# Demystifying fungal systematics: A gateway to fungal literacy and societal/ecological relevance through familiar species

Anna Vaiana, Francis Martin, Rasmus Kjøller, Toby Spribille, Garima Singh

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Anna Vaiana, Francis Martin, Rasmus Kjøller, Toby Spribille, Garima Singh. Demystifying fungal systematics: A gateway to fungal literacy and societal/ecological relevance through familiar species. Plants, People, Planet, 2025, 8 (2), pp.499-515. 10.1002/ppp3.70119 . hal-05525848

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# Demystifying fungal systematics: A gateway to fungal literacy and societal/ecological relevance through familiar species

Anna Vaiana<fnref n="1" /> ID | Francis Martin<fnref n="2" />,<fnref n="3" /> ID | Rasmus Kjøller<fnref n="4" /> ID | Toby Spribille<fnref n="5" /> ID | Garima Singh<fnref n="1" />,<fnref n="6" />,<fnref n="7" /> ID

Correspondence

Garima Singh, National Biodiversity Future Center (NBFC), Piazza Marina, 61, 90133 Palermo, Italy.

Email: garima.singh@unipd.it

Funding information

European Union - Next-Generation EU: 'PNRR Missione 4 Componente 2, "Dalla ricerca all'impresa", Investimento 1.4, Grant/Award Number: CN00000033

Societal Impact Statement

Fungal systematics can feel overwhelming given the vast species diversity within this kingdom, with numerous subgroups at every taxonomic rank. This often creates a disconnect between the general undersanding of fungal taxonomy and societal relevance and recognition of fungi. Here we anchor well-known fungi onto the taxonomic tree, using them as signposts to navigate fungal taxonomy. By presenting taxonomic ranks in relation to fungi encountered in daily life, we make the system more relatable and engaging for students, researchers, and the broader public. This approach provides both an evolutionary roadmap and an accessible guide, raising awareness of these essential yet often overlooked life forms.

Summary

Fungal systematics can feel difficult to navigate because of the high species diversity and several subgroups in each taxonomic rank. Although classification systems aim to simplify and organize this diversity, the number of subgroups/divisions often presents challenges for students and researchers. This review introduces commonly observed and well-known fungal species, including mushrooms, molds, yeasts, lichens, and fungi used in food production and medicine, as reference points to explore major clades within the fungal tree of life. Our selection spans fungi of ecological, medical, and industrial relevance, offering a tangible reference for understanding where different groups are placed within the fungal kingdom. We highlighted the position of famous fungi on a fungal tree of life, simultaneously highlighting the ecological, industrial, and medical relevance of each clade. This approach links well-known examples to broader patterns in fungal evolution and taxonomy, providing a structured overview of the fungi we have heard of or encountered in our lives. Integrating taxonomy with familiar real-world examples offers a more accessible entry point into fungal biology. This resource is intended for educational and introductory systematic purposes and supports a clearer understanding of fungal diversity and classification. By linking taxonomic groups with commonly encountered fungi, this paper aims to raise awareness of fungi and promote a

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This is an open access article under the terms of the [Creative Commons Attribution License](https://creativecommons.org/licenses/by/4.0/), which permits use, distribution and reproduction in any medium, provided the original work is properly cited.

© 2025 The Author(s). *Plants, People, Planet* published by John Wiley & Sons Ltd on behalf of New Phytologist Foundation.

<footnote n="1">¹Department of Biology, University of Padova, Padova, Italy</footnote>

<footnote n="2">²Centre INRAE Grand-Est Nancy, Université de Lorraine, INRAE, UMR Interactions Arbre/Micro-Organismes, Champenoux, France</footnote>

<footnote n="3">³The National Key Laboratory of Ecological Security and Sustainable Development in the Arid Region, Northwest Institute of Eco-Environment and Resources, Chinese Academy of Sciences, Lanzhou, China</footnote>

<footnote n="4">⁴Department of Biology, University of Copenhagen, Copenhagen, Denmark</footnote>

<footnote n="5">⁵University of Alberta, Edmonton, AB, Canada</footnote>

<footnote n="6">⁶National Biodiversity Future Center (NBFC), Palermo, Italy</footnote>

<footnote n="7">⁷Botanical Garden of Padua, University of Padova, Padova, Italy</footnote>
more integrated understanding of fungal evolution and its significance across biologi-
cal disciplines.

KEYWORDS

Ascomycota, *Fusarium*, lichens, molds, mushrooms, *Penicillium*, yeast

1 | INTRODUCTION: A RELATABLE JOURNEY INTO KINGDOM FUNGI

Fungi are a diverse and essential group of organisms that play crucial roles in ecosystems, industries, medicine, and our culture (Aly et al., 2011; Spiteller, 2015). The fungal kingdom encompasses a vast and diverse array of organisms, including molds, mushrooms, yeasts, lichens, rusts, smuts, and mycorrhizal fungi, many of which play essential ecological roles across nearly all ecosystems (Hawksworth & Lücking, 2017). Approximately one billion years ago, a group of aquatic, unicellular eukaryotes that produced sporangia with single, posteriorly flagellated zoospores split into two evolutionary lineages: one leading to animals and the other to fungi (Stajich et al., 2009). Since then, fungi have evolved in parallel with other major eukaryotic groups, such as animals and plants, diversifying across a wide range of ecological niches over hundreds of millions of years.

Overall, fungi exhibit a wide variety of lifestyles, ranging from microscopic forms such as molds and yeasts to larger forms like mushrooms and many lichens. Currently, there are about 155,000 known species of fungi; however, according to the *State of the World's Plants and Fungi 2023* report, global fungal diversity estimates this number to be approximately 2.5 million, suggesting fungi as the second largest kingdom of eukaryotes after animals. The remarkable morphological and functional diversity of fungi makes it challenging to categorize and communicate this complexity, particularly among beginners or within educational settings, even to early career researchers and academics. Some of the unique aspects of fungal taxonomy are cryptic morphology, a limited number of taxonomically informative characters, frequent revisions, the abundance of undescribed or unculturable species, and the subdivision of taxa into multiple groups at each rank. Together, these factors can make fungal systematics particularly daunting for readers encountering the field for the first time. This has contributed to the disconnect between fungal taxonomy and fungal awareness. Bridging this gap is essential, as different fungi profoundly influence different aspects of our lives, from causing diseases to producing life-saving medicines, shaping our food systems, supporting ecological resilience, and sustaining biodiversity. Raising awareness of their roles and diversity is therefore not only scientifically important but also socially and environmentally relevant. With this article, our goal is to bridge this gap by demystifying fungal taxonomy and linking major taxonomic ranks to well-known fungi that people may encounter in daily life, research, or broader cultural contexts.

