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Cuticle

A cuticle is a tough, flexible, non-mineralized outer covering secreted by the epidermis of various organisms, functioning primarily as a protective barrier against environmental stresses such as desiccation, pathogens, and mechanical damage.^[1] In biological contexts, cuticles are diverse in structure and composition, appearing in plants, fungi, and invertebrates, but they universally serve to regulate interactions between the organism and its surroundings.^[2] In plants, the cuticle forms an extracellular hydrophobic layer that coats the aerial epidermis of all land plants, consisting mainly of the polyester cutin embedded with waxes and phenolic compounds to minimize water loss through transpiration and provide defense against UV radiation and microbial invasion.^[3] This lipid-based barrier is essential for terrestrial adaptation, enabling plants to thrive in dry environments by significantly reducing water loss, while also influencing organ fusion during development and facilitating controlled gas exchange.^[4] Cuticle thickness and composition vary by organ and species, with thicker layers on leaves and fruits compared to stems, and it is synthesized by epidermal cells via the endoplasmic reticulum pathway involving fatty acid elongation and polymerization.^[5] In animals, particularly within the Ecdysozoa clade including arthropods and nematodes, the cuticle acts as an exoskeleton; in arthropods, it is composed of chitin microfibrils cross-linked with proteins, while in nematodes it is primarily collagen-based, offering structural support, preventing dehydration, and serving as an attachment site for muscles.^[6] This multilayered structure—typically including an outer epicuticle, exocuticle, and endocuticle—undergoes periodic molting (ecdysis) to allow growth, with calcification in some crustaceans enhancing rigidity for locomotion and protection.^[7] In nematodes like Caenorhabditis elegans, the cuticle additionally regulates osmotic balance and locomotion through its collagen-rich annuli and longitudinal ridges, making it critical for survival in diverse habitats.^[8] Fungal cuticles, composed of hydrophobins, chitin, and other polymers, similarly protect against environmental stresses and aid in spore dispersal.^[9]

Overview

Definition

A cuticle is a non-mineralized, often waxy or chitinous outer covering secreted by underlying cells, serving as a protective barrier against environmental stresses, desiccation, and pathogens in various organisms.^[4] This structure is characterized by its tough yet flexible nature, providing an interface between the organism and its surroundings while enabling essential exchanges like gas diffusion.^[10] In diverse taxa, including plants, animals, and fungi, cuticles have evolved convergently as analogous adaptations to terrestrial challenges, despite arising independently in unrelated lineages. The term "cuticle" derives from the Latin cuticula, a diminutive of cutis meaning "skin," evoking its role as a thin, skin-like layer.^[11] Its use in biological contexts emerged in the 17th century, with early descriptions in botany by Nehemiah Grew (1672) and Marcello Malpighi (1675), who referred to the external layer of plant organs.^[4] By the early 19th century, researchers like Adolphe Brongniart (1830) and John Stevens Henslow (1831) distinguished the cuticle as a distinct, homogeneous film separate from the epidermis, marking its recognition as a specialized structure in both botanical and entomological studies.^[10] In entomology, the term gained prominence for describing the arthropod integument's outer layer, with foundational work appearing in the 19th century alongside advances in microscopy. While functionally similar to other integumentary features, the cuticle must be distinguished from the epidermis, which comprises the underlying layer of living cells that secretes it, and from the exoskeleton, a broader term often encompassing hardened, sometimes mineralized structures in arthropods that include but extend beyond the non-mineralized cuticle.^[4] This non-cellular, acellular composition underscores the cuticle's role as a dead, protective overlay rather than a vital tissue.^[12] In fungi, the cuticle analogously refers to the outer gelatinous or protective layer on fruiting bodies, convergent in function but structurally distinct from plant or animal forms.

