Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae (Polyporales, Basidiomycota), with an emphasis on Ceriporia and Meripilus including ten new species
Abstract
Species in Ceriporia and Meripilus are important wood-decaying fungi causing white rot on both angiosperm and gymnosperm wood. Morphologically, the two genera share similar micromorphology: a monomitic hyphal system of cyanophilous generative hyphae bearing simple septa. Phylogenetic and morphological analyses of Ceriporia and other related genera in Irpicaceae and Meripilus in Meripilaceae were carried out. Ceriporia is characterized by mostly resupinate basidiomata with a white to brightly colored pore surface when fresh, usually without changing color when bruised, and cylindrical to allantoid basidiospores. Meripilus is similar to Ceriporia, but it has resupinate, effused-reflexed to pileate basidiomata, sometimes with an erubescent pore surface when bruised, and ellipsoid to globose basidiospores. Phylogenies of species in the two genera were reconstructed with multiple-loci DNA sequences, including ITS, nLSU, nSSU, TEF1, RPB1, and RPB2, as well as two combined datasets: ITS+nLSU+TEF1+RPB1+nSSU for Ceriporia and ITS+nLSU+TEF1+RPB2+nSSU for Meripilus. Three new species of Ceriporia, one new species of Meruliopsis, and six new species of Meripilus are described and illustrated. Moreover, the evolutionary times of the Polyporales, including Irpicaceae and Meripilaceae, were revealed based on the conserved regions of three-loci DNA sequences (ITS+nLSU+TEF1). Irpicaceae and Meripilaceae are estimated to have emerged at the junction of the early and late Cretaceous, with mean crown ages of 108.9 Myr and 97.23 Myr, respectively. Bayesian evolutionary analysis shows that the divergence of Ceriporia emerged with a mean crown age of 83.61 Myr [95% highest posterior density (HPD): 65.25–106.35 Myr], which occurred during the late Cretaceous; the initial diversification of Meripilus also occurred during the late Cretaceous, with a mean crown age of 81.38 Myr [95% HPD: 61.89–105.78 Myr].
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1 Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae (Polyporales, Basidiomycota), with an emphasis on Ceriporia and Meripilus including ten new species Chao-Ge Wang1* , Ying-Da Wu2* , Xin Zhang1, Yu-Cheng Dai1, Zhen-Hao Li3,4, Yuan Yuan1 1 StateKeyLaboratoryofEfficientProductionofForestResources,SchoolofEcologyandNatureConservation,BeijingForestryUniversity,Beijing100083,China 2 KeyLaboratoryofForestandGrasslandFireRiskPrevention,MinistryofEmergencyManagement,ChinaFireandRescueInstitute,Beijing102202,China 3 ZhejiangKeyLaboratoryofBiologicalBreedingandExploitationofEdibleandMedicinalMushrooms,Jinhua321200,Zhejiang,China 4 ZhejiangShouxianguPharmaceuticalCo.,Ltd,Jinhua321000,Zhejiang,China Correspondingauthor:YuanYuan([email protected]) Copyright: © Chao-Ge Wang et al. This is an open access article distributed under terms of the Creative Commons Attribution License (Attribution 4.0 International – CC BY 4.0). Research Article Abstract Species in Ceriporia and Meripilus are important wood-decaying fungi causing white rot on both angiosperm and gymnosperm wood. Morphologically, the two genera share similar micromorphology: a monomitic hyphal system of cyanophilous generative hyphae bearing simple septa. Phylogenetic and morphological analyses of Ceriporia and other related genera in Irpicaceae and Meripilus in Meripilaceae were carried out. Ceriporia is characterized by mostly resupinate basidiomata with a white to brightly colored pore surface when fresh, usually without changing color when bruised, and cylindrical to allantoid basidiospores. Meripilus is similar to Ceriporia, but it has resupinate, effused-reflexed to pileate basidiomata, sometimes with an erubescent pore surface when bruised, and ellipsoid to globose basidiospores. Phylogenies of species in the two genera were reconstructed with multiple-loci DNA sequences, including ITS, nLSU, nSSU, TEF1, RPB1, and RPB2, as well as two combined datasets: ITS+nLSU+TEF1+RPB1+nSSU for Ceriporia and ITS+nLSU+TEF1+RPB2+nSSU for Meripilus. Three new species of Ceriporia, one new species of Meruliopsis, and six new species of Meripilus are described and illustrated. Moreover, the evolutionary times of the Polyporales, including Irpicaceae and Meripilaceae, were revealed based on the conserved regions of three-loci DNA sequences (ITS+nLSU+TEF1). Irpicaceae and Meripilaceae are estimated to have emerged at the junction of the early and late Cretaceous, with mean crown ages of 108.9 Myr and 97.23 Myr, respectively. Bayesian evolutionary analysis shows that the divergence of Ceriporia emerged with a mean crown age of 83.61 Myr [95% highest posterior density (HPD): 65.25–106.35 Myr], which occurred during the late Cretaceous; the initial diversification of Meripilus also occurred during the late Cretaceous, with a mean crown age of 81.38 Myr [95% HPD: 61.89–105.78 Myr]. Key words: Meruliopsis, molecular clock, Physisporinus, taxonomy, wood-decaying fungi Academic editor: Francisco Kuhar Received: 6 June 2025 Accepted: 17 September 2025 Published: 15 October 2025 Citation: Wang C-G, Wu Y-D, Zhang X, Dai Y-C, Li Z-H, Yuan Y (2025) Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae (Polyporales, Basidiomycota), with an emphasis on Ceriporia and Meripilus including ten new species. IMA Fungus 16: e161336. https://doi. org/10.3897/imafungus.16.161336 IMA Fungus 16: e161336 (2025) DOI: 10.3897/imafungus.16.161336 * Theseauthorscontributedequallyasfirstauthorstothiswork.
2 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae Introduction Wood-decaying fungi are capable of causing white or brown rot in wood, playing a crucial role as decomposers in forest ecosystems. Polyporales is one of the main orders of wood-decaying fungi, including more than 15 families (Justo et al. 2017). Many studies focus on certain families in the Polyporales, such as Polyporaceae Fr. ex Corda, Fomitopsidaceae Jülich, and Ganodermataceae Donk, which contain many taxa and have high medicinal and ecological value (Ji et al. 2022; Sun et al. 2022; Liu et al. 2023; Spirin et al. 2024). In contrast, Irpicaceae Spirin & Zmitr., especially the poroid species, and Meripilaceae Jülich mostly have unimpressive basidiomata with light-colored hymenophores, and thus they have received less research attention. Irpicaceae,typifiedbyIrpex Fr., was established by Spirin in 2003 (Spirin 2003). It belongs to the phlebioid clade within the Polyporales Gäum. and includes thirteen genera, viz., Byssomerulius Parmasto, Ceriporia Donk, Crystallicutis El-Gharabawy, Cytidiella Pouzar, Efibula Sheng H. Wu, Leal-Dutra & G.W. Griff., Gloeoporus Mont., Irpex, Leptoporus Quél., Meruliopsis Bondartsev, Phanerochaetella C.C. Chen & Sheng H. Wu, Raduliporus Spirin & Zmitr., Resiniporus Zmitr., and Trametopsis Tomšovský, with variable hymenophores, including corticioid and irpicoid species, and resupinate to pileate polypores (Chen et al. 2021). Previous phylogenetic studies (Binder et al. 2013; Justo et al. 2017; Chen et al. 2020, 2021) determined that the genus Ceriporia is polyphyletic. Three species— Phanerochaete allantospora Burds. & Gilb., Candelabrochaete langloisii (Pat.) Boidin, and Candelabrochaete septocystidia (Burt) Burds.—were proposed to be combined into Ceriporia based on similar morphology and correlative phylogeny (Wang et al. 2023). Although two species accepted in Leptoporus Quél., L. mollis (Pers.) Quél. and L. submollis B.K. Cui & Shun Liu, are nested in the Ceriporia clade, the genus causes brown rot in dead gymnosperm wood, while Ceriporia causes white rot in dead angiosperm and gymnosperm wood (Ryvarden and Gilbertson 1993; Chen et al. 2020; Liu et al. 2023; Wang et al. 2023). Therefore, Leptoporus and Ceriporia represent two separate genera because of their different ecologicalhabits(Wangetal.2023).Wangetal.(2023)clarifiedtherelationship between Ceriporia and related genera by reconstructing the phylogenies of Irpicaceaeusingfive-lociDNAsequencesandamendedthedefinitionofCeriporia. Meripilaceae was established in 1982 as belonging to the Polyporales and contains three genera, viz., Meripilus P. Karst., Physisporinus P. Karst., and Spongipellis Pat. (Wang and Dai 2022). Wang et al. (2024) discussed the phylogenetic relationship of Meripilus and Physisporinus using multigene phylogenetic analyses and indicated that Meripilus is a separate genus nested in the Physisporinus clade. In addition, the status of three confusing species—Physisporinus vitreus (Pers.) P. Karst., P. sanguinolentus (Alb. & Schwein.) Pilát, and P. expallescens (P. Karst.) Pilát—was discussed (Wang et al. 2024). Recently, Westphalen et al. (2025) transferred all species of Physisporinus into Meripilus to propose a monophyletic classification of genera within Meripilaceae. In addition, Henningsia Möller was also combined into Meripilus based on morphological and phylogenetic evidence (Westphalen et al. 2025). Spongipellis traditionally contains four species: S. delectans (Peck) Murrill, S. litschaueri Lohwag, S. spumeus (Sowerby) Pat., and S. unicolor (Fr.) Murrill. The type species, S. spumeus, formed a separate clade addressed in Meripilaceae (Spirin et al. 2022; Wang and Dai 2022; Miettinen
