Eoscyphella luciurceolata gen. and sp. nov. (Agaricomycetes) Shed Light on Cyphellopsidaceae with a New Lineage of Bioluminescent Fungi

During nocturnal field expeditions in the Brazilian Atlantic Rainforest, an unexpected bioluminescent fungus with reduced form was found. Based on morphological data, the taxon was first identified as belonging to the cyphelloid genus Maireina, but in our phylogenetic analyses, Maireina was recovered and confirmed as a paraphyletic group related to genera Merismodes and Cyphellopsis. Maireina filipendula, Ma. monacha, and Ma. subsphaerospora are herein transferred to Merismodes. Based upon morphological and molecular characters, the bioluminescent cyphelloid taxon is described as the new genus Eoscyphella, characterized by a vasiform to urceolate basidiomata, subglobose to broadly ellipsoid basidiospores, being pigmented, weakly to densely encrusted external hyphae, regularly bi-spored basidia, unclamped hyphae, and an absence of both conspicuous long external hairs and hymenial cystidia. Phylogenetic analyses based on ITS rDNA and LSU rDNA support the proposal of the new genus and confirm its position in Cyphellopsidaceae. Eoscyphella luciurceolata represents a new lineage of bioluminescent basidiomycetes with reduced forms.


Introduction
Agaricomycetes forms a large and diverse group that includes the mushroom-forming fungi and produces the most complex basidiomata forms, such as gilled mushrooms, boletes, polypores, and puffballs [1].Some species of gilled mushroom are well known and stand out for their natural light emission with a luciferin/luciferase chemical reaction [2,3].The bioluminescent fungi are morphologically well characterized and typically known for their gilled or poroid basidiomata within the order Agaricales [4].The known bioluminescent mushrooms are distributed in tropical and temperate regions, where they grow on moist decaying wood or leaves [4].
During one of many nocturnal expeditions into the Atlantic Rainforest in the state of São Paulo (Brazil), in the same area where 12 bioluminescent species have already been described or recorded [22][23][24][25], an unusual bioluminescent fungus with cyphelloid form was discovered by co-authors of this work.The aims of this study are as follows: (i) confirm the phylogenetic position and classification of all known bioluminescent fungi based on molecular data; (ii) identify, based on morphology and molecular data, the new bioluminescent fungi with reduced form; and (iii) provide the phylogenetic placement of Maireina monacha to better understand its relationship with related genera.Based on molecular analyses, Maireina is considered a synonym of Merismodes and is herein amended.Maireina filipendula Laessøe, Ma. monacha (Speg.)W.B. Cooke, and Ma.subsphaerospora Mombert are transferred to Merismodes, and the new bioluminescent cyphelloid taxon from Brazil is described in the new genus Eoscyphella gen.nov., within Cyphellopsidaceae.Eoscyphella luciurceolata represents a new lineage of bioluminescent basidiomycetes with cyphelloid form. .The forest type is Dense Ombrophilous Forest, which is mainly composed of the Angiosperm families Annonaceae Juss., Euphorbiaceae Juss., Lauraceae Juss., Melastomataceae Juss., Moraceae Gaudich., Myrtaceae Juss., Rubiaceae Juss., and Sapotaceae Juss.[64,65].

French Site of Maireina monacha
The Butte de la Garenne is located in the Cantal department in Southern-Central France.The site is covered with a calcareous beech forest of approximately one hectare, and a pubescent oak forest in the remaining area [66].

Morphological Analyses
Macroscopic features were recorded from fresh material.Color names and codes follow Kornerup and Wanscher [67].Micromorphological analyses were performed using the methodology of Bodensteiner [53].Basidiospores were measured in lateral view using 5% KOH.Basidiospore statistics include the following: xm = arithmetic mean of basidiospore length × basidiospore width (±standard deviation) for n basidiospores measured in a single specimen; xr = range of basidiospore means; Q = quotient of basidiospore length by basidiospore width in any one basidiospore, indicated as a range of variation in n basidiospores measured; Qm = mean of Q-values in a single specimen; n = number of basidiospores measured per specimen; and s = number of specimens studied.Distilled water was used in order to visualize crystals in skeletal hyphae, whilst Melzer's reagent was used to test amyloid/dextrinoid reactions.The Brazilian specimens were deposited at the Fungarium IFungi (FIFUNGI) from the IFungiLab at the "Instituto Federal de Educação, Ciência e Tecnologia de São Paulo (IFSP)", Brazil, and the European specimens are housed at the "Muséum National d'Histoire Naturelle" (P), France ( [68], 2023, continuously updated).