General Properties and Functions

Cuticles across diverse organisms exhibit a multilayered structure that typically includes an outer protective layer, often termed the epicuticle, and an inner region such as the procuticle in animals or a cutin-wax matrix in plants, with analogous gelatinous or hydrophobic outer layers in certain fungal fruiting bodies.^[6]^[3]^[13] This architecture provides flexibility, enabling accommodation of growth or movement, while conferring toughness to resist mechanical deformation and abrasion.^[6]^[3] Hydrophobicity is a universal trait, arising from lipid-rich compositions that minimize surface wettability and adhesion of water or particulates.^[6]^[3]^[13] These properties underpin key functions, including the prevention of uncontrolled water loss by forming a diffusion barrier that maintains internal hydration in terrestrial conditions.^[6]^[3]^[13] Cuticles also shield against ultraviolet radiation via scattering, reflection, or absorption by waxes and phenolics, reducing cellular damage from solar exposure.^[3] Mechanically, they offer structural support, acting as an exoskeleton in animals or reinforcing epidermal integrity in plants and fungi.^[6]^[3]^[13] In addition, cuticles contribute to structural coloration through thin-film interference in their layered nanostructures, generating iridescent hues observed in insect exoskeletons and certain plant surfaces without relying on pigments.^[14]^[15] The independent emergence of cuticles in animals, plants, and fungi exemplifies convergent evolution, driven by the shared selective pressures of terrestrial life, such as desiccation and abiotic stress, resulting in analogous hydrophobic barriers despite distinct biosynthetic pathways.^[16]^[17]

Animal Cuticles

In Humans

In humans, the nail cuticle, known as the eponychium, is a thin fold of skin located at the base of the fingernail or toenail, extending from the proximal nail fold to adhere to the dorsal surface of the nail plate. It is composed primarily of the stratum corneum, the dead outermost layer of the epidermis, forming a thickened, soft tissue barrier. This structure grows from the proximal nail bed and creates a tight seal between the epidermis and the nail plate.^[18]^[19]^[20] The primary function of the eponychium is to protect the underlying nail matrix from external irritants, trauma, and microbial invasion by maintaining an impermeable barrier that prevents pathogens from entering the sensitive area beneath the nail. This sealing action safeguards the germinal matrix, where nail growth originates, thereby supporting overall nail integrity and digit protection.^[18]^[21]^[22] The hair cuticle, or cuticula pili, forms the outermost layer of the hair shaft, consisting of a single layer of overlapping, scale-like keratinocytes that encircle the underlying cortex and medulla. These scales, arranged in an imbricated pattern resembling roof tiles, are highly keratinized, with keratin providing structural rigidity and resistance to environmental stress. This composition ensures the cuticle remains thin yet durable, typically comprising flattened epithelial cells cemented together.^[23]^[24]^[25] The hair cuticle primarily protects the inner hair layers from mechanical abrasion, chemical damage, and daily wear, preventing fraying of the cortex and the formation of split ends. Its smooth, overlapping scales also contribute to hair shine by facilitating even light reflection and reducing friction during movement. When intact, this layer enhances the hair's overall resilience and aesthetic appearance.^[23]^[26]^[27] Unlike the rigid exoskeletal cuticles found in invertebrates, human cuticles are soft, keratin-based structures serving protective and sealing roles without providing skeletal support. Clinically, disruption of the nail eponychium through trauma, biting, or cosmetic removal heightens the risk of paronychia, an inflammatory infection of the nail fold often caused by Staphylococcus aureus or other bacteria entering via the breached barrier. Acute paronychia manifests as localized redness, swelling, tenderness, and possible pus formation, while chronic forms involve persistent irritation leading to nail plate thickening and ridging.^[28]^[29]^[18] Aggressive manicure techniques, such as those involving complete cuticle excision with electric tools, can precipitate severe outcomes like onychomadesis (nail shedding) due to matrix inflammation and temporary growth arrest. Such procedures compromise the eponychium's protective function, increasing infection susceptibility and potential for permanent nail dystrophy, particularly in individuals with frequent exposure or immunosuppression. Proper nail care, including avoiding unnecessary cuticle trimming, is recommended to mitigate these risks.^[30]^[28]^[31]