3 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae et al. 2023), while differing from the other three species nested in Cerrenaceae (Wang and Dai 2022; Spirin et al. 2022; Miettinen et al. 2023). Morphologically, Ceriporia has resupinate basidiomata with a white to brightly colored pore surface when fresh, usually without changing color when bruised; a monomitic hyphal system of cyanophilous generative hyphae bearing simple septa; and cylindrical to allantoid, hyaline, thin-walled basidiospores that are negative in Melzer’s reagent and Cotton Blue (Wang et al. 2023). Meripilus is characterized byannual,resupinate,effused-reflexedtopileatebasidiomata,sometimeswitha white pore surface when fresh and erubescent when bruised, or with a brightly colored pore surface; a monomitic hyphal system of cyanophilous generative hyphae bearing simple septa; and ellipsoid to globose, hyaline, thinto slightly thick-walled basidiospores that are negative in Melzer’s reagent and mostly negative in Cotton Blue (Wang et al. 2024). Thus, much of the micromorphology of Ceriporia and Meripilusoverlaps,andsometimesitisdifficulttodistinguishthemmacroscopically. Comparative analyses of nucleotide sequences have concluded that molecular evolution occurs at a relatively constant rate, thereby giving rise to the concept of the molecular clock (Zuckerkandl 1962; Zuckerkandl and Pauling 1965). By using fossil records or geological events as calibration points in conjunction with phylogenetic trees as the foundational framework, estimation of divergence times among different taxonomic groups has become a widely adopted methodology in evolutionary biology (Avise and Johns 1999; Yang et al. 2007; Havill et al. 2008; Lockwood et al. 2013; Zhao et al. 2016, 2017; Varga et al.2019;Jietal.2022).Moleculardivergencetimesoffungishouldreflecttheir different taxonomic ranks according to the geological ages when they evolved (Hennig 1966; Zhao et al. 2016). Ceriporia and Meripilus belong to different families, Irpicaceae and Meripilaceae, respectively, but have similar macroscopic morphological characteristics. In addition, the molecular divergence times of Ceriporia and Meripilus, and of Irpicaceae and Meripilaceae, need to be analyzed in an attempt to reveal the reasons for differences between them. Poroid fungi in the Irpicaceae, especially Ceriporia and related genera, including Meruliopsis and Gloeoporus, have long been neglected. Early studies (Jia et al. 2014; Miettinenetal.2016;Yuanetal.2017)insufficientlyaddressedspeciesdiversity andtaxonomicclassificationuntilChenetal.(2020)provedtheindependenceof Ceriporia and related genera by phylogenetic studies and sorted out the taxonomic statusoftherelevantspecies.Wangetal.(2023)clarifiedthetaxonomyandphylogeny of Ceriporia and other related taxa in Irpicaceae with more specimens from aroundtheworld,especiallyfromChina,amendedtheirdefinitions,studiedmorphologically confusing species, and speculated on the divergence time of Irpicaceae. Meripilaceae, as a small family in Polyporales, included two unimpressive genera, Meripilus and Spongipellis (Wang and Dai 2022; Wang et al. 2024; Westphalen et al. 2025). Although phylogenetic and morphological analyses of Meripilaceae have been carried out (Wang et al. 2024), more relevant specimens are being continuously collected in China, and there is a lack of research on the divergence time of Meripilaceae. In addition, Ceriporia and Meripilus have similar morphological characteristics and ecological habits, with broad-leaved trees being the most common hosts, but they belong to different families and have not been analyzed together. In this study, the morphology, phylogeny, and divergence times of Ceriporia and Meripilus are investigated. In addition, three new species of Ceriporia, one new species of Meruliopsis, and six new species of Meripilus are described and illustrated.
4 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae Materials and methods Morphological studies The studied specimens were deposited in the Fungarium of the Institute of Microbiology, Beijing Forestry University (BJFC). Morphological descriptions arebasedonfieldnotesandvoucherspecimens.Themicroscopicanalysis follows Wu et al. (2022) and Wang et al. (2023). Sections were studied at a magnificationofupto1000×usingaNikonEclipse80imicroscopewithphase contrast illumination. Microscopic features and measurements were made from slide preparations stained with Cotton Blue and Melzer’s reagent. Spores were measured from sections cut from the tubes. To represent the variation in spore size, 5% of measurements were excluded from each end of the range and are given in parentheses. In the description: KOH = 5% potassium hydroxide, IKI = Melzer’s reagent, IKI– = neither amyloid nor dextrinoid, CB = Cotton Blue, CB+ = cyanophilous in Cotton Blue, CB– = acyanophilous in Cotton Blue, L = arithmetic average of spore length, W = arithmetic average of spore width, Q = L/W ratios, and n = number of basidiospores measured from a given number of specimens. Color terms follow Anonymous (1969) and Petersen (1996). DNA extraction, amplification, and sequencing A CTAB rapid plant genome extraction kit DN14 (Aidlab Biotechnologies Co., Ltd, Beijing) was used to obtain DNA from dried specimens and to perform the polymerase chain reaction (PCR) according to the manufacturer’s instructions with some modifications (Sun et al. 2020; Wang et al. 2023). The internal transcribed spacer (ITS) and large subunit nuclear ribosomal RNA gene (nLSU) were amplified using the primer pairs ITS5/ITS4 and LR0R/LR7, respectively (White et al. 1990; Hopple and Vilgalys 1999) (https://sites. duke.edu/vilgalyslab/rdna_primers_for_fungi/). The small subunit nuclear ribosomal RNA gene (nSSU) region was amplified with primer pairs NS1 and NS4 (White et al. 1990). Part of TEF1 was amplified with primer pairs EF1-983F and EF1-1567R (Rehner and Buckley 2005). RPB1 was amplified with primer pairs RPB1-Af and RPB1-Cr (Matheny et al. 2002). RPB2 was amplified with primer pairs fRPB2-5F and fRPB2-7CR (Matheny 2005). The PCR procedure for ITS and TEF1 was as follows: initial denaturation at 95 °C for 3 min, followed by 34 cycles at 94 °C for 40 s, 54 °C for ITS and 54 °C for TEF1 for 45 s, and 72 °C for 1 min, with a final extension at 72 °C for 10 min. The PCR procedure for nLSU and nSSU was as follows: initial denaturation at 94 °C for 1 min, followed by 34 cycles of denaturation at 94 °C for 30 s, annealing at 50 °C for nLSU and 52 °C for nSSU for 1 min, and extension at 72 °C for 1.5 min, with a final extension at 72 °C for 10 min. The PCR procedure for RPB1 and RPB2 was as follows: initial denaturation at 94 °C for 2 min, followed by 10 cycles at 94 °C for 45 s, 60 °C for 45 s, and 72 °C for 1.5 min, then followed by 37 cycles at 94 °C for 45 s, 52 °C for 1 min, and 72 °C for 1.5 min, with a final extension at 72 °C. The PCR products were purified and sequenced at the Beijing Genomics Institute (BGI), China, with the same primers used in PCR. Newly generated sequences were deposited in GenBank. All sequences analyzed in this study are listed in Table 1.