Before presenting the overview of the phylogeny and our refer-
ence framework anchored in iconic fungi, we want to acknowledge
that fungal taxonomy is, and will continue to be, in a state of dynamic

transformation, driven by increasingly robust, integrative species iden-
tification methodologies, and a growing commitment to uncovering
the full scale of fungal diversity. Moving beyond industrially, medi-
cally, or ecologically relevant taxa, contemporary research actively
pursues the systematic exploration of unrecognized lineages. Such
advances are refining our understanding of species relationships and
may necessitate the addition of new taxonomic ranks, reflected in the
addition of new subclasses, orders, and families. Nonetheless, we
assert that even within a fluid classification framework, extensively
studied model and iconic fungi provide stable reference points for
both newcomers and interdisciplinary researchers, helping them famil-
iarize themselves with the current state of the art and facilitating the
integration of new discoveries and future changes. For instance, while
the taxonomic positions and genus names of nonmodel fungi are fre-
quently subjected to taxonomic revisions, model fungi such as Saccha-
romyces (e.g., Saccharomyces cerevisiae), Neurospora (e.g., Neurospora
crassa), Aspergillus, Penicillium, and Talaromyces have largely retained
their genus names and broad classifications. Their relatively stable tax-
onomic position provides both historical continuity and a reliable ref-
erence framework for interpreting future developments in fungal
systematics.

Another recurring challenge in fungal systematics is the discor-
dance among phylogenetic frameworks. As methodologies and data-
sets continue to evolve, species relationships are frequently revised,
and researchers may rely on slightly different, contemporaneously
published classifications when describing or referencing taxa. For
newcomers to the field, this variability can be a source of confusion,
particularly when encountering inconsistent numbers of phyla or con-
flicting lineage placements across sources. That said, the field has
moved toward greater cohesion in recent years. A global consortium
of mycologists now regularly publishes updated outlines of fungi and
fungus-like taxa. These collaborative efforts integrate the latest litera-
ture and revise taxonomic ranks as needed, offering a more unified
and consensus-based framework (Hyde, 2024; Li et al., 2021; Nagy &
Szöllősi, 2017; Rudramurthy & Kaur, 2024; Tedersoo et al., 2018). In
this manuscript, we adopt and refer to this framework as the founda-
tion for our exploration of fungal diversity.

Currently, nine major phylum-level clades are recognized within
the fungal kingdom: Opisthosporidia, Chytridiomycota, Neocallimasti-
gomycota, Blastocladiomycota, Zoopagomycota, Mucoromycota,
Glomeromycota, Basidiomycota, and Ascomycota (Gilando et al.
2021; Naranjo-Ortiz & Gabaldón, 2019). Among these, the phyla
Basidiomycota (e.g., mushrooms, rusts, and smuts) and Ascomycota
(e.g. yeasts, molds, truffles and morels) comprise 98% of all currently
known fungi (He et al., 2022; Lücking et al., 2017). In addition to these
**FIGURE 1** Schematic phylogenetic tree representing the dataset used in this study, illustrating the taxonomic coverage across the nine fungal phyla. Classes containing the most “famous fungi” discussed in this work are marked with stars, along with their respective phyla. This schematic tree is intended as an illustrative overview (sensu Galindo et al., 2021) and does not reflect precise phylogenetic distances or evolutionary timelines.

<figure><img src="image_4.png" /></figure>

well-known Dikarya, the five early-diverging phyla, Opisthosporidia, Chytridiomycota, Neocallimastigomycota, Blastocladiomycota, and Zoopagomycota, remain comparatively understudied, largely due to their specialized ecologies, nonstandard morphologies, and challenges in cultivation. Opisthosporidia are obligate intracellular parasites of various eukaryotic phytoplankton (Karpov et al., 2017). Chytridiomycota comprise mostly microscopic saprotrophs or parasites inhabiting freshwater, marine environments, soils, and the digestive tracts of herbivores (Longcore, 2006). Neocallimastigomycota are strictly anaerobic fungi residing in the guts of herbivores, where they aid in plant biomass degradation (Hanafy et al., 2022). Blastocladiomycota are uniflagellated, zoospore-producing fungi that function as saprotrophs and parasites, mainly in freshwater and soil (Powell, 2017). Zoopagomycota include entomopathogenic fungi, insect commensals, putative saprotrophs, mycoparasites, and parasites of microinvertebrates (Reynolds, 2021). Although almost all phyla contain some

famous fungi, the fungi which we selected for discussion in this article belong to the phyla Mucoromycota, Glomeromycota, Basidiomycota, and Ascomycota (Figure 1) for the reasons stated below.

Glomeromycota comprises fungi that form arbuscular mycorrhizal (AM) symbioses with most terrestrial plants. Most plant species engage in mycorrhizal associations, with AM fungi (AMF) representing the most abundant and widespread type. These symbioses enhance crop growth and soil health, making them vital to agriculture and ecosystems. Mucoromycota, on the other hand, includes species involved in food fermentation (e.g., *Rhizopus* in tempeh), opportunistic pathogens causing mucormycosis, and root symbionts such as *Mucoromycotina* "fine root endophytes" (MFREs) (Howard et al., 2022; Smith & Read, 2010; Soudzilovskaia et al., 2020).

Phylum Basidiomycota is the second most speciose fungal phylum and contains approximately 32,000 well-known fungi including mushrooms, encompassing a wide range of life strategies and structural
complexities (He et al., 2022). This phylum comprises club fungi with club-shaped spore-producing structures, called basidia. The typical fruiting bodies of this phylum are mushrooms, puffballs, brackets, and jelly fungi. The fruiting body, for instance, the visible mushroom, is called a basidiocarp, and it contains millions of microscopic basidia. Members of this phylum include some of the most morphologically complex fungi, particularly those with intricate life cycles involving multiple developmental stages (e.g., *Puccinia*) and extensive multicellular structures (e.g., *Armillaria*).

Many fungi of great medicinal and industrial relevance belong to the phylum Ascomycota, which is also the largest fungal phylum, comprising approximately 64,000 described species (Hawksworth & Lücking, 2017; Hibbett, 2007; Schoch et al., 2009). Ascomycota are sac fungi, characterized by the production of spores in sac-like structures called asci. The taxonomic diversity and experimental tractability of these fungi provided numerous model systems for fungal genetics and molecular biology studies, with notable examples being *Penicillium* spp., *Candida* spp., *Aspergillus* spp., and *S. cerevisiae* (yeast) (Bhunjun et al., 2024).

Taxonomically, both Ascomycota and Basidiomycota are divided into various classes, subclasses, orders, and families, based on morphological characteristics and increasingly supported by molecular phylogenetic analyses that clarify their evolutionary relationships (Galindo et al. 2021; Lücking et al., 2017). However, the taxonomy of Ascomycota is more complex, given the species richness of this phylum as compared to Basidiomycota. For instance, Basidiomycota is currently classified into 19 classes and 273 families, whereas Ascomycota includes 23 classes and 686 families, respectively (He et al., 2024). Although these taxonomic ranks aim to organize the vast diversity of currently described fungi into a manageable system, the sheer complexity and number of taxa can make navigation challenging.

