Seven of the nine new species belong to the genus Mycena, a sprawling group of roughly 1,312 described species characterized by delicate, small fruiting bodies with thin convex caps bearing sulcate margins, non-deliquescent gills, and hollow stems. The newly described Mycena species are M. albomarginata, M. oblongispora, M. lavendula, M. longnanensis, M. minispora, M. pakistanica, and M. tephroleuca. The remaining two novelties belong to the allied genera Atheniella and Phloeomana, both of which were carved out of Mycena in recent years as DNA-based phylogenetics exposed the genus as highly polyphyletic. All nine species were described on the basis of detailed morphological investigation combined with molecular analyses of the nuclear internal transcribed spacer (ITS) region and the translation elongation factor 1-alpha (tef1-α) gene.

The technical backbone of the study is a combined ITS plus tef1-α dataset comprising 152 taxa, including 140 Mycena species, eight Atheniella species, and two Phloeomana species. The aligned matrix contained 1,474 characters, of which 902 were parsimony-informative. Maximum likelihood, maximum parsimony, and Bayesian inference analyses, run through the CIPRES Science Gateway using RAxML-HPC BlackBox 8.2.6, PAUP on XSEDE, and MrBayes on XSEDE 3.2.6 respectively, produced largely congruent topologies. The best-fit substitution model for the Bayesian analysis was GTR + I + G, and the resulting phylogeny showed that each of the ten newly described species, including one additional novelty registered during the study, forms a distinct, well-supported lineage clearly separated from known species in Atheniella, Mycena, and Phloeomana. Xeromphalina campanella served as the outgroup for rooting the tree.

Within Mycena, the new species cluster into three sections, each defined by a constellation of macroscopic and microscopic traits. Five species—M. lavendula, M. longnanensis, M. minispora, M. pakistanica, and M. tephroleuca—belong to Mycena sect. Calodontes, a group characterized by pinkish, reddish, purplish to brownish, mostly hygrophanous caps, interveined gills, smooth cheilocystidia and pleurocystidia when present, and mostly amyloid spores. One species, M. albomarginata, belongs to Mycena sect. Mycena, distinguished by relatively large and robust fruiting bodies with a typically grayish to brownish, lubricous or viscid cap surface and a rigid, often rooting stipe. The final species, M. oblongispora, falls within Mycena sect. Fragilipedes, the largest section, marked by grayish-brown to dark brown caps, smooth cystidia, a pileipellis typically covered with excrescences, and amyloid basidiospores.

The morphological descriptions are meticulous. Mycena lavendula, collected in the Taohe National Nature Reserve, is immediately recognizable by its lilac cap and reddish-purple stipe that becomes lilac toward the base, with longitudinally striate surfaces and a pruinose apex. Its basidiospores average 6.68 by 2.56 micrometers, subcylindrical to ellipsoid with an oil drop, and its cheilocystidia are polymorphic—clavate, fusiform, pyriform—some with rounded apices. Mycena longnanensis, from Longnan City, presents an umbonate, radially fibrillose, lilac to dark purple pileus with deeply decurrent lilac lamellae and distinctive spheropedunculate cheilocystidia averaging 39 by 14.21 micrometers with a slightly elongated tail and rounded apex. Mycena minispora, found in Yeliguan National Forest Park, is the largest of the new Chinese species at 3.5 to 3.7 centimeters across, with a glutinous, fibrillose golden-brown pileus, milky-yellow lamellae, and remarkably small amyloid basidiospores averaging 6.04 by 3.13 micrometers.

Mycena pakistanica, the sole novelty from Pakistan, collected in Abbottabad district, is arguably the most striking. Its lavender pileus bears grape-wine fibrils, and its stipe stretches up to an extraordinary 12 centimeters—exceptional for a bonnet mushroom. The species produces small lacrymoid basidiospores averaging 5.9 by 2.6 micrometers, spathulate pleurocystidia up to 74 micrometers, and fusiform to sublageniform cheilocystidia with both ends sharp, reaching up to 91 micrometers. The presence of a pinkish to purplish pileus, large basidiomata, and smooth hymenial cystidia confirms its placement in sect. Calodontes. Mycena tephroleuca, from Diebu County, offers a contrast: small chalky-white to grayish-white fruiting bodies with adnate lamellae bearing a decurrent tooth, a white stipe with a brown tinge at the base, and clavate to narrowly fusiform or utriform cheilocystidia with cylindrical outgrowths.