5 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae Table 1. Taxa information and GenBank accession numbers of the sequences used in this study. Species name Sample no. Location GenBank accession No. References ITS nLSU nSSU TEF1 RPB1 RPB2 Agaricus campestris LAPAG370 —KM657927 KR006607 —KR006636 — — Zhou et al. (2016) Amylocorticium cebennense CFMR: HHB-2808 USA GU187505 GU187561 GU187612 GU187675 GU187439 —Binder et al. (2010) Antella americana HHB-4100-Sp USA EU232186 KP135196 ————Unpublished Aphanobasidium pseudotsugae CFMR: HHB-822 USA GU187509 GU187567 GU187620 GU187695 GU187455 —Binder et al. (2010) Athelia epiphylla CFMR: FP-100564 USA GU187501 GU187558 GU187613 GU187676 GU187440 —Binder et al. (2010) Bjerkandera adusta Dai 14516 China MW507097 MW520204 OQ509532 OQ559572 OQ542973 —Wang et al. (2021) B. fumosa Dai 21100 China MW507109 MW520211 OQ509533 OQ559573 OQ542974 —Wang et al. (2021) Boletopsis leucomelaena AFTOL-ID 1527 USA DQ484064 DQ154112 DQ435797 GU187763 GU187494 —Matheny et al. (2006a) Boletus edulis HMJAU4637 —JN563894 KF112455 —KF112202 KF112586 —Feng et al. (2012) Bondarzewia tibetica Yu 56 China KT693203 KT693205 —KX066148 KX066158 —Unpublished Byssomerulius corium FP-102382 USA KP135007 KP135230 — — KP134802 —Floudas and Hibbett (2015) Callistosporium graminicolor AFTOL-ID 978 USA DQ484065 AY745702 AY752974 GU187761 GU187493 —Matheny et al. (2006a) Ceriporia allantoidea Cui 8097 China KC182780 —————Jia et al. (2014) C. allantoidea Dai 8110 (holotype) China KC182767 KC182784 ————Jia et al. (2014) C. allantospora RLG-10478 (holotype) USA KP135039 —————Floudas and Hibbett (2015) C. arbuscula GC 1708–338 (holotype) China LC427008 LC427040 — — LC427058 —Chen et al. (2020) C. arbuscula GC 1708-340 China LC427009 LC427041 — — LC427068 —Chen et al. (2020) C. arbuscula Dai 26109 China PQ650033ªPQ642628ªPQ644088ªPQ662349ªPQ662329ª — Present study C. arbuscula Dai 26107 China PQ650034ªPQ642629ªPQ644089ªPQ662350ªPQ662330ª — Present study C. armeniaca Dai 24678A (holotype) China PQ650035ªPQ642630ªPQ644090ªPQ662351ªPQ662331ª — Present study C. aurantiocarnescens JV 0105/10 Czechia KX236482 KX236482 ————Spirin et al. (2016) C. aurantiocarnescens Dai 17951 China MW491774 MW491764 ————Unpublished C. bresadolae Rivoire 3701 France KX236467 KX236467 ————Spirin et al. (2016) C. bresadolae Dai 24539 China OQ476820 OQ476765 OQ509535 OQ559575 OQ542975 —Wang et al. (2023) C. bresadolae Dai 24541 China OQ476821 OQ476766 OQ509536 OQ559576 OR090875 —Wang et al. (2023) C. bresadolae VS 4018 Russia KX236466 —————Spirin et al. (2016) C. bubalinomarginata Dai 17937 China MW491775 MW491765 ————Chen et al. (2022) C. bubalinomarginata Dai 12113 China OQ476822 OQ476768 OQ509538 ———Wang et al. (2023) Ceriporia cf. mellita GC 1508-71 China LC427022 LC427044 — — LC427067 —Chen et al. (2020) Ceriporia cf. mellita WEI 17-024 China LC427024 LC427046 ————Chen et al. (2020) Ceriporia cf. mellita GC 1608-7 Japan LC427023 LC427045 — — LC427060 —Chen et al. (2020) Ceriporia cf. mellita Dai 8168 China KC182768 KC182785 —OQ559578 — — Jia et al. (2014) Ceriporia cf. mellita Dai 27083 China PQ650036ªPQ642631ªPQ644091ª — — — Present study Ceriporia cf. mellita Dai 27085 China PQ650037ªPQ642632ª————Present study C. crassa Dai 22034 (holotype) China OQ476823 OQ476769 OQ509539 OQ559579 OQ542977 —Wang et al. (2023) C. crassa JV 1008/41-J USA PP485892ª—————Present study C. crassiparietata Dai 26986 China PQ650038ªPQ642633ªPQ644092ªPQ662352ªPQ662332ª — Present study C. crassiparietata Dai 26988 China PQ650039ª — PQ662353ªPQ662333ª — Present study C. crassiparietata Dai 25079 (holotype) China PQ650040ªPQ642634ªPQ644093ªPQ662354ª — Present study C. crassiparietata Dai 7759 China KC182777—————Jia et al. (2014) C. daedaleoides Dai 16779 (holotype) Thailand KY825130 OR088493 OR095704 ———Wang et al. (2023) C. eucalypti Dai 18675 (holotype) Australia MW491779 MW491769 OR095705 OQ559580 — — Chen et al. (2022); Wang et al. (2023) C. excelsa Yuan 2747 China KC182778 —————Jia et al. (2014) C. excelsa Yuan 2744 China KC182773 —————Jia et al. (2014) C. gossypina Dai 23392 (holotype) China OQ476824 OQ476770 OQ509540 OQ559581 OR090876 —Wang et al. (2023) C. gossypina Dai 26113 China PQ650041ªPQ642667ªPQ644094ªPQ662355ªPQ662334ª — Present study C. griseoviolascensJV0110/26 Czechia KX236487 KX236487 ————Spirin et al. (2016)
6 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae Species name Sample no. Location GenBank accession No. References ITS nLSU nSSU TEF1 RPB1 RPB2 C. griseoviolascens Dai 13202 France OQ476825 OQ476771 OQ509541 OQ559582 — — Wang et al. (2023) C. griseoviolascens Dai 27053 China PQ650042ªPQ642635ªPQ644095ªPQ662356ªPQ662335ª — Present study C. griseoviolascens Dai 27054 China PQ650043ªPQ642636ªPQ644096ªPQ662357ª — — Present study C. hinnulea Cui 11291 (holotype) China OQ476826 OQ476772 OQ509542 OQ559583 OQ542978 —Wang et al. (2023) C. humilis Dai 7642 China KC182775 —OR095706 OR113375 — — Jia et al. (2014); Wang et al. (2023) C. humilis Spirin 4706 (holotype) Russia KX752608 —————Miettinen et al. (2016) C. langloisii FP-110343-sp USA KY948793 KY948886 — — KY948981 —Justo et al. (2017) C. macrospora Cui 6740 (holotype) China OQ476827 —OQ509543 ———Wang et al. (2023) C. macrospora Dai 24695 China OR086075 OR088494 OR095707 —OR090877 —Wang et al. (2023) “Ceriporia aff. macrospora" MEL 2382688 Australia KP013052 —————Unpublished C. manzanitaeRyvarden 21832 (holotype) USA KX236478 KX236478 ————Spirin et al. (2016) C. mellita BR 4865 France KX236485 KX236485 ————Spirin et al. (2016) C. mellita Dai 19118 China OQ476831 OQ476776 OQ509547 ———Wang et al. (2023) C. mellita Dai 18486A China OQ476832 OQ476777 OR095708 ———Wang et al. (2023) C. mpurii Dai 24426 China OQ476833 OQ476778 OR095709 —OR090878 —Wang et al. (2023) C. mpurii He 6687 China OQ476834 OQ476779 OR095710 OR113376 OR090879 —Wang et al. (2023) C. mpurii Miettinen 14381 (holotype) Indonesia KX752603 KX752603 ————Miettinen et al. (2016) C. occidentalis Spirin 8558 USA KX236475 KX236475 ————Spirin et al. (2016) C. occidentalis JV 1105/12-J USA KX236473 KX236473 ————Spirin et al. (2016) C. orientalis Li 1045 China JX623946 —OR095711 ———Jia et al. (2014); Wang et al. (2023) C. orientalis Dai 13400 (holotype) China OQ476835 OQ476780 OQ509548 OQ559585 OQ542979 —Wang et al. (2023) C. orientalis Dai 25794 China PQ650044ªPQ642637ªPQ644098ªPQ662359ª — — Present study C. pierii Dai 23500 China OQ476836 OQ476781 OQ509549 OQ559586 OR090880 —Wang et al. (2023) C. pierii Dai 23499 China OQ476837 OQ476782 OQ509550 OQ559587 OQ542980 —Wang et al. (2023) C. pierii Rivoire 1161 (holotype) France KX752604 KX752604 ————Miettinen et al. (2016) C. pseudospissa Dai 24566 (holotype) China OQ476838 OQ476783 OR095712 OR113377 — — Wang et al. (2023) C. pseudospissa Yuan 5965 China KC182772 KC182783 ————Jia et al. (2014) C. punctata Dai 15899 (holotype) China OQ476839 OQ476784 OQ509551 OQ559588 