While all organisms on Earth influence our lives in one way or another, we think fungi deserve special attention due to their vast complexity and their profound impact on society, from their roles in industry, medicine, and ecosystems in general (Niego et al., 2023). Fungi such as *Penicillium* sp. (the source of penicillin) and *S. cerevisiae* (baker's yeast) have contributed significantly to advances in medicine and have been central to several Nobel Prize-winning discoveries, alongside model organisms like the mouse (*Mus musculus*), the fruit fly (*Drosophila melanogaster*), and the gut bacterium *Escherichia coli*. Overall, fungi affect various aspects of our lives, with some providing essential benefits, others posing threats to human health and agriculture. Beyond classification, our aim is to make fungal education more accessible, interdisciplinary, and engaging in bridging taxonomy, ecology, and applied sciences. This article is intended for those looking to familiarize themselves with this vast diversity for systematic and taxonomic purposes or for educational applications.

We recognize that fungal taxonomy is, and will continue to be, in a state of dynamic transformation, driven by increasingly robust, integrative species identification methodologies and a growing commitment to uncovering the full scope of fungal diversity. Moving beyond industrially, medically, or ecologically relevant taxa, contemporary research actively pursues the systematic exploration of unrecognized

lineages. Such advances are refining our understanding of species relationships and may necessitate the addition of new taxonomic ranks, reflected in the addition of new subclasses, orders, and families. Nonetheless, we assert that even within a fluid classification framework, extensively studied model and iconic fungi provide stable reference points for both newcomers and interdisciplinary researchers, helping them familiarize themselves with the current state of the art and facilitating the integration of new discoveries and future changes.

With this aim, we highlight the most well-known fungi in fungal phylogeny and have selected the following six categories (Figure 2) based on their impact on human lives:

* Fungi Around Us: Unseen or Unnoticed but Ever-Present

* From Forests to Tables and Toasts

* Myco Mayhem: The Dark Side of Fungi

* Mind-Bending Mushrooms: Fungi That Twist our Senses

* Ecologically Important Fungi

* The Fungal Pharmacy

## 2 | FUNGI AROUND US: UNSEEN OR UNNOTICED BUT EVER-PRESENT

Molds are fungi that are commonly found in indoor environments, including our homes. Taxonomically, molds encompass many fungal species across different genera. Some of the most well-known fungi that form molds include species from the genera *Aspergillus*, *Penicillium*, *Cladosporium*, *Botrytis*, *Stachybotrys*, *Fusarium*, and *Rhizopus*.

Molds play important roles in decomposition, food production, and biotechnology. However, certain species can be harmful to plants, animals, and humans by causing diseases or by producing toxic compounds known as mycotoxins. These secondary metabolites are generated by fungi that grow on various food sources consumed by both humans and animals. To date, over 400 different types of mycotoxins have been identified (Kuhn & Ghannoum, 2003).

One of the best studied common molds is *Stachybotrys chartarum* (Sordariomycetes, Ascomycota; Figure 2), often referred to as black mold and frequently associated with buildings with damp and humid walls. Concerns regarding the potential health effects of exposure to this mold have led to extensive research on moisture-related mold damage, with some studies linking it to severe illnesses (Dylag et al., 2022; Kuhn & Ghannoum, 2003).

Another common mold is the fuzzy grey mold *Botrytis cinerea* (Leotiomycetes, Ascomycota; Figure 2), which affects a wide range of crops, including nursery plants, vegetables, ornamental plants, fruits, and legumes (Elad et al., 2007). *Botrytis* species cause diseases in greenhouses, agricultural fields, storage facilities, and during transportation among plant-related facilities, posing a major challenge to the agricultural industry (Carisse, 2016; Elad et al., 2007). Notably, as the noble rot, it is also involved in the production of highly prized sweet wines.
<figure><img src="image_5.png" /><figcaption>**FIGURE 2** Six key categories capturing the impact of fungi on society and nature. We present familiar fungi, including mushrooms, molds, and notable pathogenic species, organized into these categories, while locating them on the fungal tree of life to illustrate their taxonomic positions and to familiarize readers with fungal taxonomic ranks through recognizable examples.</figcaption></figure>

Among the bread molds, *Rhizopus* species *R. stolonifer* is particularly significant. Belonging to the Mucoromycota phylum, this fungus is one of the most common bread molds and is also a major pathogen affecting postharvest storage, especially fruit (Liu et al., 2024).

Another group of fungi commonly found around us are fungi existing in symbiotic associations with algae and other associated microbiota, forming a composite organism called lichens (Hawksworth & Grube, 2020). These organisms are commonly found in both pristine natural habitats and familiar urban areas like our own backyards on tree barks and rocks. Lichens are commonly seen on statues, tombstones, metal and wood fences, discarded cars, and house roofs, appearing as colorful patches or spots, often resembling rust. Their

colors can vary widely, from orange and yellow to green, grey, and many others, depending on the species and the metabolites contained within the thallus (Figure 3a). Lichen symbiosis has evolved multiple times throughout history, with most lichenized fungi belonging to the phylum Ascomycota, while a smaller percentage is found in Basidiomycota (Lücking et al., 2017). Some of the most common genera of lichenized fungi include *Cladonia*, *Parmelia*, *Physcia*, *Usnea*, and *Xanthoria*. Lichens are remarkable for their longevity, production of unique secondary metabolites of pharmacological interest (Singh et al., 2025), their ability to thrive in environments humans would consider extreme, and high sensitivity to certain pollutants, making them valuable bioindicators (Lutzoni & Miadlikowska, 2009).
<figure><img src="image_6.png" /></figure>

**FIGURE 3** Fungi around us: Representative phylogeny of selected *Ascomycota* genera discussed in the text, annotated with their corresponding taxonomic classes. Photographed species are highlighted in the tree. Taxa from the "Fungi Around Us: Unseen or Unnoticed but Ever-Present" section are marked in blue, while those from the "From Forests to Tables and Toasts" section are shown in green. Source: Hyde, 2024.

<figure><img src="image_7.png" /><figcaption>a) Parmelia spp., Xanthoria parietina</figcaption></figure>

<figure><img src="image_8.png" /><figcaption>b) Morchella sextelata</figcaption></figure>

<figure><img src="image_9.png" /><figcaption>c) Tuber melanosporum</figcaption></figure>

# 3 | FROM FORESTS TO TABLES AND TOASTS

## 3.1 | Edible mushrooms

Mushrooms likely top the list of the most well-known fungi, especially when considering the perception of fungi for people, thanks to their culinary value and industrial significance. The mushroom itself is the fruiting body, the reproductive structure that appears above ground during certain seasons. Much like flowers or fruits on a plant, these structures are temporary and visible only at specific times, while the main part of the organism, the mycelium, remains hidden beneath the surface throughout the year. Below the mushroom, an expansive network of mycelia grows through the substrate, supplying the nutrients and water necessary for fruiting body development. Mushrooms have been consumed and valued for their flavor, economic significance, nutritional value, and medicinal properties for centuries (Sánchez, 2004).

