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Phylogeny of Megasporoporia s.lat. and related genera of Poyporaceae: New genera, new species and new combinations

Several resupinate poroid Polyporaceae with dextrinoid skeletal hyphae and cylindrical, thinwalled basidiospores have been placed in Dichomitus and Megasporoporia. With the inclusion of DNA sequences, mostly from Chinese material, in the phylogeny of the genera, new genera were recognized, i.e., Megasporoporiella and Megasporia. In the current study, Jorgewrightia and Mariorajchenbergia are described as new genera in the Polyporaceae based on phylogenetic analyses of four gene regions: nuc rDNA ITS1-5.8S-ITS (ITS) and partial Large Subunit (28S), Translation Elongation Factor 1-alpha (TEF1), and RNA polymerase II second largest subunit (RPB2), mostly from Brazilian material. These new genera currently accommodate species formerly placed in Cerioporus, Dichomitus, Megasporia, Megasporoporiella and Pachykytospora. Dichomitus, Jorgewrightia, Mariorajchenbergia, Megasporia and Megasporoporia have mostly resupinate pale-colored basidiomata, poroid hymenophore, usually dextrinoid branched skeletal hyphae, and cylindrical basidiospores. These genera are difficult to differentiate on morphology alone and analyses based on sequences from at least two regions (ITS and 28S), as well as information about geographical distribution, are needed to separate them. Megasporia variabilicolor and Megasporoporia neosetulosa are described as new species, and one neotype and 20 new combinations are proposed. Synoptic tables including characteristics for 27 species from the five genera is presented.


DNA extraction, PCR amplification and sequencing
Fragments (1-2 mg) from the basidiomata were removed and placed in tubes of 1.5 ml and stored at -20°C until extraction. The DNA was extracted according to the method described in Goés -Neto et al. (2005). The nuc rDNA Internal Transcribed Spacer (ITS) and partial Large Subunit (nLSU), the Translation Elongation Factor 1-alpha (TEF1) and the RNA Polymerase II Second Largest Subunit (RPB2) were amplified for the sequences generated here using the primer pairs ITS4-ITS5, LR0R-LR5, RPB2 5F-RPB2 7.1R and Ef DF-EF1 2218 R, respectively (White et al. 1990, Moncalvo et al. 2000, Rehner & Buckley 2005, Frøslev et al. 2005, Smith & Sivasithamparam 2000. Negative controls containing all components of the reaction mix, but exchanging DNA by water, were used in each procedure to detect possible contamination. The amplification products were purified with GenJET PCR Purification Kit (Thermo Scientific) and sequenced at the Plataforma Tecnológica de Genômica e Expressão Gênica do Centro de Biociências, UFPE, Brazil, and at the Stab Vida, Portugal, both using the Sanger method. Cycle sequencing was carried out with the same primers as the amplification reactions (Moncalvo et al. 2000). All obtained sequences were deposited in GenBank (National Center for Biotechnology Information, Maryland, USA) (Table 1).
Phylogenetic reconstructions were performed using Maximum Likelihood (ML) and Bayesian Inference (BI) analyses. Nucleotide substitution models for each gene fragment were estimated based on Bayesian Information Criterion (BIC) on W-IQ-TREE (Kalyaanamoorthy et al. 2017) and used for ML and BI analyses.