Molecular Methods
Entire basidiomata were homogenized in lysis tubes with magnetic beads for three cycles of 2 min in SpeedMill Plus (Analytik, Jena, Germany) in an AP1 buffer, and the genomic DNA was extracted using the Qiagen Dneasy ® Plant Mini Kit (Germantown, MD, USA) according to the manufacturer's instructions.Primer pairs ITS1-F/ITS4 and LR0R/LR5 were used to amplify and sequence the ITS rDNA region and the LSU rDNA gene, respectively [44,69].Sequencing reactions were conducted at Macrogen (Seoul, Republic of Korea).

Phylogenetic Analyses
The newly generated sequences were assembled and edited in Sequencher TM v5.0 software (Gene Codes Corporation, Ann Arbor, MI, USA) and were deposited in GenBank (codes in the tree and in Supplementary Table S1).Three new ITS rDNA and two novel LSU r DNA sequences were generated in this study.Three distinct datasets were constructed: one composed only of the LSU rDNA sequences, one only with ITS rDNA sequences, and a third including the ITS rDNA + LSU rDNA sequences.To assemble the LSU rDNA dataset, our generated sequences were submitted to the BLASTn algorithm at NCBI (GenBank, https://blast.ncbi.nlm.nih.gov/Blast.cgi,accessed on 1 May 2023) to retrieve similar sequences.Other sequences of cyphelloid basidiomycetes, including those generated by Bodensteiner et al. [45], Laessøe et al. [54], Baltazar et al. [48], Karasi ński et al. [56], and Vizzini et al. [70], were downloaded and included in the dataset.Existing sequences of known bioluminescent fungi were also downloaded from GenBank to compose a final dataset that includes all known bioluminescent and cyphelloid lineages.The LSU rDNA sequence most likely misnamed G. viridilucens (EF514207), which is available on GenBank and was used in the phylogenetic analyses by Vizzini et al. [70] and Na et al. [71], is 99.7% identical to sequence M. lucentipes (DED7828) [72].For this reason, we excluded the sequence EF514207 from our phylogenetic analyses and included a new one of G. viridilucens (DED7822), originally from type locality and morphologically described and confirmed [22].For the ITS rDNA dataset, sequences of species belonging to Cyphellopsidaceae were retrieved from GenBank and then used to recover similar sequences using the BLASTn algorithm.The combined ITS+LSU rDNA dataset was constructed to focus primarily on the Merismodes clade.Boletus griseiceps B. Feng, Y.Y.Cui, J.P. Xu & Zhu L. Yang; Boletus subviolaceofuscus B. Feng, Y.Y.Cui, J.P. Xu & Zhu L. Yang; and Fistulinella ruschii Magnago were used as an outgroup in the LSU rDNA dataset.Two sequences of Cunninghammyces Stalpers were used as an outgroup in the ITS rDNA dataset, and sequences of Acanthocorticium brueggemannii Baltazar, Gorjón & Rajchenb were used for the combined analyses.

LSU rDNA Dataset
The final LSU rDNA dataset contains 206 sequences (including 2 that are newly generated), consisting of 1051 nucleotide sites, including gaps.The most appropriate evolutionary model estimated was TrN+I+G.The bootstrapping criteria from the ML analyses stopped after 350 replicates.Both the RAxML analysis and Bayesian inference yielded similar tree topologies.The LSU rDNA tree generated from the ML analysis, including bootstrap values and posterior probabilities, is shown in four parts (Figure 1a-d).
The new proposed genus Eoscyphella formed a well-supported clade (100% BS, 1.0 BPP) sister to the cyphelloid genus Woldmaria.The LSU rDNA sequences of Eoscyphella and Woldmaria are from 92.6% to 92.7% similar.Sequences of taxa of the genus Maireina were represented in our analyses by Ma. filipendula, Ma. subsphaerosphora, and Ma.monacha (type species of Maireina), with the latter sampled, sequenced, and identified by Mombert [55] and fully described and epityfied in this study.Maireina formed a paraphyletic group, but represents a monophyletic clade when including sequences of Merismodes anomala (Pers.)Singer (as = Cyphellopsis anomala) and Me.fasciculata (Schwein.)Earle, with the latter being the type species of the genus.The clade formed with Maireina and Merismodes is well supported (75% BS, 0.99 BPP).