Majority of the edible mushrooms belongs to the phylum Basidiomycota, primarily in the class Agaricomycetes (Figure 4), for example, Agaricus species. Edible mushrooms within Ascomycota belong to the class Pezizomycetes particularly within the order Pezizales, for example, truffles (Tuber spp.) and morels (Morchella spp.) (O'Donnell et al., 1997) (Figure 3). Only a few edible fungi exist outside Dikarya, such as the Rhizopus species in Mucoromycota, but mainly as fermentation agents.

The most widely cultivated mushrooms worldwide include Agaricus bisporus (sold as button mushrooms when small and portobello mushrooms when mature) (Ambhore et al., 2024), Agaricus campestris (field mushroom) (Figure 4a), Lentinus edodes (shiitake), Pleurotus spp. (oyster mushrooms), Auricularia auricula (wood ear mushroom), Flammulina velutipes (winter mushroom), and Volvariella volvacea (straw mushroom) (Valverde et al., 2015; Ambhore et al., 2024; Sharma et al., 2024). Recent studies indicate that Lentinus edodes is now the world's leading cultivated edible mushroom, representing approximately 22% of global production, with China being the main producer (Royse et al., 2017).
<figure><img src="image_10.png" /><figcaption>**FIGURE 4** From forests to tables and toasts (green) and fungal fury (red): Representative phylogeny of selected genera from the phyla Basidiomycota, Mucoromycota, and Glomeromycota, annotated with their respective taxonomic classifications. Photographed species are highlighted on the tree. Taxa featured in the “From Forests to Tables and Toasts” section are shown in green, while those from the “Fungal Fury: Toxic Fungi” section are indicated in red. Source: He et al., 2022, 2024.</figcaption></figure>

Other well-known edible fungi include milk-cap mushrooms (Laccarius deliciosus), morels (Morchella esculenta), porcini mushrooms (Boletus edulis), and truffles, which are considered a delicacy worldwide (Figure 3). Truffles belong to the genus Tuber, one of the few ectomycorrhizal Ascomycetes, and include black truffles (Tuber melanosporum), white truffles (Tuber magnatum), summer truffles (Tuber aestivum), Burgundy truffles (Tuber uncinatum), and Chinese truffles (Tuber indicum) (Figure 3c). Unlike saprotrophic species, ectomycorrhizal fungi form symbiotic relationships with plant roots, and some lineages are notoriously difficult to cultivate under artificial conditions. This ecological distinction not only affects cultivation practices but also contributes to the rarity and high market value of certain edible fungi (Graziosi et al., 2022; Mello et al., 2006; Stobbe et al., 2013; Zambonelli et al., 2015).

## 3.2 | Fungi used in fermentation

Fungi also play an essential role in fermentation and baking: microorganisms such as S. cerevisiae (also known as “Baker’s yeast” or “Brewer’s yeast”) and Penicillium spp., both belonging to Ascomycota, can transform basic ingredients into complex food and beverage products, including bread, beer, wine, and cheese thanks to their fermentative metabolism (Parapouli et al., 2020).

In bread-making, S. cerevisiae converts glucose and fructose from more complex carbohydrates, such as sucrose, maltose, and starch, into carbon dioxide and ethanol (Kong et al., 2018; Parapouli et al., 2020). Fermentation of carbon-based sources by S. cerevisiae for alcoholic beverage production is one of the oldest and most economically significant biotechnological processes. This yeast plays a key role in the production of various fermented and distilled beverages, such as wine, beer, cider, rum, vodka, whiskey, brandy, and sake (Eliodório et al., 2019; Kong et al., 2018).

The close association between humans and S. cerevisiae is further supported by a study that analyzed the genetic diversity of over 600 yeast strains from 56 geographic locations worldwide (Legras et al., 2007). This study highlights how numerous S. cerevisiae strains are closely associated with human activities such as baking, brewing, and winemaking—practices that have been integral to human culture for millennia. These findings suggest that through sustained use and selection, humans have played a significant role in shaping the evolutionary trajectory and genetic landscape of S. cerevisiae, turning it into one of the earliest examples of microbial domestication (Legras et al., 2007).

In winemaking, S. cerevisiae ferments grape juice sugars to generate ethanol and carbon dioxide. The fermentation process is complex and involves numerous biosynthetic pathways, regulated by specific genes associated with sugar metabolism, stress responses, and
fermentation efficiency. It is influenced by multiple parameters, including the composition of the fermentation medium, environmental conditions, and the yeast inoculum (Contreras-Ruiz et al., 2024; Eldarov et al., 2016). Both the specific yeast strain inoculated and fermentation conditions play an important role in shaping the characteristics of wine, primarily through the production of secondary metabolites such as esters, which contribute to aroma and flavor (Eldarov et al., 2016; Kong et al., 2018). However, during the initial stages of spontaneous fermentation, various non-Saccharomyces yeast species, including *Candida*, *Cryptococcus*, *Debaryomyces*, *Hanseniaspora* (and its asexual form *Kloeckera*), *Kluyveromyces*, *Metschnikowia*, *Pichia*, *Rhodotorula*, *Saccharomycodes*, *Schizosaccharomyces*, and *Zygosaccharomyces*, can influence the fermentation dynamics (Escribano-Viana et al., 2018; Jolly et al., 2014).

In beer production, *S. cerevisiae* ferments the sugars present in malted barley, producing ethanol and carbon dioxide. Yeast strain and wort composition play a major role in defining beer characteristics (Eliodório et al., 2019; Legras et al., 2007). Additionally, key factors such as flocculation, wort fermentation efficiency (the process of fermenting the sugar-rich liquid extracted from malted grains, known as wort, including the uptake of sugars, amino acids, and peptides), ethanol and osmotic stress tolerance, and oxygen requirements significantly influence the fermentation process. Natural and industrial hybrids, such as those between *S. cerevisiae* and *S. eubayanus*, have given rise to diverse brewing strains, including lager yeasts, demonstrating how hybridization has contributed to the evolution of yeast metabolism and the broad spectrum of beer styles and flavor profiles (González et al., 2008; Krogerus et al., 2017; Peris et al., 2018).

*S. cerevisiae* has also been employed in various important biotechnological processes, including the production of biofuels, pharmaceuticals, and industrial enzymes, due to its well-characterized genetics and fermentation capabilities (Maneira et al., 2025; Parapouli et al., 2020; Tippmann et al., 2013; Turner et al., 2018).

## 3.3 Fungi in cheese and other fermented food

*Penicillium* spp. are involved in the production of various cheeses, particularly blue cheeses such as Roquefort, Gorgonzola, and Stilton and soft cheeses such as Camembert and Brie (Cleere et al., 2024; Fröhlich-Wyder et al., 2019; Ropars et al., 2020). Bacteria are used to first acidify milk and make it suitable for fungal growth, after which *Penicillium* filamentous hyphae develop during ripening. In most cheeses, the fungal growth is restricted to the surface. This is the case for *Penicillium candidum* (P. camembertii), used in soft cheese production, which forms a white rind while breaking down fats, contributing to the cheese's characteristic texture and mild flavor. In contrast, *Penicillium roqueforti* grows throughout the interior of the cheese and sporulates along air channels introduced during piercing, producing the characteristic blue "veins" and contributing to the cheese's strong taste by breaking down fats and proteins (Cleere et al., 2024).

