across North America and Europe. As current machine-learning viral-detection tools are not trained on complex eukaryotic metagenomes, we first developed efficient methods to remove eukaryotic reads prior to viral detection and a custom pipeline to validate viral contigs predicted with three machine-learning methods. Our resulting high-quality viral data illustrate that every cyanolichen thallus contains diverse viruses that are distinct from viruses in other terrestrial ecosystems. In addition to cyanobacteria, predicted viral hosts include other lichen-associated bacterial lineages and algae, although a large fraction of viral contigs had no host prediction. Functional annotation of cyanolichen viral sequences predicts numerous viral-encoded auxiliary metabolic genes (AMGs) involved in amino acid, nucleotide, and carbohydrate metabolism, including AMGs for secondary metabolism (antibiotics and antimicrobials) and fatty acid biosynthesis. Overall, the diversity of cyanolichen AMGs suggests that viruses may alter microbial interactions within these complex symbiotic assemblages. 

Abstract Understanding how species-rich communities persist is a foundational question in ecology. In tropical forests, tree diversity is structured by edaphic factors, climate, and biotic interactions, with seasonality playing an essential role at landscape scales: wetter and less seasonal forests typically harbor higher tree diversity than more seasonal forests. We posited that the abiotic factors shaping tree diversity extend to hyperdiverse symbionts in leaves—fungal endophytes—that influence plant health, function, and resilience to stress. Through surveys in forests across Panama that considered climate, seasonality, and covarying biotic factors, we demonstrate that endophyte richness varies negatively with temperature seasonality. Endophyte community structure and taxonomic composition reflect both temperature seasonality and climate (mean annual temperature and precipitation). Overall our findings highlight the vital role of climate-related factors in shaping the hyperdiversity of these important and little-known symbionts of the trees that, in turn, form the foundations of tropical forest biodiversity. 

Biological nitrogen fixation (BNF) by microorganisms associated with cryptogamic covers, such as cyanolichens and bryophytes, is a primary source of fixed nitrogen in pristine, high-latitude ecosystems. On land, low molybdenum (Mo) availability has been shown to limit BNF by the most common form of nitrogenase (Nase), which requires Mo in its active site. Vanadium (V) and iron-only Nases have been suggested as viable alternatives to countering Mo limitation of BNF; however, field data supporting this long-standing hypothesis have been lacking. Here, we elucidate the contribution of vanadium nitrogenase (V-Nase) to BNF by cyanolichens across a 600-km latitudinal transect in eastern boreal forests of North America. Widespread V-Nase activity was detected (∼15–50% of total BNF rates), with most of the activity found in the northern part of the transect. We observed a 3-fold increase of V-Nase contribution during the 20-wk growing season. By including the contribution of V-Nase to BNF, estimates of new N input by cyanolichens increase by up to 30%. We find that variability in V-based BNF is strongly related to Mo availability, and we identify a Mo threshold of ∼250 ng·g lichen −1 for the onset of V-based BNF. Our results provide compelling ecosystem-scale evidence for the use of the V-Nase as a surrogate enzyme that contributes to BNF when Mo is limiting. Given widespread findings of terrestrial Mo limitation, including the carbon-rich circumboreal belt where global change is most rapid, additional consideration of V-based BNF is required in experimental and modeling studies of terrestrial biogeochemistry. 

Summary Bryophytes harbour microbiomes, including diverse communities of fungi. The molecular mechanisms by which perennial mosses interact with these fungal partners along their senescence gradients are unknown, yet this is an ideal system to study variation in gene expression associated with trophic state transitions. We investigated differentially expressed genes of fungal communities and their hostDicranum scopariumacross its naturally occurring senescence gradient using a metatranscriptomic approach. Higher activity of fungal nutrient‐related (carbon, nitrogen, phosphorus and sulfur) transporters and Carbohydrate‐Active enZyme (CAZy) genes was detected toward the bottom, partially decomposed, layer of the moss. The most prominent variation in the expression levels of fungal nutrient transporters was from inorganic nitrogen‐related transporters, whereas the breakdown of organonitrogens was detected as the most enriched gene ontology term for the hostD. scoparium, for those transcripts having higher expression in the partially decomposed layer. The abundance of bacterial rRNA transcripts suggested that more living members ofCyanobacteriaare associated with the photosynthetic layer ofD. scoparium, while members ofRhizobialesare detected throughout the gametophytes. Plant genes for specific fungal–plant communication, including defense responses, were differentially expressed, suggesting that different genetic pathways are involved in plant‐microbe crosstalk in photosynthetic tissues compared to partially decomposed tissues.