In Invertebrates

In invertebrates, the cuticle serves as the primary exoskeleton, providing structural integrity and protection, particularly in phyla such as Arthropoda and Nematoda.^[32] This acellular layer is secreted by the underlying epidermis and is essential for locomotion, environmental interaction, and physiological regulation, differing markedly from the softer, non-molting cuticles in vertebrates.^[33] In arthropods, including insects and crustaceans, the cuticle is a composite of chitin and proteins, forming a chitin-protein matrix that imparts rigidity and flexibility.^[34] It consists of three main layers: the outermost epicuticle, a thin waxy barrier primarily composed of lipoproteins, fatty acids, and a wax monolayer that prevents water loss and blocks pathogens; the exocuticle, a hardened layer where proteins are cross-linked by quinones during sclerotization (tanning), creating durable sclerites; and the endocuticle, an inner flexible region of chitin microfibrils embedded in a protein matrix, arranged in lamellae for enhanced strength.^[33] This layered structure enables molting (ecdysis), where the old cuticle is shed and a new one secreted, allowing growth and metamorphosis.^[33] Additionally, nanoscale arrangements in the cuticle produce structural colors through light interference and diffraction, as seen in iridescent beetle elytra and butterfly wings, serving camouflage or signaling functions.^[35] In nematodes, the cuticle is collagen-based, reinforced by insoluble proteins called cuticlins that are cross-linked by dityrosine bonds for durability.^[36] It features distinct zones: the outer cortex, rich in cuticulin for surface resistance; a median zone with fluid-filled structures and minimal organization; and a basal zone with fibrous layers oriented at angles (approximately 75° and 135°) that support elasticity.^[36] Unlike the chitin-dominant arthropod cuticle, this collagenous structure lacks extensive sclerotization but maintains body shape through hydrostatic pressure.^[36] Across invertebrates, the cuticle provides mechanical support as a scaffold for muscle attachment and body rigidity, facilitating locomotion—such as the undulating waves in nematodes via their hydrostatic skeleton or the powered flight in insects through exoskeletal leverage.^[32]^[36] It integrates sensory functions through embedded sensilla, specialized cuticular structures like campaniform sensilla in insects that detect strain and mechanosensory hairs for tactile input, enabling environmental navigation.^[32] Adaptations for waterproofing, particularly in terrestrial arthropods, rely on epicuticular waxes that minimize desiccation, with insects losing up to 90% less water compared to unwaxed surfaces under dry conditions.^[33] In nematodes, the cuticle aids osmoregulation by modulating permeability to maintain ionic balance, crucial for survival in varying salinities or host tissues.^[37]

In Vertebrates

In vertebrates, cuticles manifest primarily as scales, which provide protection and serve adaptive functions distinct from the hair and nails seen in mammals. These structures arise through complex interactions between the epidermis and dermis, forming layered keratinous or mineralized coverings that enhance survival in diverse environments. Unlike the chitin-based exoskeletons of invertebrates, vertebrate scales are predominantly composed of keratins and often incorporate minerals, reflecting an evolutionary shift toward flexible, renewable integumentary protections.^[38] Reptile scales exemplify this keratin-dominated cuticle, featuring beta-keratin layers that create a robust, overlapping armor resistant to abrasion, desiccation, and predation. Beta-keratin, a hard corneous material unique to sauropsids, polymerizes into filaments that interlock to form the scale's outer surface, providing waterproofing essential for terrestrial life. These scales also facilitate thermoregulation by absorbing or reflecting solar radiation based on pigmentation and structure, while their coloration patterns enable camouflage against varied substrates. For instance, in snakes, periodic shedding renews the scale layer through a cyclical epidermal process, removing the outer beta-keratin stratum to maintain flexibility and prevent cracking.^[39]^[40]^[41]^[42] Fish scales, in contrast, often exhibit a mineralized composition suited to aquatic habitats, with cycloid scales featuring smooth, rounded edges and ctenoid scales displaying comb-like spines for enhanced grip or sensory function. Primarily built from hydroxyapatite crystals embedded in a collagen matrix, these scales form an enamel-like outer layer that offers puncture resistance and flexibility, allowing body contouring during movement. Hydrodynamically, the overlapping arrangement minimizes drag and turbulence, optimizing propulsion efficiency in water, while also contributing to ion regulation by serving as a calcium reservoir that buffers osmotic stress in fluctuating salinities.^[43]^[44]^[45] Evolutionarily, vertebrate scales originated from iterative epidermal-dermal signaling during embryogenesis, where dermal papillae induce epidermal thickening and keratinization, a process conserved across fish, reptiles, and birds but absent in chitin-synthesizing invertebrates. This dermal-epidermal interplay allows for scalable regeneration and patterning, contrasting sharply with the rigid, molted chitin cuticles of arthropods that require complete exoskeletal replacement. Beta-keratins in reptile scales share a distant homology with alpha-keratins in mammalian hair, underscoring a common integumentary heritage.^[38]^[46]