Well-known examples of fungi used for food production and processing in Asian cuisine include *Aspergillus*-fermented products such

as miso and soy sauce, and *Rhizopus*-fermented products such as tempeh. Soy sauce is produced by fermenting a mixture of cooked soybeans and roasted wheat with *Aspergillus oryzae* to create Koji, which is then further processed to extract the liquid product (Archer, 2000; Dunn-Coleman et al., 1991). Miso, for its part, is made from cooked soyabeans mixed with grains as steamed polished rice or barley, inoculated with *A. oryzae*. Similarly, fermenting partially cooked soybean cotyledons with *Rhizopus oligosporus* (Mucoromycota) result in Indonesian tempeh (Mei Feng et al., 2007). It is worth mentioning that both *Saccharomyces* and *non-Saccharomyces* yeasts can be used to improve the aroma of chocolate and coffee through nonalcoholic fermentation, and *Monascus purpureus* can be used to produce the characteristic pigments and ethanol used in red rice wine and food coloring (Niego et al., 2023; Singh et al., 2021).

## 4 | MYCO MAYHEM: THE DARK SIDE OF FUNGI

Fungi are essential to ecosystems, but they also have a "dark side" as some fungi harm humans, animals, or plants. Fungi can negatively impact us in several ways, including causing diseases in humans and animals, producing deadly mycotoxins, damaging crops and food supplies, and contributing to environmental and structural degradation. Raising awareness about fungal diversity is essential for a deeper understanding of their ecology and evolution. Such knowledge is crucial for developing effective strategies to mitigate their harmful impacts and to foster more sustainable ways of coexisting with these remarkable organisms. Here are a few ways fungi show their darker side, at least from a human point of view.

### 4.1 | Pathogens and parasites

#### 4.1.1 | Plant pathogens

Certain fungi have harmful effects, as they are responsible for various diseases affecting crops, fruits, and other economically important plants. These fungal infections cause significant damage and lead to substantial economic losses. Fungal plant pathogens predominantly belong to phyla Ascomycota and Basidiomycota. Plant pathogenic fungi are distributed across various classes in Ascomycota, including Dothideomycetes (e.g., *Cladosporium* spp.), Sordariomycetes (e.g., *Magnaporthe* spp.), and Leotiomycetes (e.g., *Botrytis* spp.). In Basidiomycota, the most significant plant pathogens are rust fungi (Pucciniomycetes) and smut fungi (Ustilaginomycotina) (Doehlemann et al., 2017).

Throughout history, fungal diseases have had devastating impacts on agriculture, leading to famines and mass migration. A major historical event was the Bengal Rice Famine of 1942–1943, caused by Rice Blast (*Magnaporthe oryzae*) and Brown Spot Disease (*Cochliobolus miyabeanus*). Fungal infections had led to widespread crop failure, causing millions of deaths owing to starvation, making it one of the
deadliest agricultural disasters in history (Greenough, 1980; Nalley et al., 2016; Shamim et al., 2024).

During the Middle Ages, ergotism outbreaks caused widespread mortality and social upheaval across Europe. Called "St. Anthony's Fire," the disease came from eating rye bread contaminated with *Claviceps purpurea* fungus. Ergot is an overgrown grain caryopsis (a type of dry fruit where the seed is fused to the fruit wall) containing toxic alkaloids, developing through parasitic fungal mycelium that produces black or purple spores in early spring. The dramatic effects of ergotism, which killed thousands, significantly influenced medieval art, literature, and religious practices, while recurring epidemics shaped both agricultural methods and community responses (Grzybowski et al., 2021).

Some of the most impactful fungal diseases in agriculture and forestry include smut disease (Ustilago spp., Tilletia spp.), rust diseases (Puccinia spp.), damping-off diseases (Rhizoctonia, Fusarium), apple scab (Venturia inaequalis), brown rot of stone fruits (Monilinia fructicola), red rot of sugarcane (Colletotrichum falcatum), black arm, wilt, and root rot of cotton (Fusarium oxysporum and Macrophomina phaseolina). Beyond cultivated crops, nonagricultural fungal pathogens like Hymenoscyphus fraxineus (ash dieback) and Ophiostoma spp. (Dutch elm disease) have caused extensive damage to tree species in natural and managed environments (Combes et al., 2024; Jürisoo et al., 2025). In addition to pathogenic fungi, some species, such as Serpula lacrymans, commonly referred to as dry rot fungi, cause significant structural damage by decomposing dead wood in buildings and destroying timber and lumber (Schilling et al., 2021). While not pathogens in the strict sense, these wood-decaying fungi are nonetheless ecologically and economically destructive.

## 4.1.2 | Human and animal pathogens

Fungal infections, although less common than bacterial and viral diseases, represent a growing public health concern, especially due to increasing antifungal resistance and limited treatment options. In 2022, the World Health Organization (WHO) published the Fungal Priority Pathogen List (FPPL) to draw attention to the growing impact of fungal diseases, which are estimated to cause approximately 6.5 million cases of invasive infections and around 3.8 million deaths globally each year (Denning, 2024; van Rhijn et al., 2024). These infections primarily affect immunocompromised individuals and are becoming increasingly difficult to treat due to rising antifungal resistance. While climate change has been proposed as a contributing factor, the increasing burden of fungal disease is also directly associated with medical advances that extend the lives of vulnerable patients, as well as changes in agricultural practices and land use. These trends underscore the relevance of a One Health perspective, which emphasizes the interconnectedness of human, animal, and environmental health (CDC, https://www.cdc.gov/fungal/about/one-health.html).

Superficial fungal infections, defined as infections restricted to the stratum corneum with little tissue reaction (Gupta et al., 2017), affect hundreds of millions globally. Common manifestations include Tinea pedis (athlete's foot) and Tinea cruris (jock itch), caused by

*Trichophyton rubrum* and *T. mentagrophytes*, leading to itching and scaling between the toes or groin area (Garber, 2001; Gupta et al., 2017; Piérard et al., 1996). Onychomycosis (nail infections) represents one of the most prevalent fungal conditions worldwide, with higher occurrence in elderly and diabetic individuals (Garber, 2001). Tinea capitis (scalp ringworm), caused by *T. tonsurans* and *Microsporum audouinii*, disproportionately affects children in urban communities, resulting in hair loss, scaling, and social stigma (Hay et al., 1996). Tinea versicolor, caused by *Malassezia furfur*, presents as scaly, pigmented lesions and is particularly common in tropical and subtropical regions (Marcon & Powell, 1987).

Subcutaneous fungal infections penetrate deeper tissues, with conditions like Sporotrichosis (*Sporothrix schenckii*) and Chromoblastomycosis (*Dematiaceae*) fungi particularly affecting agricultural workers and those in contact with contaminated soil (Sharkey-Mathis et al., 1993).

