Drylands account for 45% of the Earth’s land area, supporting ~40% of the global population. These regions support some of the most extreme environments on Earth, characterized by extreme temperatures, low and variable rainfall, and low soil fertility. In these biomes, microorganisms provide vital ecosystem services and have evolved distinctive adaptation strategies to endure and flourish in the extreme. However, dryland microbiomes and the ecosystem services they provide are under threat due to intensifying desertification and climate change. In this review, we provide a synthesis of our current understanding of microbial life in drylands, emphasizing the remarkable diversity and adaptations of these communities. We then discuss anthropogenic threats, including the influence of climate change on dryland microbiomes and outline current knowledge gaps. Finally, we propose research priorities to address those gaps and safeguard the sustainability of these fragile biomes.

In an era of rapid climate change and expansion of desertification, the extremely harsh conditions of drylands are a true challenge for microbial life. Under drought conditions, where most life forms cannot survive, rocks represent the main refuge for life. Indeed, the endolithic habitat provides thermal buffering, physical stability, and protection against incident ultraviolet (UV) radiation and solar radiation and, to some extent, ensures water retention to microorganisms. The study of these highly specialized extreme-tolerant and extremophiles may provide tools for understanding microbial interactions and processes that allow them to keep their metabolic machinery active under conditions of dryness and oligotrophy that are typically incompatible with active life, up to the dry limits for life. Despite lithobiontic communities being studied all over the world, a comprehensive understanding of their ecology, evolution, and adaptation is still nascent. Herein, we survey the fungal component of these microbial ecosystems. We first provide an overview of the main defined groups (i.e., lichen-forming fungi, black fungi, and yeasts) of the most known and studied Antarctic endolithic communities that are almost the only life forms ensuring ecosystem functionality in the ice-free areas of the continent. For each group, we discuss their main traits and their diversity. Then, we focus on the fungal taxonomy and ecology of other worldwide endolithic communities. Finally, we highlight the utmost importance of a global rock survey in order to have a comprehensive view of the diversity, distribution, and functionality of these fungi in drylands, to obtain tools in desert area management, and as early alarm systems to climate change.

Asexually reproducing fungi play a significant role in essential processes in managed and wild ecosystems such as nutrients cycling and multitrophic interactions. A large number of such taxa are among the most notorious plant and animal pathogens. In addition, they have a key role in food production, biotechnology and medicine. Taxa without or rare sexual reproduction are distinguished based on their sporulating structures and conidiomata in traditional morphology-based taxonomy. The number, variation and diversity of asexually reproducing taxa are insufficiently known, even though fungi capable of asexual reproduction may provide an untapped, rich biological resource for future exploitation. Currently, ca. 30,000 asexual species belonging to ca. 3800 genera have been reported (including 1388 coelomycetous and 2265 hyphomycetous genera). Recent reports (2017–2020) reiterate that the number of asexually producing fungi is higher than the number of frequently sexually-reproducing fungi. With the advent of molecular tools and the abandonment of the dual nomenclature system for pleomorphic fungi, priority criteria were established and revisited in the latest outline of fungi and fungus-like taxa. However, species numbers and taxonomic boundaries of pleomorphic taxa and their synanamorphs or synasexual morphs have yet to be addressed. The number of species of speciose genera (e.g. Alternaria, Aspergillus, Cercospora, Fusarium, Phoma and Pseudocercospora), cryptic species, species of pleomorphic genera, less studied life modes (such as lichenicolous taxa, taxa from extreme environments) and species from biodiversity-rich areas still need evaluation to achieve more reliable estimates of their diversity. This paper discusses the current knowledge on the matter, with diversity estimates, and potential obstacles in several chapters on (1) speciose genera; (2) pleomorphic genera; (3) cryptic species; (4) well-studied but insufficiently resolved taxa, e.g. leaf inhabiting species, marine fungi, (5) less studied life modes, e.g. lichenicolous, rock-inhabiting fungi, insect-associated and yeast-forming taxa and (6) species from biodiversity-rich areas.