ITS rDNA Dataset
The final ITS rDNA dataset has 44 sequences (including 3 that are newly generated), consisting of 1051 nucleotide sites, including gaps.The best evolutionary models estimated for each part of the alignments were ITS1: TPM2uf+G, 5.8S: TPM2+G, and ITS2: HKY+G.The bootstrapping criteria from the ML analysis stopped after 300 replicates.Both the RAxML analysis and Bayesian inference yielded similar tree topologies.The ITS rDNA tree generated from the ML analysis, including bootstrap and posterior probabilities, is shown in Figure 2.  The family Cyphellopsidaceae (100% BS, 1.0 BPP) is represented by 42 sequences, with no representatives of Woldmaria nor Peyronelina due to lack of available sequences.Eoscyphella luciurceolata sp.nov.and the non-bioluminescent Eoscyphella sp.formed a well-supported clade (84% BS, 1.0 BPP), sister to (but not supported) a clade that contains sequences of Dendrothele microspora (H.S. Jacks.& P.A. Lemke) P.A. Lemke, Dendrothele incrustans (P.A. Lemke) P.A. Lemke, and Dendrothele griseocana (Bres.)Bourdot & Galzin (Figure 2).The genus Maireina, represented by the same species as in the LSU rDNA analyses, is again confirmed as paraphyletic with the ITS rDNA data.However, as in the nLSU analyses, the included Maireina sequences form a monophyletic and well-supported clade (91% BS, 1.9 BPP) when including sequences of Merismodes anomala, Me. fasciculata, and Merismodes sp.(MZ919217).

Combined LSU rDNA + ITS rDNA Dataset
The final combined LSU rDNA plus ITS rDNA dataset contains 20 ITS rDNA and 16 LSU rDNA sequences (including 5 generated as part of this study) for 21 terminals, and consists of 1806 nucleotide sites, including gaps.The most appropriate evolutionary models estimated for each part of the alignments were ITS1: TPM2uf+G, 5.8S: TPM2, ITS2: TPM2uf+G, and LSU: TIM3+G.
The bootstrapping criteria from the ML analysis stopped after 50 replicates.The most likely tree generated with the ML analysis is shown in Figure 3.The family Cyphellopsidaceae (100% BS, 1.0 BPP) is represented by 19 terminals, with emphasis on the Merismodes clade (100% BS, 1.0 BBP), represented by 12 terminals of Maireina and Merismodes.Consistent with the other previous analyses, both the Bayesian inference and ML analysis recover Maireina as a paraphyletic group (Figure 3).Merismodes.Consistent with the other previous analyses, both the Bayesian inference and ML analysis recover Maireina as a paraphyletic group (Figure 3).or herbarium number as well as GenBank accession numbers and country of origin follow taxon name.Thicker lines represent branches with maximum bootstrap values and posterior probabilities (100% BS, 1.0 BPP).Bootstrap values and Bayesian posterior probabilities are indicated if they are equal to or greater than 70%, and 0.95, respectively.The scale bar represents the expected number of nucleotide changes per site.
Notes: After the very brief protologue, Knudsen and Vesterholt [61] included in their description of Merismodes include some morphological characteristics of the genera Maireina, Cyphellopsis, and Phaeocyphellopsis.In our emendation, we include additional distinctive morphological characteristics of the species recently described [52][53][54] and of Maireina based on Bodensteiner [57].In all our analyses, the genus Maireina is resolved as paraphyletic, forming a well-supported monophyletic lineage with the sequences of Mersimodes included.Based on these results and those of previous investigators [61], we consider Maireina a synonym of the latter genus and propose an amendment.The name Merismodes, proposed in 1909 [79], has priority against Maireina erected in 1961 [43].Thus, to better accommodate the Maireina species sampled in our analyses (which includes sequences from holotype material), we propose the combination of Ma. filipendula and Ma.subsphaerosphora in Merismodes.Additionally, a recently collected sample of Ma. monacha (type species of Maireina) from France (same country locality of the holotype) was also included in our analyses.The taxon is herein re-analyzed and confirmed in Merismodes and an epitype is designated.(Fagaceae) [57].Distributed in Europe and Oceania [43].