Plant Cuticles

Structure and Composition

The plant cuticle is an extracellular hydrophobic layer that covers the aerial epidermis of all primary land plants, serving as the interface between the plant and its environment. It consists of two main layers: the cuticle proper, which is the dominant structural component, and the underlying cuticular layer that merges with the cell wall. The primary polymer in the cuticle proper is cutin, a polyester composed mainly of inter-esterified C16 and C18 hydroxy and epoxy fatty acids, such as ω-hydroxyacids and mid-chain hydroxylated fatty acids, providing a flexible and insoluble matrix. Embedded within this cutin matrix are intracuticular waxes, which are complex mixtures of very-long-chain lipids including alkanes, primary alcohols, aldehydes, ketones, and secondary alcohols, contributing to hydrophobicity and mechanical properties. Epicuticular waxes form crystalline structures on the outer surface, often appearing as tubules, platelets, or rods that enhance water repellency. Additionally, the cuticle incorporates polysaccharides from the cell wall, such as pectins and hemicelluloses, and phenolic compounds like flavonoids and ferulic acid, which add cross-linking and UV-absorbing capabilities.^[3]^[10] Cuticle thickness varies from less than 0.1 µm to over 10 µm depending on the organ, species, and environmental conditions, with thicker cuticles on adaxial leaf surfaces and fruits. Biosynthesis occurs in epidermal cells, primarily through the endoplasmic reticulum where fatty acids are elongated to C16-C18, oxidized, and transported via ABC transporters to the extracellular space for polymerization by cutin synthase enzymes and self-assembly into the cuticle. Wax biosynthesis involves similar pathways but with decarboxylation and reduction steps, leading to deposition both inside and outside the cutin matrix. Unlike fungal structures, the plant cuticle is a true lipid-based barrier without chitin, relying on its polyester-wax composition for impermeability.^[3]^[47]

Functions and Adaptations

The plant cuticle serves as a primary diffusion barrier that minimizes uncontrolled water loss from aerial plant surfaces, thereby regulating transpiration and enabling survival in terrestrial environments. By forming a hydrophobic layer over the epidermis, it restricts non-stomatal water efflux, which can constitute a significant portion of total transpiration under closed-stomatal conditions; for instance, in species like maize, enhanced cuticular wax deposition under drought significantly reduces water loss compared to untreated controls.^[48] This function is complemented by the cuticle's role in pathogen resistance, where its lipophilic composition and thickness act as a physical and chemical shield against microbial invasion, limiting fungal and bacterial penetration in crops such as tomato and wheat.^[49] Additionally, the cuticle provides UV screening through phenolic compounds that absorb harmful ultraviolet radiation, dissipating most incident UV energy as heat via radiationless mechanisms, thus protecting underlying tissues from photodamage in various species.^[50] Adaptations of the plant cuticle enhance its protective roles in diverse ecological niches, often involving modifications to its nanostructure and composition. A prominent example is the lotus effect, observed in Nelumbo nucifera leaves, where hierarchical micro- and nanopapillae covered by epicuticular wax tubules create superhydrophobic surfaces with contact angles exceeding 150°, promoting self-cleaning by repelling water and contaminants to prevent pathogen adhesion and maintain photosynthetic efficiency.^[51] In floral structures, cuticular ridges and multilayers produce structural coloration through interference and diffraction, generating iridescent hues that attract pollinators; for example, in hibiscus petals, variations in cutin monomers drive the formation of diffraction gratings restricted to pigmented regions, enhancing visual signaling without pigment reliance.^[14] Under abiotic stresses like drought, plants dynamically increase cuticular wax biosynthesis and deposition—often by 2- to 3-fold in leaves of Arabidopsis thaliana and sorghum—thickening the barrier to further curb transpiration while maintaining gas exchange.^[52] The ecological significance of plant cuticles extends to their ancient origins and contemporary inspirations. Fossil evidence indicates that cuticles, characterized by preserved sporophyte tissues with stomatal complexes, first appeared in the Lower Devonian period (approximately 419-393 million years ago) in early vascular plants from regions like Yunnan, China, facilitating the colonization of land by mitigating desiccation.^[53] In modern contexts, these properties inspire biomimicry applications, such as developing water-repellent coatings and selective barriers modeled on cuticular waxes to improve crop resilience and create sustainable materials like polyester films mimicking cutin for moisture protection.^[54] The waxy components of the cuticle, primarily long-chain hydrocarbons, underpin these adaptations by modulating surface hydrophobicity and permeability.^[3]