OQ542981 —Wang et al. (2023) C. punctata Dai 15904 China OQ476840 OR088495 OQ509552 OQ559589 OQ542982 —Wang et al. (2023) C. punctata Yuan 438 China PQ650045ª—————Present study C. punctata Yuan 439 China PQ650046ª—————Present study C. punicans Dai 13376 China OQ476841 OQ476785 OQ509553 OQ559590 OQ542983 —Wang et al. (2023) C. punicans JV 0808/30 (holotype) USA KX236479 KX236479 ————Spirin et al. (2016) C. purpurea Dai 22445 China OQ476842 OQ476786 OQ509554 OQ559591 — — Wang et al. (2023) C. purpurea Dai 16368 China KX494577 KX494581 OQ509555 OQ559592 OQ542984 —Wang et al. (2023) C. purpurea Rivoire 4413 (neotype) France KX236461 KX236461 ————Yuan et al. (2017) C. reticulata Li 1316 China JX623947 —OR095713 ———Jia et al. (2014); Wang et al. (2023) C. reticulata KHL11981 Norway JX109845 JX109845 ————Binder et al. (2013) “C. reticulata"RLG-11354-Sp USA KP135041 KP135204 — — KP134794 —Floudas and Hibbett (2015) “C. reticulata"Dai 27072 China PQ650047ªPQ642638ªPQ644099ªPQ662360ªPQ662337ª — Present study C. septocystidia RLG-9759-sp USA MZ636934 GQ470631 —MZ913692 MZ748442 —Chen et al. (2021) C. septocystidia RMJ-119-sp USA KY948783 ———KY948959 —Justo et al. (2017) C. sericea Spirin 4944 (holotype) Russia KX752609 KX752609 ————Miettinen et al. (2016) C. sericea Dai 27086 China PQ650048ªPQ642639ªPQ644100ªPQ662361ª — — Present study C. sericea Dai 26044 China PQ650049ªPQ642640ªPQ644101ªPQ662362ªPQ662338ª — Present study C. sinospissa Cui 11282 (holotype) China OQ476843 OQ476787 OQ509556 OQ559593 — — Wang et al. (2023) C. sinospissa Dai 10477 China KC182769 KC182781 ————Jia et al. (2014) C. sinospissa Dai 16831 China OQ476844 OQ476788 OQ509557 OQ559594 — — Wang et al. (2023)
7 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae Species name Sample no. Location GenBank accession No. References ITS nLSU nSSU TEF1 RPB1 RPB2 C. sinoviridans Dai 13621A (holotype) China MW491781 MW491771 OQ509558 ———Chen et al. (2022) C. sinoviridans Li 1046 China KC182776 —————Jia et al. (2014) C. sordescens Miettinen 15492.2 (holotype) USA KX752606 KX752606 ————Miettinen et al. (2016) Ceriporia sp. Dai 26805 China PP479793ªPP479793ªPQ644097ªPQ662358ªPQ662336ª — Present study C. spissa JV 0108/6 USA KX236483 KX236483 ————Spirin et al. (2016) C. spissa Dai 19164 Canada OQ476845 OQ476789 OQ509559 ———Wang et al. (2023) C. subbadia Dai 15062 China OQ476846 —OQ509560 ———Wang et al. (2023) C. subviridans Cui 8012 (holotype) China KC182774 —OR095714 OR113378 — — Jia et al. (2014); Wang et al. (2023) C. subviridans GC 1704-54 China LC427026 LC427048 ————Chen et al. (2020) C. torpida Murdoch 90 (holotype) Finland KX236477 KX236477 ————Spirin et al. (2016) C. triumphalis Kout 18 (holotype) Spain KX236476 KX236476 ————Spirin et al. (2016) C. viridans Dai 17003 China OQ476847 OQ476790 OQ509561 ———Wang et al. (2023) C. viridans Miettinen 11701 Netherlands KX752600 KX752600 ————Miettinen et al. (2016) C. viridans Yuan 5702 Netherlands KC182779 —————Jia et al. (2014) C. wuyiana Dai 24998 (holotype) China PQ650050ªPQ642641ªPQ644102ªPQ662363ªPQ662339ª—Present study Cerrena albocinnamomea Dai 12892 China KC485522 KC485539 ————Yuan (2014) C. albocinnamomea Dai 12955 China KC485521 KC485538 ————Yuan (2014) C. aurantiopora NIBRFG0000102423 Rep. Korea FJ821532 FJ821521 ————Lee and Lim (2010) C. aurantiopora SNU-m 03110102 Rep. Korea FJ821531 FJ821520 ————Lee and Lim (2010) C. consors F20080702KCM29 Rep. Korea FJ821527 FJ821516 ————Lee and Lim (2010) C. consors F20080208 LYW10 Rep. Korea FJ821528 FJ821517 ———FJ821543 Lee and Lim (2010) C. unicolor KHL-GB Sweden JQ031127 JQ031127 —JX109891 —JX109863 Sjoekvist et al. (2012) C. unicolor FD 299 USA KP135304 KP135209 ————Floudas and Hibbett (2015) C. zonata Dai 7821 China KC485529 KC485547 ————Yuan (2014) C. zonata Dai 7359 China KC485528 KC485546 ————Yuan (2014) Cotylidia sp.MB5 —AY854079 AY629317 AY705958 AY885148 AY864868 —Unpublished Crystallicutis serpens HHB-15692-Sp USA KP135031 KP135200 — — KP134785 —Floudas and Hibbett (2015) Crystallicutis sp. Miettinen-16854.3 USA KY948742 KY948890 — — KY948964 —Justo et al. (2017) Crystallicutis sp. Dai 6090 China JX623934 JX644066 ————Jia et al. (2014) Cytidiella albida GB-1833 Spain KY948748 KY948889 —MZ913675 KY948960 —Justo et al. (2017) C. nitidula T-407 Canada KY948747 — — MZ913676 KY948961 —Justo et al. (2017) Efibula americana FP-102165 USA KP135016 KP135256 —MZ913669 KP134808 —Floudas and Hibbett (2015) Flaviporus minutus Dai 16222 China KY131881 KY131938 ————Wu et al. (2017) F. minutus Dai 16240 China KY131883 KY131940 ————Wu et al. (2017) Gloeophyllum sepiarium Wilcox-3BB USA HM536091 HM536061 HM536062 HM536110 — — Garcia-Sandoval et al. (2011) Gloeoporus africanus O 918063 Uganda MG572763 MG572747 ————Jung et al. (2018) G. africanus O 918572 (holotype) Uganda MG572764 MG572748 ————Jung et al. (2018) G. citrinoalbus Dai 16238 (holotype) China KU360396 KU360404 OR095715 ———Yuan et al. (2016); Wang et al. (2023) G. citrinoalbus Dai 19547 Sri Lanka OQ476849 OQ476792 OQ509563 OQ559595 OQ542985 —Yuan et al. (2016); Wang et al. (2023) G. citrinoalbus Dai 15293 China OQ476850 OQ476793 OQ509564 OQ559596 — — Yuan et al. (2016); Wang et al. (2023) G. dichrous Dai 23626 China OQ476851 OQ476794 OQ509565 OQ559597 OQ542986 —Wang et al. (2023) G. dichrous Dai 23260 China OQ476852 OQ476795 OQ509566 OQ559598 OQ542987 —Wang et al. (2023) G. dichrous Dai 22633 China OQ476853 OQ476796 OQ509567 OQ559599 OQ542988 —Wang et al. (2023) G. hainanensis Dai 15259 China KU360403 KU360410 OQ509568 OQ559600 — — Yuan et al. (2016); Wang et al. (2023) G. hainanensis Yuan 4397 China KU360400 KU360409 ————Yuan et al. (2016) G. hainanensis Dai 15268 (holotype) China KU360401 KU360411 OQ509569 OQ559601 OQ542989 —Yuan et al. (2016); Wang et al. (2023)
8 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae Species name Sample no. Location GenBank accession No. References ITS nLSU nSSU TEF1 RPB1 RPB2 G. orientalis Cui 17922 China OQ476854 OQ476797 OQ509570 OQ559602 — — Wang et al. (2023) G. orientalis Dai 18536A China OQ476856 —————Wang et al. (2023) G. pannocinctus FP 135015 USA MG572755 MG572739 ————Jung et al. (2018) G. pannocinctus L-15726-Sp USA KP135060 KP135214 — — KP134867 —Jung et al. (2018) G. septatus Dai 22221 (holotype) China OQ476857 OQ476798 OQ509572 OQ559604 OQ542990 —Wang et al. (2023) G. thelephoroides BZ-2896 Belize MG572757 MG572741 ————Jung et al. (2018) G. thelephoroides JV 1808/26 French Guiana OQ476858 OQ476799 OQ509573 OQ559605 — — Wang et al. (2023) G. variiformis Dai 20655 China OQ476859 OQ476801 OQ509574 OQ559606 OQ542991 —Wang et al. (2023) G. variiformis Dai 22225 (holotype) China OQ476860 OQ476802 OQ509575 OQ559607 OQ542992 —Wang et al. (2023) Hapalopilus ochraceolateritius Miettinen-16992.1 USA KY948741 KY948891 — — KY948965 —Justo et al. (2017) Hydnochaete duportii AFTOL-ID 666 —DQ404386 AY635770 AY662669 DQ435793 — — Matheny et al. (2006a) Hyphoderma