Merismodes monacha (
Notes: Our specimen agrees with the description of Me. monacha presented by Cooke (ref.[43], as Ma.monacha), who analyzed authentic material of all names included here as synonyms, including the types of Cyphella obscura Roum.and Cyphella sydowii Bres.According to Cooke [43], Me. monacha is characterized by brown receptacles with long hairs around the cup edge and at the hymenial surface, elongate to cylindrical basidiospores, four-spored basidia, and cylindrical, yellowish brown to brown external hyphae with paler apices.Although our material has had slightly broader receptacles (1.5-3 mm diam.) and basidia (9.0-110 µm diam.)than reported by Cooke [43] (receptacles, 0.5-1 mm diam.; basidia, 5.5-8.0 µm diam.), other macro-and micromorphological characteristics are sufficient for the identification of this sample as Ma.monacha sensu Cooke [43] and Bodensteiner [57].Merismodes monacha was originally described from samples collected in France but it has a distribution recorded in many European countries, including Germany, Austria, Italy, the Czech Republic, Hungary, the United Kingdom, and one record from Australia [43].The holotype of Cyphella monacha Speg.[anon.s.n.(Fung.Gall.768) Spegazzini s.n.] was deposited at the New York Botanical Garden Herbarium (NY).Considering the complete morphological and molecular data recovered from our sample that is from a region close to the type locality, we decided to designate the voucher ALV30536 as epitypus.Macro-and micro-morphological description: see Mombert [55].

MycoBank: MB 849403
Etymology: Eos = light of day; the goddess of dawn (Greek); cyphella (from kyfos in Greek) = shape of a cup, something hollow.The prefix "Eos" is in reference to the light emitted by the bioluminescent basidiomata of the type species.Additionally, the Roman equivalent refers to Eosforos as Lucifer, which is the entity's name that was later considered into Christianity as the devil, and it also refers to the name of the protected area (Devil's Cave State Park) near where the specimens of the type species were found.The name cyphella is a reference to the genus Cyphella and to the cyphelloid body form.
Type species: Eoscyphella luciurceolata Silva-Filho, Stevani & Desjardin (described below).Diagnosis: Eoscyphella is morphologically similar to Merismodes and Woldmaria but differs from Woldmaria in lacking conspicuous long hairs in the receptacle, subglobose to broadly ellipsoid basidiospores, regularly bi-spored basidia, and unclamped hyphae; and from Merismodes by the absence of conspicuous hairs in the receptacle, absence of cystidia, regularly bi-spored basidia, and the characteristic external hyphae that are always pigmented and encrusted at the tips.
Notes: Eoscyphella, typified here using Eoscyphella luciurceolata sp.nov., represents a new lineage of bioluminescent fungi.It is supported with phylogenetic data (Figures 1a, 2 and 3) and morphological characteristics, including the absence of conspicuous long hairs on the receptacle, subglobose to broadly ellipsoid basidiospores, regularly bi-spored basidia, the absence of clamp connections, and the consistent presence of pigmented and encrusted external hyphae.An additional collection (FIPBIO 01) of a related non-bioluminescent cyphelloid species was found in the same region of the type species.The ITS rDNA sequence data (OR260255) resolves this taxon as sister to E. luciurceolata and suggests that it represents an additional species of Eoscyphella (Figures 2 and 3).The presence of a second species indicates that Eoscyphella is likely a non-monospecific genus that includes both bioluminescent and non-bioluminescent members.Until additional material of the non-bioluminescent taxon can be collected to confirm these initial observations, we prefer to leave it undescribed.
Habitat and known distribution: On bark of "fumeiro" tree (Solanum swartzianum) in the Atlantic Rainforest, southern Brazil.Known only from the type locality.