Systemic infections pose the greatest threat, particularly to immunocompromised populations. *Candida* species cause approximately 1.6 million cases of bloodstream infections or invasive candidiasis globally each year, with an estimated mortality rate of 64% (Denning, 2024; Guinea, 2014; McCarty et al., 2021). These infections range from mucosal candidiasis affecting the mouth and genitals to invasive candidemia that can spread to vital organs (Hoepelman & Dupont, 1996; Singh et al., 2014; Vazquez & Sobel, 2011).

Similarly, invasive aspergillosis caused by *Aspergillus* species is a leading cause of infection-related mortality in immunocompromised patients, particularly those with hematological malignancies, lung cancer, or chronic respiratory diseases (Caillot et al., 1997; Denning, 2024; Lortholary et al., 2000; Segal, 2009). *Cryptococcus neoformans* causes cryptococcosis, a life-threatening form of meningitis, especially in HIV/AIDS patients, with the highest burden in sub-Saharan Africa (Denning, 2024; Maziarz & Perfect, 2023). Other infections such as pneumocystis pneumonia and mucormycosis (*Rhizopus, Absidia, Mucor*) also significantly affect at-risk populations, especially in healthcare settings and regions with limited medical access (Denning, 2024; Gonzalez et al., 2002; Roden et al., 2005).

Endemic fungal infections highlight the intersection of geography, climate, and health equity. Coccidioidomycosis (Valley fever), caused by *Coccidioides immitis*, significantly affects populations in the southwestern United States, with migrant agricultural workers facing disproportionate risk due to occupational dust exposure (Kirkland & Fierer, 1996).

Fungal diseases such as blastomycosis, histoplasmosis, paracoccidioidomycosis, and penicilliosis demonstrate how environmental and socioeconomic factors shape the distribution and impact of endemic mycoses. For instance, blastomycosis, caused by *Blastomyces dermatitidis*, disproportionately affects Indigenous communities in the Great Lakes region and southeastern regions of the United States, where it can involve skin, bones, and the central nervous system (Klein et al., 1986). Similarly, histoplasmosis, caused by *Histoplasma capsulatum*, affects substantial populations globally, particularly in river valleys where bat and bird droppings create ideal growth conditions (Kurowski & Ostapchuk, 2002; Wheat et al., 1990). Climate change is increasingly altering these patterns, as flooding, extreme weather
events, and shifts in animal migration contribute to expanding exposure zones.

In Latin America, *Paracoccidioides brasiliensis* primarily affects rural agricultural workers, while *Penicillium marneffei* has emerged in Southeast Asia as a major opportunistic infection among HIV patients (Brummer et al., 1993; Sirisanthana & Supparatpinyo, 1998). Together, these examples highlight how fungal disease burden is often amplified by ecological disturbance, human vulnerability, and limited access to healthcare.

Fungal pathogens not only affect humans but also pose serious threats to animal health, often with serious consequences for wildlife and domesticated animals. Among fungal pathogens, *Batrachochytrium dendrobatidis* (Chytridiomycota), often shortened to “Bd,” has caused global declines in amphibian populations by triggering chytridiomycosis, a deadly skin disease that interferes with vital physiological functions (Kilpatrick et al., 2010; Sewell et al., 2021). Similarly, *Pseudogymnoascus destructans* (Ascomycota), the fungus responsible for white-nose syndrome, has devastated North American bat populations by interfering with hibernation and depleting fat reserves, ultimately leading to high mortality rates (Blehert et al., 2009; Hoyt et al., 2021). Fungal pathogens also present serious health risks to domesticated animals. Ringworm, previously mentioned in the context of human infections and caused by *Microsporum* and *Trichophyton* species, is a highly contagious skin disease that also commonly affects cats, dogs, and cattle and can easily spread between animals and humans (Seyedmousavi et al., 2015; Smith et al., 1969; Wright, 1989). Additionally, opportunistic fungi such as *Candida* and *Aspergillus* can lead to severe infections in livestock and poultry, especially in animals with weakened immune systems or those kept in poor living conditions (Osman et al., 2019; Sartori et al., 2014; Seyedmousavi et al., 2015).

## 4.2 | Fungal fury: Toxic Fungi

Many fungi produce a diverse array of secondary metabolites that act as chemical defences against mycophagy, particularly from fungivores including slugs, insects, and mammals. These compounds include phenolics, alkaloids, terpenoids, and polyketides, which deter, inhibit growth, or induce toxicity in organisms that ingest them. These metabolites are typically concentrated in various parts of the fungal organism, such as the fruiting bodies (mushrooms), mycelial networks, and spores, to mitigate consumption.

*Amanita muscaria*, recognized for its iconic bright red cap with white spots, has inspired various myths, legends, and artistic representations (Figure 4c). Its distinctive red-and-white appearance has influenced elements of popular culture, including *Super Mario* video games and Lewis Carroll's *Alice's Adventures in Wonderland*. This fungus belongs to the division Basidiomycota, order Agaricales, and the family Amanitaceae. Commonly referred to as fly agaric or fly amanita, it has earned these names because of its historical use in Europe to catch flies. *A. muscaria* is considered toxic because of its high content of muscarine, which can cause muscarine poisoning, tropane alkaloids, and isoxazole compounds, which affect the central nervous system

and can induce hallucinations (Rampolli et al., 2021). Its distinctive appearance makes accidental and severe intoxication extremely rare. In certain regions, it is consumed after undergoing specific preparation methods to remove the toxic components.

Unlike the fly agaric, the death cap (*Amanita phalloides* (Vetter 1998)) does not have a distinctive red and white appearance to warn potential consumers, and it comes perhaps as no surprise that this mushroom is responsible for the majority of fatal mushroom poisonings worldwide (Figure 4d). This is because of its widespread distribution and resemblance to certain edible mushrooms, which can lead to accidental consumption. Its toxicity is attributed to the presence of three mycotoxins: amatoxins, phallotoxins, and virotoxins. Among these, amatoxins, particularly α-amanitin, are the most lethal because they inhibit RNA polymerase II in the human liver, thereby preventing messenger RNA synthesis. Phallotoxins bind to F-actin filaments, preventing cell movement and division; however, little is known about the effects of virotoxins (Vetter, 1998).

## 5 | MIND-BENDING MUSHROOMS: FUNGI THAT TWIST OUR SENSES

“Magic mushrooms” are hallucinogenic mushrooms that have been used recreationally and in religious ceremonies for centuries and employed by shamans in healing rituals, divination practices, and other spiritual rites. Their psychoactive properties are due to the presence of psychotropic tryptamines, particularly psilocybin and psilocin, which are structurally similar to serotonin neurotransmitters. These compounds are present in *Psilocybe cubensis*, the most well-known and widely cultivated magic mushrooms because of their relatively large size, ease of cultivation, and potent hallucinogenic effects. Besides *Psilocybe*, there are many mushroom species, including the genera *Pluteus*, *Panaeolus*, *Inocybe*, *Gymnopilus*, *Galerina*, and *Pholiotina*, that contain high levels of psychotropic tryptamines (Gotvaldová et al., 2021). Recent genomic studies suggest that the genetic pathway for psilocybin biosynthesis has been horizontally transferred among different fungal lineages, contributing to the broad distribution of this trait. A leading ecological hypothesis suggests that psilocybin and related neuroactive compounds serve as a defensive or competitive strategy by altering the behavior of mycophagous animals and wood-eating invertebrate competitors (Reynolds et al., 2018).