praetermissum AFTOL-ID 518 —AY854081 AY700185 AY707094 AY885150 AY864871 —Matheny et al. (2006a) Irpex flavus WHC 1381 China LC427029 LC427052 — — LC427064 —Chen et al. (2020) I. laceratus Dai 16433 China OQ476861 OQ476803 OQ509576 OQ559608 OQ542993 —Wang et al. (2023) I. laceratus PBU 0048 Thailand KC570339 KU760725 ————Permpornsakul et al. (2016) I. laceratus Dai 21940 China OQ476862 OQ476804 OQ509577 OQ559609 OQ542994 —Wang et al. (2023) I. laceratus SFFPS MZ-340 (holotype) —AB091675 —————Suhara et al. (2003) I. lacteus Dai 11230 China OQ476863 OQ476805 OQ509578 OQ559610 OQ542995 —Wang et al. (2023) I. lacteus FD-9 —KP135026 KP135224 — — KP134806 —Floudas and Hibbett (2015) I. latemarginatus FP-55521-T USA KP135024 KP135202 — — KP134805 —Floudas and Hibbett (2015) I. latemarginatus Marcin Piatek 4.IX.1997 Poland KX752592 KX752592 ————Miettinen et al. (2016) I. rosettiformis Meijer 3729 Brazil JN649346 JN649346 ————Sjökvist et al. (2012) Irpiciporus pachyodon SP-Lgt Italy AY849307 —————Pilotti et al. (2005) I. pachyodon PRM 846564 Czechia HQ728293 HQ729003 ————Tomšovský (2012) I. sinuosus PW 17/171 Thailand MK589288 —————Thamvithayakorn et al. (2019) I. sinuosus Dai 12234 China KX161649 KX161658 —OM982699 — — Unpublished I. xuchilensis Ryvarden 44669 Ecuador KX161650 KX161659 ————Unpublished Jaapia argillacea CBS: 252.74 Netherlands GU187524 GU187581 —GU187711 — — Binder et al. (2010) Junghuhnia fimbriatella Miettinen 2091 Russia JN710555 JN710555 ————Miettinen et al. (2012) Lactarius deceptivus AFTOL-ID 682 USA AY854089 AY631899 AY707093 AY885158 AY864883 —Matheny et al. (2006a) Leptoporus mollis RLG-7163 USA KY948794 MZ637155 —MZ913693 KY948956 —Justo et al. (2017) L. mollis Dai 21062 Belarus MW377302 MW377381 —MW337129 — — Liu et al. (2023) L. submollis Dai 20182 (paratype) China ON468434 ON468246 —ON468452 ON468448 —Liu et al. (2023) L. submollis Cui 18379 (paratype) China ON468433 ON468245 —ON468451 ON468447 —Liu et al. (2023) Leptosporomyces raunkiaeri CFMR: HHB-7628 USA GU187528 GU187588 GU187640 GU187719 GU187471 —Binder et al. (2010) Macrohyporia dictyopora PBU 0051 Thailand KC570331 KU760726 ————Permpornsakul et al. (2016) Meripilus albomarginatus Dai 19796 (holotype) China PQ650057ªPQ642661ªPQ644116ªPQ662374ª — — Present study M. albomarginatus Dai 25241 China PQ650058ªPQ642662ªPQ644117ªPQ662375ª — PQ662347ªPresent study M. albomarginatus Dai 24711 China PQ650059ª—————Present study M. albostygius Kout 1807/15.1 (holotype) Puerto Rico OM669892 OM669976 OM670027 — — OM810070 Wang et al. (2024) M. albostygius Kout 1508/18.1 Puerto Rico OM669893 —————Wang et al. (2024) M. albostygius RP 185 Brazil KP859303 —————Wang et al. (2024) M. brasiliensis RP 215 Brazil PP259422 PP259408 ————Westphalen et al. 2025 M. brasiliensis RP 200 Brazil PP259421 PP259407 ————Westphalen et al. 2025 M. caesiomarginatus Dai 19793 China OM669891 OM669975 OM670024 OM810094 —OM810067 Wang et al. (2024) M. castanopsidis Dai 20396 (holotype) China MT309485 MT309470 OM670025 OM810095 —OM810068 Wang et al. (2024) M. castanopsidis Dai 20397 China MT309486 MT309472 OM670026 OM810096 —OM810069 Wang et al. (2024)
9 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae Species name Sample no. Location GenBank accession No. References ITS nLSU nSSU TEF1 RPB1 RPB2 M. castanopsidis Dai 11693 China KY131865 KY131922 ————Wang et al. (2024) M. cinereus Cui 3266 China KY131844 KY131903 ————Wu et al. (2017) M. cinereus Dai 22427 China PQ650060ªPQ642648ª————Present study M. cinereus Dai 24688 China PQ650061ªPQ642649ªPQ644108ªPQ662368ª—PQ662343ªPresent study M. cinereus Dai 17581 China PQ650062ªPQ642650ª————Present study M. cinereus Dai 24690 China PQ650063ªPQ642651ªPQ644109ªPQ662369ª — — Present study M. concrescens MV 690 Brazil PP259426 PP259411 ————Westphalen et al. (2025) M. crataegi Dai 15499 China KY131846 KY131905 OM670028 OM810097 —OM810071 Wu et al. (2017); Wang et al. (2024) M. crataegi Dai 15497 (holotype) China KY131845 KY131904 OM670029 OM810098 —OM810072 Wu et al. (2017); Wang et al. (2024) M. crocatus Dai 12800 USA KY131869 KY131925 ————Wu et al. (2017) M. crocatus Dai 15917 China KY131870 KY131926 OM670063 OM810124 — — Wu et al. (2017); Wang et al. (2024) M. crocatus JV 0509/40 USA OM669894 OM669977 OM670030 OM810099 — — Wang et al. (2024) M. crocatus DLL 2009-061 USA JQ673152 —————Brazee et al. (2012) M. crocatus MJ 19/09 Slovakia JQ409466 OM669978 ————Vampola and Vlasák (2012) M. crocatus JV 0808/33 USA OM669895 OM669979 OM670031 OM810100 — — Wang et al. (2024) M. crystallinus Cui 10491 (holotype) China PQ650064ª — — ———Present study M. crystallinus Cui 10475 China PQ650065ªPQ642647ª — ———Present study M. dollingerii Dollinger 880 USA OM669897 —OM670032 OM810101 —OM810073 Wang et al. (2024) M. dollingerii Dollinger 1000 USA OM669899 OM669982 OM670034 — — OM810075 Wang et al. (2024) M. emarginatus Dai 24682A (holotype) China PQ650066ªPQ642652ªPQ644110ªPQ662370ª — PQ662344ªPresent study M. emarginatus Dai 24683A China PQ650067ªPQ642653ªPQ644111ªPQ662371ª — PQ662345ªPresent study M. emarginatus Dai 16971 China PQ650068ªPQ642654ª————Present study M. emarginatus Dai 24694A China PQ650069ªPQ642655ªPQ644112ªPQ662372ª — PQ662346ªPresent study M. emarginatus Dai 26696 China PQ650070ªPQ642656ª————Present study M. eminens Dai 11400 China KY131852 KY131909 OM670035 OM810103 — — Wu et al. (2017), Wang et al. (2024) M. eminens Dai 20832 China MT279689 MT279689 OM670036 OM810104 — — Wang et al. (2024) M. eminens Dai 22472 China OM669900 OM669983 OM670037 OM810105 — — Wang et al. (2024) M. eminens Dai 20868 China MT840117 MT840135 ————Wang et al. (2024) M. expallescens Dai 21060 Belarus MT840130 MT840148 OM670077 — — OM810090 Chen and Dai (2021), Wang et al. (2024) M. expallescens MJ 332/94 Czechia OM669935 —————Wang et al. (2024) M. expallescens MJ 642/93 Czechia OM669936 —————Wang et al. (2024) M. galapagensis MV 513 Brazil PP259430 PP259414 —PP239252 PP239254 —Westphalen et al. (2025) M. giganteus CBS 421.48 Germany MH856418 —————Vu et al. (2019) M. giganteus FP-100460-Sp Netherland KP135306 —————Floudas and Hibbett (2015) M. giganteus Cui 9202 UK OM669888 OM669973 OM670021 ———Wang et al. (2024) M. giganteus Cui 9203 UK OM669889 —OM670022 ———Wang et al. (2024) M. giganteus FP-135344-Sp UK KP135307 KP135228 ————Floudas and Hibbett (2015) M. furcatus TAA 150972 (holotype) Russia KY131853 KY131910 ————Wu et al. (2017) M. lavendulus Dai 9925 China KY131858 KY131915 ————Wu et al. (2017) M. lavendulus Dai 13587A (holotype) China KY131859 KY131916 OM670038 OM810106 — — Wu et al. (2017) M. lineatus Dai 17986 China MT840121 MT840139 OM670039 OM810107 —OM810076 Chen and Dai (2021), Wang et al. (2024) M. lineatus Dai 18281 Vietnam MT840123 MT840141 OM670040 OM810108 —OM810077 Chen and Dai (2021), Wang et al. (2024) M. lineatus JV 1008/18 USA OM669902 OM669985 OM670042 ———Wang et al. (2024) M. lineatus JV 1407/37 Costa Rica OM669903 OM669986 OM670043 OM810109 —OM810078 Wang et al. (2024) M. longicystidius PDD 70600 (holotype) New Zealand KY131863 —————Wu et al. (2017)