Discussion
The morphological delimitation of Merismodes, Cyphellopsis, and Maireina has been the cause of debates about the morphological limits of these genera [43,51,57,62,[80][81][82][83].Reid [81] considered Cyphellopsis and Maireina as synonyms and suggested that the depth of the cavity that lined the hymenium is a character insufficient for the separation of Cyphellopsis (=Maireina) and Merismodes.Singer [83] synonymized the genus Cyphellopsis and Maireina with Merismodes and listed both Maireina and Cyphellopsis as sections.The first broad research on cyphelloid fungi based on molecular phylogenetic analyses resolved Merismodes and Cyphellopsis as a monophyletic group, recognizing them as a single genus [45].Another broad study of Maireina without molecular data led Bodensterner [53,57] to recognize the genus Maireina as an independent lineage from Merismodes and Cyphellopsis.Knudsen and Vesterholt [61] recognized Cyphellopsis, Maireina, and Phaeocyphellopsis as synonyms of Merismodes, providing a broad description for the genus.The first works to describe new species of Maireina based in-part on molecular data are those of Laessøe et al. [54] and Mombert [55].In both, the sequences of Maireina clustered with Merismodes and Cyphellopsis in a large clade, making it possible to determine the phylogenetic position within Cyphellopsidaceae.Our phylogenetic analyses in separate and combined LSU rDNA and ITS rDNA recognized Merismodes, Cyphellopsis, and Maireina as a monophyletic group, supporting the proposal of Knudsen and Vesterholt [61] for a broad morphological concept of Merismodes.The samples and sequences of Me. monacha, type species of Maireina, first studied by Mombert [55] were extremely important for the recognition and the phylogenetic positioning of the genus Maireina.Although the sequences are not of the holotype specimen, the collection is from a region very close to the type locality, and the morphological description agrees with the complete redescription presented by Cooke [43].
Our cyphelloid bioluminescent samples were initially identified within the morphological concept of Maireina sensu Bodensteiner [53,57].However, our phylograms (Figures 1a, 2 and 3) showed a phylogenetic distance between E. luciurceolata and Me.monacha, which are only 90.6% to 90.7% similar in the LSU rDNA and 64.6% to 65.9% similar in the ITS rDNA.Eoscyphella is closely related to the genus Woldmaria in our analyses, but the included taxa are 7.3% to 7.4% divergent in their LSU rDNA sequences, a high value considering a similarity threshold of around 96.91% to discriminate genera using LSU rDNA in Basidiomycota [84].These data and results support the proposition of a new cyphelloid genus and distinct molecular lineage.Additionally, Eoscyphella is also morphologically well delimited with receptacles that lack conspicuous long hairs, subglobose to broadly ellipsoid basidiospores, frequently bi-spored basidia, unclamped hyphae, and weakly to densely incrusted overall external hyphae, which are always pigmented and encrusted at the tips.
Regarding the cyphelloid genera within Agaricomycetes, our LSU rDNA analyses retrieved 11 lineages of cyphelloid fungi and the phylogenetic relationship of the cyphelloid genera agrees with recent phylogenetic studies [45,50,56].However, we highlight that sequences of the collection PB327 named as Calathella columbiana appear in different positions and for this reason were excluded from the combined analyses: in the ITS rDNA tree within Cyphellopsidaceae (Figure 2), and in the LSU rDNA tree (Figure 1c) in a clade close to representatives of Entolomataceae.Additionally, Phaeosolenia densa (Berk.)W.B. Cooke was shown by Bodensteiner et al. [45] to be close to the genus Tubaria (W.G.Sm.) Gillet, whilst in our analyses, it forms an isolated clade without support (Figure 1c).Desjardin et al. [4] performed the second review of bioluminescent fungi worldwide, referring 64 luminescent species into three lineages, Armillaria, Mycenoid, and Omphalotus, indicating that Gerronema viridilucens and Mycena lucentipes do not belong to the Mycenoid lineage.Later, Desjardin et al. [24] referred G. viridilucens and M. lucentipes to a new and unnamed lineage, which was later named the Lucentipes lineage by Oliveira et al. [35].Our LSU rDNA phylogram (Figure 1b) shows and confirms G. viridilucens plus M. lucentipes as a separate bioluminescent lineage.The Eoscyphella lineage is here recognized as a new and fifth bioluminescent lineage in Cyphellopsidaceae (Figure 1a).