Fungal Cuticles

Structure and Composition

The pileipellis serves as the outermost layer of the fruiting body in many fungi, particularly basidiomycetes, forming a protective skin composed of compacted hyphae arranged in a plectenchymatous tissue. This layer is typically gelatinous or filamentous, consisting of an outer hyphal covering that can range from a thin, parallel arrangement to a more intricate network with intercellular spaces less than 10 µm wide. Its primary structural elements derive from fungal hyphal walls, which are rich in chitin—a β-1,4-linked polymer of N-acetylglucosamine providing rigidity—and β-glucans, such as β-1,3- and β-1,6-linked polysaccharides that contribute to flexibility and matrix embedding.^[55]^[56]^[13] Pigments, often melanins, are incorporated into the hyphae, imparting coloration and potentially enhancing photoprotection, as observed in species like Stropharia rugosoannulata where pileipellis-derived melanins exhibit alkali-soluble properties characteristic of eumelanin.^[57] In basidiomycetes, pileipellis morphology shows significant variation, including dry types with a non-gelatinized, appressed hyphal cutis and viscid types embedded in a gelatinous matrix that appears glutinous when moist due to mucilaginous polysaccharides.^[58] These differences influence surface texture and water retention, with viscid forms often featuring interwoven hyphae 2–7 µm wide in a gelatinous epicutis up to 250 µm thick. Amyloid reactions, detectable via staining with Melzer's reagent, occur in certain hyphal elements or associated structures, turning bluish-black and aiding taxonomic identification, particularly in genera like Russula where spore ornamentation and pileipellis components react positively. Thickness can vary from 10–400 µm, with ornamentation such as scales or fibrils arising from erect hyphal tufts in trichodermial arrangements, as seen in species with heterogeneous, non-gelatinized layers.^[59]^[60] Biosynthesis of the pileipellis occurs through extension of the underlying hyphal network during fruiting body development, incorporating cell wall polymers synthesized via enzymes like chitin synthases and glucan synthases, resulting in a transient structure integrated with the trama. Unlike plant cuticles, it lacks a true waxy, cutin-based barrier, relying instead on hyphal density and minor lipid components for hydrophobicity.^[56]^[13]

Functions and Role in Fungi

The fungal cuticle, often referred to as the pileipellis in the context of basidiomycete fruiting bodies, serves primarily as a protective barrier that minimizes water loss through its low permeability, ranging from 2.8 to 9.8 × 10^{-4} m s^{-1}, thereby reducing transpiration by factors of 10 to 30 compared to an uncovered water surface.^[13] This function is crucial for maintaining the structural integrity of ephemeral mushroom fruiting bodies in variable terrestrial environments, preventing desiccation that could impair spore production. Additionally, pigmentation within the pileipellis, such as melanin or other phenolic compounds incorporated into the hyphal walls, absorbs ultraviolet (UV) radiation, converting it to heat and shielding underlying tissues from photodegradation and DNA damage.^[61]^[62] The pileipellis also contributes to defense against microbial pathogens and herbivorous invertebrates by forming a physical and chemical barrier; its interwoven hyphal structure impedes penetration by antagonistic fungi and bacteria, while surface waxes and secondary metabolites deter grazing by slugs and insects.^[13] In species like Lentinula edodes, pigments in the outer layer further enhance resistance to microbial colonization by absorbing harmful wavelengths that could otherwise promote opportunistic infections.^[63] In reproductive biology, the fungal cuticle facilitates spore dispersal by sustaining moisture levels in the fruiting body, which prolongs viability and supports convective airflows beneath the cap driven by temperature gradients, thereby aiding the release and transport of basidiospores.^[13] In hygrophanous fungi, such as species in the genus Hygrophorus, the gelatinous nature of the pileipellis—composed of gelatinized hyphae—enhances moisture retention, allowing the cap to absorb and hold water, which stabilizes color changes and maintains humidity around the hymenium for optimal spore maturation and discharge.^[64] The structural variations in the pileipellis hold significant taxonomic value in mycology, particularly for identifying basidiomycete genera; for instance, a trichodermium—characterized by erect, perpendicular hyphae—distinguishes boletes in genera like Suillus and Rubinoboletus, while ixotrichodermia with gelatinous elements aid in classifying hygrophanous agarics.^[65]^[66] Evolutionarily, the pileipellis represents an elaboration of ancestral fungal cell wall layers, such as the chitin-glucan matrix, which provided early protection against desiccation in terrestrial-colonizing fungi, adapting over time to specialized roles in fruiting body defense and reproduction.^[67]