The two allied novelties illuminate the broader evolutionary picture. Atheniella parvispora, from Lianhuashan National Forest Park, is characterized by a papillate, umbilicate, orangish-yellow pileus with off-white margins, small lacrymoid basidiospores averaging 6.71 by 3.70 micrometers, and abundant fusiform cheilocystidia. Atheniella was elevated from Mycena sect. Adonideae in 2012 and is differentiated from Mycena by brightly colored caps—yellow, orange, pink, or red—and tissues that are unreactive in Melzer’s reagent. Phloeomana qilianensis, from the Qilian Mountains National Nature Reserve, produces small clustered basidiomata up to 1 centimeter across with an off-white to pale yellow pileus, fimbriate lamellae, a furfuraceous stipe, and inamyloid basidiospores averaging 7.39 by 3.99 micrometers. Phloeomana was formerly placed in the family Porotheleaceae but has recently been reassigned to the family Cyphellaceae, a small monophyletic family comprising a heterogeneous assemblage of lignicolous fungi with agaricoid, cyphelloid, clavarioid, pleurotoid, corticioid, or stereoid fruiting bodies.

Both families belong to the suborder Marasmiineae, circumscribed on a phylogenomic basis by Dentinger and colleagues in 2016, corresponding to the Marasmioid clade sensu Matheny and Binder. The reclassification of these mycenoid lineages reflects a broader revolution in fungal systematics. Recent phylogenetic studies have placed the so-called hydropoid fungi across several independent evolutionary lineages within the families Porotheleaceae and Cyphellaceae. In recent years, new genera such as Atheniella, Phloeomana, Leucoinocybe, Pruinomycena, and Chrysomycena have been proposed to accommodate species formerly squeezed into Mycena, yet several unresolved clades persist and the phylogenetic relationships within the genus remain unclear. The authors emphasize that the phylogenetic framework of Mycena sensu lato should be continuously refined, particularly through the inclusion of an increasing number of type materials and sequences from a broader geographic range.

The ecological significance of these fungi is considerable. Mycena species play important roles in ecosystems as decomposers, promoting nutrient cycling, forest metabolism, and natural renewal. Some form symbiotic associations with orchid seeds, enhancing germination through orchidoid mycorrhizae. Others are bioluminescent, and the genus represents a promising source of bioactive secondary metabolites. The study’s authors note that northwest China contains the largest tracts of virgin forest in the country, serving as essential refuges for a wide variety of unusual macrofungi. From Pakistan, only eight species of Mycena had previously been described based on morpho-anatomical characters, underscoring how little is known about the region’s fungal diversity.

According to the authors’ preliminary molecular and morphological studies, the species diversity of mycenoid taxa in Asia is substantially underestimated, and many additional new species remain to be described. At present, the ITS region remains the most effective DNA barcode for identifying mycenoid fungi, but additional specimens accompanied by multigene data—SSU, LSU, rpb1, and rpb2—are needed to further investigate species diversity and phylogenetic relationships within Mycena sensu lato. With further fieldwork and the availability of more collections, additional new taxa will undoubtedly be revealed, and the bonnets of Asia’s forests may yet yield many more surprises.

Subject of Research: Mycology

Article Title: Unveiling the diversity of the families Cyphellaceae and Mycenaceae from Southeast Asia

Article References: Fan, L. F., Naseer, A., Khalid, A., Khalid, A. N., Deng, Y., Wang, C. Y., Wang, Y., & Ma, S. X. (2026). Unveiling the diversity of the families Cyphellaceae and Mycenaceae from Southeast Asia. IMA Fungus, 17, Article e182174. https://doi.org/10.3897/imafungus.17.182174

Image Credits: AI Generated

DOI: 10.3897/imafungus.17.182174

Keywords: mycology, fungi, taxonomy, phylogenetics, Mycena, biodiversity, Southeast Asia, new species, Unveiling, diversity, families, Cyphellaceae