Bioluminescent fungi are found mainly in temperate and tropical regions (Stevani et al., 2024). More than 125 bioluminescent fungi have been identified so far, which belong to the order Agaricales (Basidiomycota), with one potential ascomycete exception in the order Xylariales (Ascomycota) (Seas-Carvajal & Avalos, 2013). All documented bioluminescent *Agaricales* are mushroom-forming, white-spored agarics that fall into four fungal lineages: *Omphalotus*, *Armillaria*, and mycenoid fungi, with *Armillaria mellea* being the most widely distributed species. It has been proposed that bioluminescent basidiomycetes may attract invertebrates to aid in fungal spore dispersal or serve as a beneficial mechanism providing antioxidant protection (Desjardin et al., 2008, 2010).
Numerous fungi synthesize toxins that can harm animals, including mammals, and some even display remarkable biological activity, as exemplified by entomopathogenic species. Entomopathogenic fungi, particularly the so-called “zombie-ant” fungus *Ophiocordyceps unilateralis*, are widely recognized for their ability to control the behavior of insects, a phenomenon that has gained mainstream attention through media for being an inspiration for the 2013 video game *The Last of Us* and its subsequent television adaptation. Fungi from the genus *Ophiocordyceps* are parasites that infect insects, altering their behavior to ensure the optimal conditions for fungal growth and spore dispersal, often in sync with natural circadian rhythms, before ultimately leading to their death (Araújo & Hughes, 2019; Hughes et al., 2011). One particularly well-known species, *Ophiocordyceps sinensis*, also known as caterpillar fungus, holds significant industrial value, especially in traditional Chinese medicine, due to its immunomodulatory, anti-inflammatory, antioxidant, antitumoral, and cholesterol-lowering properties (Xu et al., 2016). Beyond the *Ophiocordyceps* genus, other fungi with similar host-manipulating abilities exist outside the order Hypocreales (Phylum: Ascomycota). Examples include species in the order Entomophthorales (Phylum: Zoopagomycota). The structural and life cycle differences between fungi in Ascomycota and Zoopagomycota likely influence the molecular mechanisms underlying their ability to control host behavior (Hywel-Jones, 2002). These particular entomopathogenic fungi are primarily found in tropical forest ecosystems, where they play a vital ecological role in regulating insect populations and contributing to environmental stability, despite their lethal effects on their hosts (de Bekker et al., 2021). These interactions are highly species-specific, and the possibility of a host switch to humans is considered extremely unlikely.

# 6 | ECOLOGICALLY IMPORTANT FUNGI

## 6.1 | Decomposers and nutrient recyclers

Fungi are fundamental drivers of ecosystem processes and act as decomposers, symbionts, and pathogens across a wide range of habitats. Their ecological roles are particularly notable in terrestrial ecosystems, where they influence nutrient availability, host population dynamics, and soil physicochemical composition (Bagchi et al., 2014; Clemmensen et al., 2013; Tedersoo et al., 2020). Fungi serve as primary agents of organic matter decomposition, especially recalcitrant plant polymers such as lignin and cellulose, facilitating the release and recycling of essential elements such as carbon, nitrogen, and phosphorus (Figure 5a).

A striking example of a fungal ecological impact even at the landscape level is *Armillaria ostoyae*, a species that forms vast subterranean mycelial networks (Figure 5b). One individual, located in Oregon, USA, spans approximately 9.6 km², making it the largest terrestrial organism known by area. A. ostoyae exhibits both saprotrophic and pathogenic behavior, contributing to wood decomposition and causing root rot in a range of tree species. This dual role exemplifies the

complex ecological strategies employed by fungi and their widespread effects on forest dynamics (Kim et al., 2023; Sipos et al., 2018).

Fungi play critical roles in aquatic ecosystems. Aquatic hyphomycetes and chytrids, for example, contribute to the fragmentation of organic particles and infection of phytoplankton, rendering energy and nutrients more accessible to higher trophic levels. These processes enhance trophic transfer efficiency and regulate food-web dynamics and biogeochemical cycling in freshwater and marine systems (Grossart et al., 2019).

Mycorrhizal fungi are among the most important symbiotic organisms in terrestrial ecosystems. They play a critical role in enhancing plant nutrient uptake in return for carbohydrates produced through plant photosynthesis. This mutualistic exchange not only improves plant fitness and productivity but also enhances the stability of plant communities under environmental stress (Read & Perez-Moreno, 2003; Smith & Read, 2010; Soudzilovskaia et al., 2020; Martin & van der Heijden, 2024). Over 90% of plant families form associations with mycorrhizal fungi, highlighting their ecological significance (Soudzilovskaia et al., 2020; Martin & van der Heijden, 2024). In addition to facilitating nutrient acquisition, these fungi help to protect plants against pathogens and environmental stressors. They exert profound effects on plant population and community dynamics, influencing processes such as seed dispersal, seedling establishment, soil niche differentiation, and both interspecific and intraspecific competition, thereby shaping patterns of plant diversity (Bahram & Netherway, 2022; Kadowaki et al., 2018; Kendrick, 2011; Smith & Read, 2010).

Mycorrhizal associations are commonly categorized into four principal types based on distinct anatomical and evolutionary characteristics (reviewed by Tedersoo et al., 2020). Arbuscular mycorrhiza (AM), formed by fungi in Glomeromycota and so-called fine root endophytes (Mucoromycota), are obligate mutualists and represent the most widespread form of mycorrhizal symbiosis. In AM, the fungi penetrate root cortical cells and form highly branched arbuscules surrounded by a plant-derived membrane, which serves as the site for nutrient exchange (Smith & Read, 2010; Willis et al., 2013). Ectomycorrhiza (EcM), involving more than 80 lineages of Basidiomycota (e.g., *Amanita* spp., *Russula* spp.), Ascomycota (e.g., *Tuber* spp., *Elaphomyces* spp.), and some Endogonomycetes, are obligate mutualists that are commonly associated with woody plant species (Figure 5c,d). In EcM, the fungi do not penetrate the plant cells but instead form a dense hyphal network between the root cortical cells, facilitating nutrient exchange. Ericoid mycorrhiza (ErM) is a symbiosis between various Ascomycota and some Basidiomycota species and plants in the Ericaceae family. ErM fungi proliferate extensively in the rhizosphere, enzymatically mining for nutrients, and form hyphal coils within the root epidermal cells, which are the primary interface for nutrient transfer (Read, 1996; Straker, 1996). Orchids form orchid mycorrhiza (OM), in association with fungi from the Ceratobasidiaceae, Serendipitaceae, and Tulasnellaceae families (all Basidiomycota), and occasionally with EcM fungi. OM fungi are typically free-living saprotrophs that facilitate seed germination and early growth in orchids; however, myco-heterotrophic achlorophyllous orchids often
<figure><img src="image_11.png" /><figcaption>**FIGURE 5** Ecologically important Fungi (purple): Representative phylogeny of selected genera from the phyla Ascomycota, Basidiomycota, Mucoromycota, and Glomeromycota, annotated with their respective taxonomic classifications. Photographed taxa mentioned in the "Ecologically Important Fungi" section are marked in purple. Source: He et al., 2022, 2024; Hyde, 2024.</figcaption></figure>