16 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae Figure 1. ML analysis of Irpicaceae based on the dataset of ITS+nLSU+TEF1+RPB1+nSSU. ML bootstrap values higher than 50% and Bayesian posterior probability values more than 0.90 are shown. New taxa are in bold. 0.06 Ceriporia sinospissa Cui 11282 Irpex laceratus PBU 0048 Ceriporia purpurea Dai 16368 Meruliopsis variegata Dai 19886 Meruliopsis rhizomorpha Dai 25816 Ceriporia torpida Murdoch 90 Ceriporia mellita Dai 19118 Gloeoporus citrinoalbus Dai 16238 Leptoporus submollis Dai 20182 Meruliopsis taxicola GC 1704-60 Irpex latemarginatus FP-55521-T Ceriporia mpurii Dai 24426 Meruliopsis marginata Dai 14737 Meruliopsis nanlingensis Dai 13414 Meruliopsis rosea Dai 18640A Ceriporia viridans Miettinen 11701 Meruliopsis pseudocystidiata Dai 18405 Leptoporus mollis RLG 7163 Gloeoporus dichrous Dai 22633 Meruliopsis taxicola Dai 21878 Ceriporia mpurii Miettinen 14381 Ceriporia purpurea Rivoire 4413 Meruliopsis crystallina Dai 26217 Ceriporia cf. mellita WEI 17-024 Ceriporia sinoviridans Dai 13621A Ceriporia sordescens Miettinen 15492-2 Ceriporia humilis Dai 7642 Meruliopsis faginea LEF-334408 Meruliopsis crassitunicata Dai 10833 Trametopsis cervina Cui 9985 Meruliopsis leptocystidiata Li 1011 Meruliopsis pseudocystidiata Li 1704 Ceriporia subbadia Dai 15062 Meruliopsis crystallina He 7477 Meruliopsis tarda LE 247365 Meruliopsis taxicola Dai 22625 Gloeoporus orientalis Dai 18536A Meruliopsis variegata Dai 19791 Ceriporia griseoviolascens Dai 27053 Meruliopsis taxicola Dai 22636 Ceriporia sinospissa Dai 10477 Phanerochaetella exilis HHB 6988 Ceriporia purpurea Dai 22445 Ceriporia reticulata Li 1316 Ceriporia manzanitae Ryvarden 21832 Gloeoporus variiformis Dai 20655 Ceriporia sp. Dai 26805 Trametopsis cervina AJ 185 Cytidiella albida GB 1833 Gloeoporus hainanensis Dai 15268 Ceriporia daedaleoides Dai 16779 Efibula americana FP 102165 Meruliopsis bambusicola Dai 21944 Raduliporus aneirinus HHB-15629-Sp Gloeoporus septatus Dai 22221 Ceriporia sinoviridans Li 1046 Ceriporia gossypina Dai 26113 Gloeoporus hainanensis Yuan 4397 Phanerochaete s.l. sp. RLG 13408 Gloeoporus africanus 918063 Ceriporia arbuscula GC1708-340 Byssomerulius corium FP 102382 Meruliopsis cystidiata ICN 139059 Ceriporia allantoidea Cui 8097 Ceriporia crassa JV 1008/41-J Ceriporia humilis Spirin 4706 Ceriporia sericea Dai 26044 Resiniporus resinascens BRNM 710169 Ceriporia mpurii He 6687 Ceriporia septocystidia RMJ-119-sp Gloeoporus dichrous Dai 23626 Meruliopsis rhizomorpha Dai 24733 Ceriporia orientalis Li 1045 Meruliopsis pseudocystidiata Dai 3204 Ceriporia crassiparietata Dai 26986 Ceriporia arbuscula Dai 26107 Ceriporia mellita Dai 18486A Meruliopsis tarda Dai 10226 Ceriporia arbuscula GC 1708-338 Phanerochaetella sp. HHB 11463 Ceriporia spissa Dai 19164 Ceriporia allantospora RLG 10478 Ceriporia griseoviolascens Dai 13202 Ceriporia sinospissa Dai 16831 Irpex flavus WHC 1381 Ceriporia crassa Dai 22034 Ceriporia punicans Dai 13376 Meruliopsis crystallina Dai 26052 Leptoporus mollis Dai 21062 Ceriporia macrospora MEL 2382688 Ceriporia bresadolae Dai 24541 Ceriporia cf. mellita Dai 27085 Meruliopsis albomellea Dai 15223 Ceriporia punctata Yuan 438 Meruliopsis albomellea Dai 15205 Ceriporia wuyiana Dai 24998 Meruliopsis taxicola Dai 17248 Irpex laceratus MZ 340 Ceriporia punctata Dai 15899 Resiniporus pseudogilvescens Wu 1209-46 Gloeoporus pannocinctus FP 135015 Meruliopsis parvispora CHWC 1505-129 Ceriporia allantoidea Dai 8110 Meruliopsis rhizomorpha Dai 25742 Ceriporia punicans Vlasak 0808-30 “Meruliopsis crassitunicata” Dai 9995 Ceriporia excelsa Yuan 2747 Irpex lacteus Dai 11230 Ceriporia viridans Dai 17003 Ceriporia punctata Yuan 439 Ceriporia bubalinomarginata Dai 17937 Ceriporia hinnulea Cui 11291 Ceriporia punctata Dai 15904 Ceriporia griseoviolascens JV 0110-26 Ceriporia gossypina Dai 23392 Meruliopsis nanlingensis Dai 8173 Ceriporia pierii Dai 23500 Bjerkandera fumosa Dai 21100 Gloeoporus dichrous Dai 23260 Ceriporia occidentalis JV 1105/12 Macrohyporia dictyopora PBU 0051 Ceriporia occidentalis VS 8558 Meruliopsis variegata Li 1780 Ceriporia aurantiocarnescens Dai 17951 Ceriporia crassiparietata Dai 26988 Gloeoporus thelephoroides BZ 2896 Gloeoporus citrinoalbus Dai 15293 Ceriporia bresadolae Rivoire 3701 Ceriporia cf. mellita Dai 8168 Irpex rosettiformis Meijer 3729 Ceriporia sericea Spirin 4944 Meruliopsis marginata Wei 3388 Crystallicutis sp. Miettinen 16854 Irpex laceratus Dai 16433 Ceriporia bresadolae VS 4018 Meruliopsis nanlingensis Dai 17172 “Ceriproia reticulata” Dai 27072 Gloeoporus citrinoalbus Dai 19547 Ceriporia reticulata KHL 11981 Ceriporia spissa JV 0108/6 Gloeoporus pannocinctus L-15726-Sp Meruliopsis leptocystidiata Wu 1708-43 Meruliopsis cystidiata 776308 Phanerochaetella xerophila HHB 8509 Ceriporia griseoviolascens Dai 27054 Ceriporia armeniaca Dai 24678A Meruliopsis marginata Cui 6878 Ceriporia triumphalis Kout 18 Ceriporia subviridans GC 1704-54 Irpex laceratus Dai 21940 Ceriporia eucalypti Dai 18675 Leptoporus submollis Cui 18379 Ceriporia septocystidia RLG-9759-sp Ceriporia pierii Rivoire 1161 Ceriporia macrospora Dai 24695 Gloeoporus variiformis Dai 22225 Meruliopsis marginata Cui 11626 Ceriporia pseudospissa Yuan 5965 Crystallicutis serpens HHB 15692 Ceriporia sericea Dai 27086 Cytidiella nitidula T 407 Ceriporia cf. mellita Dai 27083 Ceriporia viridans Yuan 5702 Ceriporia macrospora Cui 6740 Crystallicutis sp. Dai 6090 Ceriporia orientalis Dai 25794 Ceriporia cf. mellita GC 1508-71 Ceriporia bubalinomarginata Dai 12113 Ceriporia crassiparietata Dai 25079 Irpex lacteus FD 9 Ceriporia orientalis Dai 13400 Ceriporia subviridans Cui 8012 Ceriporia pseudospissa Dai 24566 Meruliopsis albostramineus HHB 10729 Gloeoporus thelephoroides JV 1808/26 Ceriporia langloisii FP-110343-sp Bjerkandera adusta Dai 14516 Ceriporia crassiparietata Dai 7759 “Ceriproia reticulata” RLG 11354 Hapalopilus ochraceolateritius Miettinen 16992 Ceriporia arbuscula Dai 26109 Meruliopsis parvispora Wu 1209-58 Gloeoporus africanus 918572 Ceriporia pierii Dai 23499 Ceriporia mellita BR 4865 Gloeoporus orientalis Cui 17922 Meruliopsis sp. FD 278 Irpex latemarginatus Marcin Ceriporia aurantiocarnescens JV 0105-10 Ceriporia bresadolae Dai 24539 Meruliopsis crassitunicata CHWC 1506-46 Ceriporia excelsa Yuan 2744 Ceriporia cf. mellita GC 1608-7 Gloeoporus hainanensis Dai 15259 96/1 63/0.9 99/0.98 93/1 97/1 77/0.97 82/1 99/1 94/1 99/1 99/1 94/1 100/1 98/1 54/- 91/1 91/1 99/1 54/0.98 58/- 100/1 81/0.96 100/- 100/1 58/1 89/1 100/1 66/0.91 99/1 77/0.99 82/- 84/0.96 97/1 76/0.97 99/1 52/0.93 73/0.96 51/- 65/0.9 87/0.99 100/1 79/- 58/0.97 72/0.99 77/0.98 88/- 89/1 98/0.91 90/1 92/1 99/1 97/0.99 100/1 88/1 55/1 96/1 63/- 96/1 99/1 95/1 60/- 71/0.93 99/1 96/0.99 99/1 92/0.99 100/1 97/1 84/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 -/0.91 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 Ceriporia Meruliopsis Gloeoporus The Ceriporia purpurea group The Ceriporia mpurii group The Ceriporia viridans group The Ceriporia mellita group