Conclusions
Our systematic study confirms the findings of previous studies regarding the existence of multiple bioluminescent lineages in Agaricales.All bioluminescent fungi have currently been described in suborder Marasmiineae Aime, Dentinger & Gaya.The newly described Eoscyphella luciurceolata was confirmed from molecular phylogenies in the family Cyphellopsidaceae, currently accepted within the suborder Schizophyllineaeae Aime, Dentinger & Gaya [62].Additionally, our study reveals a new lineage within a group primarily consisting of reduced forms.Fungal bioluminescence engages in a cyclical process of biosynthesis known as the Caffeic Acid Cycle (CAC), which relies on a sequence of four consecutive enzymes: hispidin synthase (HispS), hispidin-3-hydroxylase (H3H), luciferase (Luz), and caffeylpyruvate hydrolase (CPH) [3].At present, there are limited genomic data concerning bioluminescent fungi in the existing literature [85], with the majority originating from the Mycenoid and Armillaria lineages.By identifying this recently discovered bioluminescent lineage and uncovering the sequences of the hisps, h3h, luz, and cph genes, there is potential for enhancing our understanding of the evolutionary progression of the bioluminescent trait within the fungal kingdom.
A high diversity of bioluminescent fungi has been discovered in Brazil, with 23 species (including our new described species) reported theretofore, see [86].In the Brazilian Atlantic Rainforest, new species of bioluminescent fungi have been described or reported, e.g., [87], with emphasis to the southwestern portion of the state of São Paulo, the same area where E. luciurceolata was found and where another 12 species of Mycenoid and Lucentipes lineage taxa have already been described or reported [22][23][24][25].Even so, new bioluminescent samples collected at the same area are currently in the process of molecular and morphological characterization and may represent taxonomic novelties, demonstrating that the Atlantic Rainforest in the southwestern region of the São Paulo state is one of the most studied areas of bioluminescent fungi and may represent a biodiversity hot spot for these organisms.

2 . 1 . 1 .
Brazilian Site of the New Luminescent Taxon Basidiomata of the new bioluminescent taxon were collected during expeditions to the Atlantic Rainforest in the municipality of Eldorado, state of São Paulo, Brazil.More specifically, at a 546 m altitude and 500 m west of the entrance to the "Caverna do Diabo" (Devil's Cave) State Park at coordinates 24 • 38 14.0100 S and 48 • 24 37.6812 W. The climate there is classified as humid subtropical, and the mean annual temperatures are usually between 20 and 22 • C and have a high pluviometric index, with average annual rainfall ranging from 1500 to 2000 mm [63]

Figure 1 .
Figure 1.(a-d) The ML phylogeny of representative collections of Agaricomycet complete LSU rDNA.Voucher/strain/clone or herbarium number as well as GenBa numbers and country of origin follow taxon name.Cyphelloid species are noted with t and bioluminescent species with the symbol •.The new species is highlighted in luminescent lineages are in gray.Thicker lines represent branches with maximum boo and posterior probabilities (100% BS, 1.0 BPP).Bootstrap values and Bayesian posterior are indicated if they are equal to or greater than 70%, and 0.95, respectively.The scale b the expected number of nucleotide changes per site.

Figure 1 .Figure 1 .
Figure 1.(a-d) The ML phylogeny of representative collections of Agaricomycetes based on complete LSU rDNA.Voucher/strain/clone or herbarium number as well as GenBank accession numbers and country of origin follow taxon name.Cyphelloid species are noted with the symbol

Figure 1 .
Figure 1.(a-d) The ML phylogeny of representative collections of Agaricomycetes based on complete LSU rDNA.Voucher/strain/clone or herbarium number as well as GenBank accession numbers and country of origin follow taxon name.Cyphelloid species are noted with the symbol ◆ and bioluminescent species with the symbol •.The new species is highlighted in red, and the luminescent lineages are in gray.Thicker lines represent branches with maximum bootstrap values and posterior probabilities (100% BS, 1.0 BPP).Bootstrap values and Bayesian posterior probabilities are indicated if they are equal to or greater than 70%, and 0.95, respectively.The scale bar represents the expected number of nucleotide changes per site.