Phylogeny
Sixteen specimens were sequenced, generating 14 ITS, 14 nLSU, seven RPB2 and seven TEF1 sequences. The concatenated alignment resulted in 3012 positions including gaps, of which 1652 are constant sites and 1191 parsimony-informative. The best-fit models selected were TPM2u+F+I+G4 for ITS, SYM+I+G4 for nLSU, TN+F+I+G4 for RPB2, and TIMe+I+G4 for TEF1.
Five different datasets, one for each locus and the concatenated one, were analyzed using ML and BI analyses. No strong conflicts were detected among the datasets, consequently only the analyses from the concatenated data set are presented here. The results of the phylogenetic analyses generated from ML and BI showed similar tree topologies. Thus, the BI trees with bootstrap support values (BS), UFBoot2 (UB) and posterior probabilities (PP) were used to show the results (Fig. 1).
The newly generated sequences are placed in two clades of Polyporaceae: three in Megasporoporia s.str. and nine in a redefined Megasporia, both with strong support (BS = 99.9%; UB = 83%; PP = --and BS = 99.5%; UB = 100; PP = 0.91, respectively). The results also support the description of two new species (Fig. 1). In addition, several sequences of species previously placed in Cerioporus, Dichomitus, Megasporia, Megasporoporiella and Pachykytospora are recovered in two strongly supported clades that could not be assigned to known taxa of Polyporaceae ( Fig. 1) and are described below as new genera. Li & Cui (2013a) typified Megasporoporiella with M. cavernulosa based on the placement of a 28S sequence from Chinese material identified as M. cavernulosa (Wu 9508-328). However, the type of M. cavernulosa originated from Brazilian Amazonia. In our analyses, the ITS, 28S and TEF1 sequences of what we consider the true M. cavernulosa recently collected in Brazilian Amazonia were placed in the Megasporia clade (Fig. 1). This makes Megasporoporiella a synonym of Megasporia and renders nameless the clade that corresponds to Megasporoporiella sensu Li & Cui (2013a). Recently, Wang et al. (2021), without further notice, selected M. pseudocavernulosa as the type species of Megasporoporiella, because they had discovered that the type specimen (Wu 9508-328) selected by Li & Cui (2013a) actually represents M. subcavernulosa. This typification does not follow the procedures for changing a genus type outlined in the International Code of Nomenclature for algae, fungi, and plants (Turland 2018), neither has the typification been registered, as required by the same Code. Consequently, the solution suggested by Wang et al. (2021) for Megasporoporiella is not valid. Instead, the new genus Mariorajchenbergia is described below to include M. australiae, M. hubeiensis, M. pseudocavernulosa, M. rhododendri and M. subcavernulosa.
Megasporia was typified by M. hexagonoides Li & Cui (2013a), and the species was represented in the phylogeny by sequences from Chinese collections (He 2608 and Cui 6592). The type specimen of M. hexagonoides, however, is from Argentina and the inclusion of an Argentinean sequence of M. hexagonoides (CBS 464.63, deposited by J.E. Wright) revealed that the clade that corresponds to Megasporia is a different one (Fig. 1). The species placed in Megasporia sensu Li & Cui (2013a) are now referred to the new genus Jorgewrightia including J. bambusae, J. cystidiolophora, J. ellipsoidea, J. fusiformis, J. guangdongensis, J. major, J. rimosa, J. hengduanensis, J. tropica, J. violacea and J. yunnanensis as well the Chinese specimens misidentified as M. hexagonoides (He 2608, Cui 6592, Cui 13853 and Cui 13855), whose identity should be re-examined (Fig. 1).
The addition of the Argentinean sequence of M. hexagonoides also indicated that a revised Megasporia includes the neotropical M. amazonica, M. anoectopora, M. cavernulosa, M. cylindrospora, M. hexagonoides and the new species M. variabilicolor (described below) (Fig. 1).
At last, Megasporoporia sensu Li & Cui (2013a) included a specimen identified as M. setulosa (MG38, only nLSU), as well as M. bannaensis and M. minor, all from China. With the inclusion of the neotype of M. setulosa (designated below), collected in north Tanzania close to the type locality, Megasporoporia currently includes, besides the neotype and the Chinese species, the neotropical, new species M. neosetulosa (described below) (Fig. 1). Also, the identity of M. setulosa (MG38) should be re-examined. Observations -According to Dai et al. (2004), this species is characterized by the lack of hyphal pegs, dendrohyphidia and polyhedric crystals. In addition, generative hyphae dominate the tramal structure, the basidiospores are ellipsoid and it grows in boreal forests on fallen trunks and dead trees of Rhododendron. Observations -According to Dai & Wu (2004), this species has both hyphal pegs and dcndrohyphidia, and basidiospores distinctly smaller than in Megasporia cavernulosa.
Observations -The description above is updated from Megasporoporia sensu Ryvarden et al. (1982), typified by Poria setulosa Henn., collected in Usambara, Tanga Region in Tanzania, and deposited at herbarium B. Ryvarden et al. (1982) stated that the type is lost, probably during World War II. Upon our request, R. Lücking [current curator for cryptogams (lichens, fungi, bryophytes) in herbarium B, pers. comm.] confirmed that the material or similar is absent from herbarium B. Material identified as Megasporoporia setulosa (Henn.) Rajchenb. and also collected in north Tanzania had its DNA sequenced and selected here to be the neotype of P. setulosa.
Megasporoporia is characterized by strongly dextrinoid, cyanophilous hyphae, which are also unbranched to sparingly branched, lack of dendrohyphidia, presence of hyphal pegs in most of species, and tropical and subtropical distribution. Basidiomata annual, resupinate, easily detachable from the substrate, 1.8-5.1 × 0.4-1.6 × 0.1-0.2 mm. Pore surface beige to cream (52 Buff), angular pores, 1.5-2 per mm, dissepiments thin and entire. Margin sterile, up to 1 mm width, lighter than the pore surface (6 F). Context very thin, invisible to naked eye, 0.1-0.2 mm. Tubes concolorous to the pore surface, up to 2 mm deep.