17 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae Figure 2. ML analysis of Meripilaceae based on the dataset of ITS+nLSU+TEF1+RPB2+nSSU. ML bootstrap values higher than 50% and Bayesian posterior probability values more than 0.90 are shown. New taxa are in bold. 0.06 Meripilus albostygius RP 185 Meripilus crystallinus Cui 10475 Meripilus cinereus Dai 17581 Meripilus sp. 1 JV 0309/45 Meripilus yunnanensis CLZhao 21647 Meripilus roseus Dai 19877 Pseudospongipellis delectans BRNM 686401 Spongipellis spumeus BRNM 712630 Cerrena unicolor FD 299 Meripilus stillicidiorum HCFC 1088 Radulodon yunnanensis He 6183 Meripilus subfurcatus Dai 11313 Flaviporus minutus Dai 16240 Pseudospongipellis litschaueri Dai 13845 Meripilus lavendulus Dai 9925 Pseudolagarobasidium baiyunshanense Han 405 Meripilus emarginatus Dai 26696 Meripilus vitreosanguineus Kout 0609/1 Meripilus vitreus Dai 12685 Meripilus albomarginatus Dai 25241 Meripilus lineatus JV1407/37 Meripilus sanguinolentus Dai 20976 Meripilus pouzarii MJ 27/04 Pseudolagarobasidium belizense VPB 197 Radulodon americanus CFMR-HHB 11240 Meripilus eminens Dai 22472 “Rigidoporus hypobrunneus” CM 108b Meripilus subfurcatus Dai 26167 Meripilus sp. 2 MJ 6003-Beneschová Meripilus cinereus Dai 22427 Meripilus crocatus Dai 12800 Meripilus vitreosanguineus MJ 144/95 Meripilus sp. 3 JV 8908/19 Meripilus eminens Dai 11400 Steccherinum tenue KHL 12316 Meripilus galapagensis MV 513 Cerrena consors F20080702KCM29 Meripilus sublineatus Dai 22598 Spongipellis spumeus He 6736 Meripilus sanguinolentus JV 1610/2-Tejklova Meripilus expallescens MJ 332/94 Meripilus crataegi Dai 15499 “Rigidoporus hypobrunneus” Dai 10569 “Rigidoporus hypobrunneus” Cui 16874 Meripilus malayanus Dai 18529 Meripilus neovitreus JV 0509/127 Meripilus subfurcatus Dai 25999 Cerrena consors F20080208LYW10 Meripilus pouzarii Dai 21043 Cerrena albocinnamomea Dai 12892 Meripilus robledoi HCFC 1095 Meripilus albomarginatus Dai 24711 Meripilus vinctus Cui 16903 Meripilus sp. 1 JV 0709/83 Meripilus niveomarginatus Dai 26373 Meripilus sp. 3 JV 1310/15-1 Meripilus crystallinus Cui 10491 Meripilus niveomarginatus Dai 18268 Meripilus vinctus JV 1407/36 Meripilus albostygius Kout 1807/15.1 Cerrena aurantiopora SNUm 03110102 Pseudolagarobasidium belizense CFMR-DCL04-31 Meripilus cinereus Cui 3266 Meripilus lineatus Dai 17986 Pseudospongipellis delectans OSM-F925 Meripilus revolubilis MCW 702 Pseudospongipellis litschaueri Dai 20266 Meripilus neovitreus JV 1009/59 Meripilus vitreus Miettinen 13591 Meripilus emarginatus Dai 24683A Meripilus niveomarginatus Dai 17695 Meripilus subfurcatus Dai 2105 Meripilus dollingerii Dollinger 880 Meripilus neovitreus JV 0709/188 Radulodon erikssonii CBS 126044 Cerrena aurantiopora NIBRFG 0000102423 Meripilus rhododendri Dai 22272 Radulodon americanus RLG 6350 Meripilus lineatus JV1008/18 Meripilus longicystidius Cui 16630 Radulodon casearius HHB-9567-sp Meripilus emarginatus Dai 24682A Meripilus crataegi Dai 15497 Meripilus lineatus Dai 18281 Meripilus brasiliensis RP 215 Meripilus concrescens MV 690 Meripilus sp. 3 MJ 4738-53/02 Meripilus lavendulus Dai 13587A Irpiciporus xuchilensis Ryvarden 44669 Irpiciporus sinuosus PW 17/171 Antella americana HHB-4100-Sp Radulodon erikssonii X 3536 Meripilus eminens Dai 20868 “Meripilus rigidus” F 2061 Meripilus tamilnaduensis A164FB3 Meripilus sanguinolentus MJ 111/04 Meripilus dollingerii Dollinger 1000 Meripilus eminens Dai 20832 Meripilus obscurus MCW 590 Meripilus castanopsidis Dai 20397 Meripilus niveomarginatus Dai 18540A Meripilus caesiomarginatus Dai 19793 Meripilus cinereus Dai 24688 Meripilus tamilnaduensis MKDM 01 Spongipellis spumeus BRNM 734877 Meripilus vitreus Cui 10340 Pseudolagarobasidium acaciicola CBS 115543 Meripilus furcatus TAA 150972 Meripilus albomarginatus Dai 19796 Meripilus tibeticus Cui 9588 Meripilus neovitreus JV 0509/47 Irpiciporus pachyodon SP-Lgt Meripilus castanopsidis Dai 20396 Meripilus srilankensis Dai 19535 Flaviporus minutus Dai 16222 Meripilus expallescens Dai 21060 Cerrena zonata Dai 7359 Meripilus expallescens MJ 642/93 Meripilus stillicidiorum Cui 16620 Cerrena albocinnamomea Dai 12955 Meripilus giganteus Cui 9202 Meripilus giganteus FP-100460-Sp Meripilus crocatus Dai 15917 Meripilus rhododendri Dai 22279 Meripilus minutissimus JV 1704/83 Meripilus vinctus Kout 1807/3 Cerrena uniclor KHLGB Meripilus rigidus JV 1704/79 Meripilus sublineatus Dai 17885 Meripilus sanguinolentus Dai 21030 Meripilus emarginatus Dai 24694A Cerrena zonata Dai 7821 Pseudospongipellis litschaueri BRNM 67093 Meripilus crocatus MJ 19/09 Meripilus tibeticus Cui 9381 Meripilus giganteus CBS421-48 Meripilus sumstinei Russell 5913 Spongipellis sibirica Dai 1723 Meripilus sulphureus Dai 17839 Meripilus giganteus FP-135344-sp Spongipellis quercicola Dai 20899 Junghuhnia fimbriatella Miettinen 2091 Pseudospongipellis unicolor CFMRcc-FP-71791-T Pseudolagarobasidium baiyunshanense Han 406 Irpiciporus pachyodon PRM 846564 Meripilus crocatus DLL 2009/061 Meripilus sp. 1 JV 0308/66 Meripilus crocatus JV 0509/40 Meripilus vitreus Cui 10341 Meripilus giganteus Cui 9203 “Rigidoporus hypobrunneus” Dai 10503 Meripilus albostygius Kout 1508/18.1 Irpiciporus sinuosus Dai 12234 Meripilus cinereus Dai 24690 Meripilus yunnanensis CLZhao 21583 Meripilus sp. 2 CWU 3874 Meripilus sublineatus Dai 20523 Meripilus sublineatus Dai 17553 “Rigidoporus hypobrunneus” JV 1712/13-J Pseudolagarobasidium acaciicola CBS 115544 Pseudospongipellis delectans MUcc 838 Meripilus noncontusus Dai 24718 Meripilus sp. TUFC 100564 Meripilus sublineatus Dai 19639 Meripilus longicystidius PDD 70600 Meripilus sulphureus Dai 17841 Meripilus obscurus MCW 722 Meripilus emarginatus Dai 16971 Spongipellis spumeus Dai 20901 “Rigidoporus hypobrunneus” Dai 19451 Meripilus subfurcatus Dai 2544 Radulodon casearius KRT-Iso-26 Meripilus vitreus JV 0110/48 Meripilus tamilnaduensis MKDM 01a Radulodon yunnanensis Cui 17979 Meripilus brasiliensis RP 200 Meripilus sanguinolentus MJ 39/00 Meripilus sanguinolentus JV 1310/11 Trametes ochracea HHB 13445 Meripilus vitreosanguineus JV 0909/3 Meripilus pouzarii JV 0511/23 Meripilus vitreus MJ 129/04 Spongipellis quercicola Cui 10009 Meripilus crocatus JV 0808/33 Pseudospongipellis unicolor CFMRcc-FP-59199-T 98/1 99/0.99 100/1 89/0.94 99/0.99 100/1 80/- 100 96/0.98 99/0.99 98/0.99 63/- 91/- 98/1 76/- 100/1 100/1 99/0.98 99/1 95/1 100/1 100/1 79/- 98/1 95/0.9 94/0.98 95/1 98/- 99/1 67/- 96/- 99/1 95/1 79/- 100/1 100/0.99 95/1 98/1 100/1 93/0.97 69/- 94/0.98 99/1 98/1 100/- 100/1 98/0.99 73/- 74/- 100/- 96/1 96/- 85/- 99/1 88/- 88/- 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 100/1 Meripilus Subclade Ⅰ Meripilus Subclade Ⅱ Meripilus Subclade Ⅲ Meripilus Subclade Ⅳ Meripilus Subclade Ⅴ Meripilaceae Spongipellis