.
The new species is highlighted in red, and the luminescent lineages are in gray.Thicker lines represent branches with maximum bootstrap values and posterior probabilities (100% BS, 1.0 BPP).Bootstrap values and Bayesian posterior probabilities are indicated if they are equal to or greater than 70%, and 0.95, respectively.The scale bar represents the expected number of nucleotide changes per site.

Figure 2 .
Figure 2. ML phylogeny of collections of Cyphellopsidaceae representatives based on complete ITS rDNA.The new species is highlighted in red.Voucher/strain/clone or herbarium number as well as GenBank accession numbers and country of origin follow taxon name.Thicker lines represent branches with maximum bootstrap values and posterior probabilities (100% BS, 1.0 BPP).Bootstrap values and Bayesian posterior probabilities are indicated if they are equal to or greater than 70%, and 0.95, respectively.The scale bar represents the expected number of nucleotide changes per site.The bootstrapping criteria from the ML analysis stopped after 50 replicates.The most likely tree generated with the ML analysis is shown in Figure 3.The family Cyphellopsidaceae (100% BS, 1.0 BPP) is represented by 19 terminals, with emphasis on the Merismodes clade (100% BS, 1.0 BBP), represented by 12 terminals of Maireina and

Figure 2 .
Figure 2. ML phylogeny of collections of Cyphellopsidaceae representatives based on complete ITS rDNA.The new species is highlighted in red.Voucher/strain/clone or herbarium number as well as GenBank accession numbers and country of origin follow taxon name.Thicker lines represent branches with maximum bootstrap values and posterior probabilities (100% BS, 1.0 BPP).Bootstrap values and Bayesian posterior probabilities are indicated if they are equal to or greater than 70%, and 0.95, respectively.The scale bar represents the expected number of nucleotide changes per site.

Figure 3 .Figure 3 .
Figure 3. ML phylogeny of Cyphellopsidaceae focusing on collections of Merismodes representatives based on combined ITS rDNA and LSU rDNA.The new species is highlighted in red.Voucher/strain/clone or herbarium number as well as GenBank accession numbers and country of origin follow taxon name.Thicker lines represent branches with maximum bootstrap values and posterior probabilities (100% BS, 1.0 BPP).Bootstrap values and Bayesian posterior probabilities are indicated if they are equal to or greater than 70%, and 0.95, respectively.The scale bar represents the expected number of nucleotide changes per site.3.2.Taxonomic Part

Figure 6 .
Figure 6.Eoscyphella luciurceolata basidiomata in light (above) and dark (below) on removed bark of "fumeiro" tree (Solanum swartzianum).Note that mushrooms are in wetter conditions and all of them emit light.(a) A dried mushroom is shown next to a scalpel blade to demonstrate its size; (b) FBIPBio 93.20220802 (Paratype-FIFUNGI00249). Photos by Adão Henrique Rosa Domingos and Isaias Santos.

Figure 6 .
Figure 6.Eoscyphella luciurceolata basidiomata in light (above) and dark (below) on removed bark of "fumeiro" tree (Solanum swartzianum).Note that mushrooms are in wetter conditions and all of them emit light.(a) A dried mushroom is shown next to a scalpel blade to demonstrate its size; (b) FBIPBio 93.20220802 (Paratype-FIFUNGI00249). Photos by Adão Henrique Rosa Domingos and Isaias Santos.

Figure 6 .
Figure 6.Eoscyphella luciurceolata basidiomata in light (above) and dark (below) on removed bark of "fumeiro" tree (Solanum swartzianum).Note that mushrooms are in wetter conditions and all of them emit light.(a) A dried mushroom is shown next to a scalpel blade to demonstrate its size; (b) FBIPBio 93.20220802 (Paratype-FIFUNGI00249). Photos by Adão Henrique Rosa Domingos and Isaias Santos.
, who considered Maireina, Cyphellopsis, and Phaeocyphellopsis W.B. Cooke synonyms of Merismodes.We herein propose the combination of Ma. monacha, Ma. filipendula, and Ma.subsphaerophora into Merismodes, as well as the description of the genus Eoscyphella to accommodate the novel bioluminescent cyphelloid species from Brazil.