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Jul 21, 2025 · This observation indicates that the pigments derived from the pileipellis of S. rugosoannulata belong to the melanin family, while exhibiting ...
[52]
[PDF] Glossary - New York Botanical Garden
pileus surface embedded in a gelatinous matrix; surfaces appear viscid or glutinous to the naked eye. Ixotrichodermium - long and usually multi-celled ...
[53]
Testing spore amyloidity in Agaricales under light microscope
Nov 11, 2020 · It has been known for about 150 years that some ascomycete and basidiomycete sporomata may contain elements which stain grey to blue-black with ...
[54]
New Data on Boletaceae (Agaricomycetes, Basidiomycota) from ...
Mar 19, 2024 · The heterogeneous structure of the pileipellis, which is not a true ... Do H.T., Nguyen P.D.N. Composition of edible macrofungi species ...
[55]
Leaf cuticle and water loss in maize lines differing in dehydration ...
Aug 10, 2025 · The plant cuticle plays an important role in protecting plants against water loss. The objectives of this study were to investigate the ...<|separator|>
[56]
Multifunctional Roles of Plant Cuticle During Plant-Pathogen ...
This review describes multiple roles of plant cuticle during plant-pathogen interactions and its responses to both leaf and fruit pathogens.
[57]
Radiationless mechanism of UV deactivation by cuticle phenolics in ...
Apr 4, 2022 · As such, the cuticle can be regarded as the first and foremost protective barrier against UV radiation. This photostable and photodynamic ...
[58]
Superhydrophobicity in perfection: the outstanding properties ... - NIH
Mar 10, 2011 · The lotus plant has successfully developed an excellent protection for this delicate epistomatic surface of its leaves. Keywords: epicuticular ...
[59]
Wax biosynthesis in response to danger: its regulation upon abiotic ...
Apr 3, 2020 · Drought treatments significantly increased cuticular wax load in A. thaliana (Kosma et al., 2009; Seo et al., 2011), as well as in various ...
[60]
Studies on plant cuticles from the Lower–Middle Devonian of China
Numerous plant cuticles were collected from five sections in the Lower–Middle Devonian of Yunnan and Sichuan provinces, China. Based on these cuticles and ...
[61]
Plant Surfaces: Structures and Functions for Biomimetic Innovations
The cuticle is basically a biopolymer made of polyester called cutin, impregnated with integrated (intracuticular) waxes. Additionally, waxes on the cuticle ...
[62]
2.3.2: Characteristics of Fungi - Biology LibreTexts
Jul 28, 2025 · Pigments in fungi are associated with the cell wall and some play a protective role against ultraviolet radiation. Some fungal pigments are ...
[63]
Synthesis and structural characteristics analysis of melanin ... - NIH
Sep 29, 2023 · In this study, elemental composition analysis of melanin pigments revealed the presence of elements C, N, H, O, and S in both samples (Table 1).
[64]
Characterization of brown film formed by Lentinula edodes
Eumelanins are dark brown to black pigments containing 6–9 % nitrogen and 0–1 % sulfur. Pheomelanins are red–brown pigments containing 8–11 % nitrogen and 9–12 ...
[65]
Hygrophorus chrysodon: A Mushroom Between Myth and Reality
Sep 17, 2025 · The cap cuticle is viscous in consistency under humid conditions, especially in young specimens, while it becomes drier in dry weather. This ...
[66]
[PDF] suillus - USDA Forest Service
Cuticle of pileus a deep trichodermium of tangled (loosely) hyphae with short more or less differentiated cells near apex 9-15 J.l in diam., and with orange ...
[67]
About Boletes - MykoWeb
However, there is considerable variation in the trichodermial cuticle, and noticeable changes may occur as the basidiocarp matures. If at all possible, young ...
[68]
Fungal evolution: major ecological adaptations and evolutionary ...
Herein we survey the main evolutionary and ecological processes that have guided fungal diversity.Missing: convergence | Show results with:convergence