**4. Studies about bryophilic mushrooms**

Bryophilic mushrooms have been known for a long time, their habitats are well known (swamps, moss-covered tree trunks and mounds). In the 1980's the works about bryophilous fungi around the world include optical microscopy observations, in both Ascomycota and Basidiomycota. In 1981, the fungal species *Lyophyllum palustre* (Peck) Singer was found in mosses and it was constantly associated with necrotic areas of the species, and relationships with bryophytes discussed. Apparently less aggressive species of associated mushrooms were obtained from pure cultures made of *Sphagnum capillaceurn* (Weiss) Schrank, isolating *Lyophyllum palustre* and *Galerina paludosa* (Fr.) Kühner [26]. Research was carried out in 1987 on bryophilic fungi found in samples from the main herbaria in central and northern Europe, identifying ascomycetes associated with the bryophyte *Polytrichum sexangulare*: *Bryochiton heliotropicus* Döbb., *Bryochiton perpusillus* Döbb., *Lizonia sexangularis* Döbb. & Poelt., *Protothelenella polytrichi* Döbb. & Mayrh., *Gloeopeziza interlamellaris* Döbb. and *Hymenoscyphus norvegularis* Döbb [27].

Bryophytes are involved in a variety of competitive, parasitic, symbiotic, mutualistic and not yet specified interactions with fungi [28]. Some of Bryophilous species have very specific substrates such as *Galerina paludosa* found only in *Sphagnum* swamps, *Cyphellostereum laeve* was found on polytrichoid mosses in coastal regions, in addition to *Rickenella fibula* (Bull.) Raithelh. and *Rickenella setipes* which descriptions reported that a mound formed by *Alnus glutinosa* roots, covered by bryophytes served as substrate for the development of the species [29, 30]. In the 1990s, the main genera of bryophytic fungi were well known: *Galerina*, *Omphalina*, *Rickenella*, *Hypholoma*, *Mycenella* and *Psilocybe*, whose collection area was more widespread, knowing that their nature is more delicate they must be collected with a considerable amount of their substrate of origin [30]. Some islands were targets of ecological studies, such as South Georgia (Southern South America) and Iceland where the bryophyte substrate used by some Agaricomycetes was reported. Collections focused further south of the island of South Georgia were found to have species of the genus *Galerina*, *Gerronema*, *Phaeogalera* and *Hypholoma* [31]. Experiments about growth symbiosis using *Laccaria* spp. on different substrates, including *Sphagnum*, showed that when *Sphagnum* was mixed with vermiculite it was beneficial for the development of the *Laccaria* species and its symbiotic effects [32]. Another research reports that *Sphagnum* and other bryophytes can increase the presence of macrofungi in the substrate [33]. Studies of capture and evasion of nitrogen in soils with suspension of mosses verified that rhizomes of bryophytes have covering links with hyphae of Basidiomycetes fungi, forming a sheath around the rhizoids. This is important since the soil alone cannot handle nitrogen excess, nor accumulating without moss [34].

Research related to Agaricomycetes fungi and mosses among the years 2000 to 2010 cover mainly themes involved in the taxonomy, phylogeny, distribution, diversity, and classification of these organisms. Based on morphological characteristics and phylogenetic analysis, it was proposed that the *Omphalina giovanellae* Bres. systematic position is better maintained in another genus, making the mushroom that grows between mosses and low grass belonging to the genus *Clitopilus* (Fr. ex Rabenh.) P. Kumm [35]. *Multiclavula ichthyiformis* Nelsen, Lücking, L. Umaña, Trest & Will-Wolf, was identified as a new basidiolichen from Costa Rica with terricolous habits that grows with bryophytes [36]. *Psathyrella laurentiana* A.H. Sm and *Omphalina philonotis* (Lasch) Quél. (currently classified as *Arrhenia philonotis* (Lasch) Redhead,

Lutzoni, Moncalvo & Vilgalys) had their occurrence linked to bryophytes and/or peatlands, suggesting a more specialized niche for some basidiomycetes [37]. The occurrence of decomposing species of the genus *Galerina* Earle and *Coprinus* Pers. in bryophyte swamps including *Polytrichum alpestre* Hoppe in South Georgia Island [38], previously reported [32] can suggest a niche specialization in some species of these genera. *Chromocyphella muscicola* (Fr.) Donk was reported for the first time in Turkey [39], a species that grows near or on mosses, or even parasitizes them while alive [40]. The occurrence in association with mosses is used as identifying character for *Maireina callostoma* (Pilát) W.B. Cooke when using an identification key of this genus [41], showing that associations between Bryophytes and Fungi is an important character also to the taxonomy of Fungi.

Phylogenetic analysis helped to understand the bryophilic habits and high concentration of basidiolichens in Hygrophoraceae (Basidiomycota, Agaricales), suggesting a predisposition of these fungi to change their mutualist nutrition associated with photobionts to saprotrophic [42]. The occurrence of *Marasmius epidryas* Kühner, currently classified as *Rhizomarasmius epidryas* (Kühner ex A. Ronikier) A. Ronikier & Ronikier, among mosses was recorded in cold areas of Canada, Denmark and Russia highlights the occurrence of this relationships in cold and harshest areas [43].

Among 2015–2022 most analyses focused on more complex analyses about bryophilic mushrooms. Association among *Sphagnum* mosses and 26 species of Agarics and Boleti (Agaricomycetes, Basidiomycota) were found in Ukraine, whereas *Galerina cerina*, *G. paludosa*, *G. sphagnicola*, *Hypholoma elongatum*, *H. udum,* and *Tephrocybe palustris* could be considered as closely associated by substrate links [44]. In 2018, it was demonstrated that many bryophilous Hymenochaetales have values of stable isotope indicating ectomycorrhizal habits or a biotrophic cluster indicative of parasitism or an endophytic lifestyle [18].

The mycorrhizal-like associations, diversity and distribution of fungal associations in bryophytes, as between liverworts of the Jungermaniidae and Marchantiidae with Basidiomycota clades were explored by bibliographic review [4]. Although several works have been published reporting the occurrence of Agaricomycetes fungi growing among mosses, the importance of more specialized studies on the subject is highlighted. Part of these works did not perform microscopic and phylogenetic analysis in order to prove the relationship between Fungi and Bryophytes, but only reported the growth of certain species among mosses. The use of microscopic and phylogenetic analysis will contribute not only to prove the association, but also to understand the phylogeny and evolution of these organisms.