associate with mycorrhizal fungi and thereby parasitize plants via a commonly shared mycorrhizal mycelia. These associations are often called tripartite associations as they involve the orchid, the ectomycorrhizal fungus, and its photosynthetic host plant. Through this shared fungal network connecting multiple plants, the white orchids essentially take carbon from the photosynthetic trees, making them indirect parasites (Dearnaley, 2007; Rasmussen, 2002; Sathiyadash et al., 2012).

7 | THE FUNGAL PHARMACY

The discovery of penicillin has revolutionized modern medicine and laid the foundation for the development of antibiotics. It was discovered in 1929 when Alexander Fleming observed that mycelia growing from Penicillium spores, which had accidentally contaminated a bacterial culture plate, inhibited the growth of Staphylococcus bacteria,

revealing the antibiotic properties of penicillin (Fleming, 1929). To enhance production, more efficient Penicillium strains have been identified, including P. chrysogenum, which produced six times more penicillin than Fleming's original strain, enabling large-scale industrial production by 1943 (Chain et al., 2005; Gaynes, 2017).

The mass availability of penicillin during World War II has saved millions of lives and transformed the treatment of bacterial infections. The large-scale production of penicillin not only revolutionized healthcare but also drove advancements in microbial strain enhancement and fermentation technologies, which became essential for developing subsequent antibiotic classes (Kardos & Demain, 2011).

By 1950, extensive screening initiatives led by pharmaceutical companies had revealed a wide variety of microbial products with significant pharmaceutical potential. Since then, thousands of compounds that exhibit antibacterial, antifungal, antiparasitic, antiviral, and anticancer properties have been discovered. Additionally, numerous other bioactive molecules with cytotoxic, mutagenic, carcinogenic,
teratogenic, immunosuppressive, enzyme-inhibitory, allelopathic, and other biological activities have been identified in several fungal species beyond Penicillium, demonstrating the immense pharmaceutical value of fungal metabolites (Niego et al., 2023; Keller et al., 2005). Key examples include cephalosporin antibiotics, lovastatin (cholesterol-lowering drug), cyclosporin immunosuppressants, and other essential drugs such as griseofulvin, mycophenolic acid, mizoribine, and mevastatin (Devi et al., 2020; Gaynes, 2017; Keller et al., 2005; Niego et al., 2023; Pitt et al., 2014).

Drug discovery has predominantly focused on a few Ascomycota classes, particularly Saccharomycetes, Eurotiomycetes (Penicillium, Aspergillus), and Sordariomycetes (Acremonium spp., Claviceps spp., Melanocarpus spp., Ophiocordyceps spp., Tolypocladium spp.) (Niego et al., 2023). However, recent studies suggest that many fungi, including nonmodel and Basidiomycota species, may be even richer sources of biosynthetic genes responsible for producing natural products with potential medicinal value (Pasinato & Singh, 2025). Psilocybin (previously discussed in Section 4) has received renewed attention for its therapeutic potential in treating mental health disorders (Johnson & Griffiths, 2017; Lowe et al., 2021; Passie et al., 2002). Similarly, recent genomic analyses have revealed that Lecanoromycetes, an Ascomycota class composed almost entirely of lichenized fungi, have emerged as one of the richest fungal classes in terms of secondary metabolite gene content (Singh et al., 2025). A key feature that enhances the biotechnological promise of fungi, including the nonmodal taxa, lies in their osmotrophic mode of nutrition, which not only facilitates the uptake of dissolved nutrients but also renders them highly effective at secreting the specialized metabolites they synthesize. This inherent capacity for metabolite exudation makes fungi particularly attractive platforms for industrial-scale natural product production. These findings indicate that fungi traditionally known only for their ecological roles or environmental presence may soon become prominent players in industrial and pharmaceutical industry.

8 | CONCLUSIONS

The presentation of fungal taxonomy along with recognizable, day-to-day examples creates an approachable gateway to study fungi. This review aims to serve as an educational and introductory systematic resource that enhances comprehension of fungal diversity and classification. By contextualizing taxonomic groups using familiar species, this study encourages a more cohesive perspective on fungal evolution and underscores its relevance across various biological fields. We illustrated how fungi from diverse taxonomic groups are deeply intertwined with our daily lives, via medicines or impacting human health and plant health through both disease and root-associated symbiosis. We also highlight the ways in which fungi have become integral to human life, particularly through their role in medicine and food. Ultimately, we advocate that fostering fungal awareness and education is essential to understanding their relevance in our everyday lives

and to building a more harmonious future alongside these remarkable organisms.

AUTHOR CONTRIBUTIONS

Anna Vaiana and Garima Singh planned and structured the review. Anna Vaiana and Garima Singh wrote the first manuscript draft. Anna Vaiana, Rasmus Kjøller, Francis Martin, Toby Spribille, and Garima Singh contributed sections to the manuscript.

ACKNOWLEDGMENTS

G.S. was supported by the Italian Ministry of University and Research (project funded by the European Union - Next-Generation EU: 'PNRR Missione 4 Componente 2, "Dalla ricerca all'impresa", Investimento 1.4, Progetto CN00000033'). Open access publishing facilitated by Universita degli Studi di Padova, as part of the Wiley - CRUI-CARE agreement.

CONFLICT OF INTEREST STATEMENT

The authors declare no conflicts of interest.

DATA AVAILABILITY STATEMENT

Data sharing is not applicable to this article as no datasets were generated or analyzed during the current study.

ORCID

Anna Vaiana ID https://orcid.org/0009-0003-0289-4722

Francis Martin ID https://orcid.org/0000-0002-4737-3715

Rasmus Kjøller ID https://orcid.org/0000-0002-2027-4119

Toby Spribille ID https://orcid.org/0000-0002-9855-4591

Garima Singh ID https://orcid.org/0000-0003-3753-5431

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**How to cite this article:** Vaiana, A., Martin, F., Kjøller, R., Spribille, T., & Singh, G. (2026). Demystifying fungal systematics: A gateway to fungal literacy and societal/ecological relevance through familiar species. *Plants, People, Planet*, **8**(2), 499-515. https://doi.org/10.1002/ppp3.70119