18 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae Divergence time estimation The MCMC tree (Fig. 3) shows that the ancestor of the Polyporales evolved during the late Jurassic at 148.25 Myr [95% HPD of 116.72–186.39 Myr], which is largely consistent with the divergence time of the Polyporales by Ji et al. (2022). The two main clades, the Irpicaceae main clade and the Meripilaceae main clade, had strong support (0.98 PP; 0.94 PP, Fig. 5). The divergence time of the Irpicaceae main clade emerged with a mean stem age of 115.35 Myr [95% HPD of 88.98–147.46 Myr] and a mean crown age of 108.9 Myr [95% HPD of 84.92–139.28 Myr], which belongs to the early Cretaceousperiod.TheinitialdiversificationoftheMeripilaceae main clade occurred during the late Cretaceous with a mean stem age of 108.17 Myr [95% HPD of 82.75–140.53 Myr] and a mean crown age of 97.23 Myr [95% HPD of 74.77–126.72 Myr]. Their divergence times mostly overlapped. Three clades in the Irpicaceae are the Ceriporia clade, the Meruliopsis clade, and the Gloeoporus clade, and two clades in the Meripilaceae are the Meripilus clade and the Spongipellis clade. The divergence time of the Ceriporia clade emerged with a mean stem age of 91.17 Myr [95% HPD of 70.92–116.08 Myr] and a mean crown age of 83.61 Myr [95% HPD of 65.25–106.35 Myr], which belongs to the late Cretaceous period. The divergence time of the Meripilus clade emerged with a mean stem age of 97.23 Myr [95% HPD of 74.77–126.72 Myr] and a mean crown age of 81.38 Myr [95% HPD of 61.89–105.78 Myr], which belongs to the late Cretaceous period. The divergence times of the main nodes are shown in Fig. 3 and summarized in Table 2. The international chronostratigraphic chart follows Cohen et al. (2013; updated) (URL: http:// www.stratigraphy.org/ICSchart/ChronostratChart2022-10.pdf). Taxonomy Ceriporia Donk, Rev. Niederl. Homob. Aphyll. 2: 170 (1933). Type species. Ceriporia viridans (Berk. & Broome) Donk [as ‘Ceraporia’], Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 9: 171 (1933). Description. For a detailed description of this genus, see Wang et al. (2023). Ceriporia armeniaca Y.C. Dai, Chao G. Wang & Yuan Yuan, sp. nov. MycoBank No: 856725 Figs 4, 5 Etymology. Armeniaca (Lat.): refers to the species having an apricot pore surface when dry. Diagnosis. Differs from other Ceriporia species by resupinate basidiomata with a white pore surface when fresh, apricot when dry, round to angular pores of 5–7 per mm, subicular hyphae distinctly wider than tramal hyphae, lunate to allantoidbasidiosporesmeasuring4–4.5×2–2.2µm. Type. CHINA • Guangdong Province, Guangzhou, Baiyunshan Forest Park, on fallen angiosperm branch, 18 April 2023, Dai 24678A (BJFC042232, holotype).
19 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae Figure 3. Divergence time estimation of Irpicaceae and Meripilaceae from Bayesian evolutionary analysis sampling tree based on the conserved regions of three DNA fragments (ITS+nLSU+TEF1). Posterior probabilities are not less than 0.80, and the mean ages (Myr) of each node are annotated. The 95% highest posterior densities of divergence time estimation are marked by horizontal bars. 20.0 -200 -150 -100 -50 0 Hydnochaete duportii AFTOL-ID666 Junghuhnia fimbriatella Miettinen 2091 Ganoderma hochiminhense Cui 18229 Hyphoderma setigerum FD 312 Meripilus vitreus Dai 12685 Ceraceomyces sp. Dai 6090 Meripilus giganteus Cui 9202 Podoscypha multizonata Jahn 751012 Meripilus sp. 1 JV 0709/83 Meruliopsis faginea MW 673659 Meripilus srilankensis Dai 19535 Callistosporium graminicolor AFTOL-ID978 Meruliopsis nanlingensis Dai 13414 Meripilus albomarginatus Dai 25241 Ceriporia pierii Dai 23499 Lactarius deceptivus AFTOL-ID682 Spongipellis sibirica Dai 1723 Meripilus minutissimus JV 1704/83 Irpiciporus pachyodon PRM 846564 Ceriporia spissa Dai 19164 Gloeoporus pannocinctus FP 135015 Tomentella sp. AFTOL-ID1016 Russula emeticicolor FH 12253 Ceriporia reticulata Li 1316 Ceriporia gossypina Dai 23392 Hydnophanerochaete odontoidea CWN 00776 Efbula americana FP 102165 Hyphoderma mutatum HHB-15479-Sp Meripilus vinctus Cui 16903 Ceriporia orientalis Dai 13400 “Rigidoporus hypobrunneus” Kout 1807/8 Scopuloides hydnoides FP 150473 Pseudospongipellis litschaueri Dai 13845 Ceriporia aurantiocarnescens Dai 17951 Irpiciporus xuchilensis Ryvarden 44669 Ceriporia humilis Dai 7642 Ceriporia eucalypti Dai 18675 Pseudolagarobasidium baiyunshanense Han 405 Spongiporus leucospongia OKM 4335 Meripilus albostygius Kout 1807/15.1 Aphanobasidium pseudotsugae CFMR-HHB-822 Meripilus vitreosanguineus JV 0909/3 Flaviporus liebmannii X 249 Irpex rosettiformis Meijer 3729 Meripilus rhododendri Dai 22272 Meruliopsis rosea Dai 18640A Meripilus eminens Dai 22472 Meruliopsis tarda Dai 10226 Ceriporia langloisii FP-110343-sp Leptoporus mollis Dai 21062 Antrodiella trivialis MCW 369/12 Ceriporia allantoidea Dai 8110 Hypochnicium polonense NH 12117 Ceriporia wuyiana Dai 24998 Ganoderma guangxiense Cui 14453 Steccherinum tenue KHL 12316 Hydnophlebia chrysorhiza FD 282 Meripilus longicystidius Cui 16630 Ceriporia purpurea Dai 22445 Ceriporia torpida Murdoch 90 Trametes elegans FP-105679-sp Ceriporia arbuscula GC 1708-340 Phanerochaete alnea FP 151125 Meruliopsis marginata Cui11626 Meruliopsis variegata Dai 19886 Byssomerulius corium FP-102382 Ceriporia hinnulea Cui 11291 Gloeoporus africanus 918572 Ceriporia griseoviolascens Dai 13202 Radulodon americanus CFMR-HHB 11240 Meruliopsis albostramineus HHB 10729 Ceraceomyces sp. Miettinen16854 Ceriporia triumphalis Kout 18 Agaricus campestris LAPAG 370 Gloeoporus dichrous Dai 23260 Ceriporia allantospora RLG 10478 Antrodia favescens FP 103723 Podoserpula ailaoshanensis ZJL 2015015 Hypochnicium karstenii NH 10924 Ceriporia occidentalis JV 1105/12-J Meruliopsis albomellea Dai 15205 Meripilus castanopsidis Dai 20396 Ceriporia viridans Dai 17003 Ceriporia punctata Dai 15899 Jaapia argillacea CBS-252-74 Meruliopsis taxicola Dai 22625 Boletus edulis HMJAU 4637 Ceriporia pseudospissa Dai24566 Leptoporus submollis Dai 20182 Meripilus malayanus Dai 18529 Ceriporia subviridans Cui 8012 Meripilus pouzarii Dai 21043 Steccherinum ochraceum KHL 11902 Boletopsis leucomelaena AFTOL-ID1527 Meripilus noncontusus Dai 24718 Meripilus sp. 3 JV 1310/15-1 Meripilus lineatus Dai 17986 Cerrena uniclor KHL GB Phanerochaetella exilis HHB 6988 Neolentinus adhaerens DAOM 214911 Irpex latemarginatus FP-55521-T Hyphoderma litschaueri FP-101740-Sp Gloeoporus hainanensis Dai 15259 Hapalopilus ochraceolateritius Miettinen 16992 Meruliopsis crassitunicata Dai 10833 Meripilus niveomarginatus Dai 18540A Amylocystis lapponica FP 105131 Ceriporia punicans Dai 13376 Ceriporia sinoviridans Dai 13621A Meripilus sublineatus Dai 19639 Raduliporus aneirinus HHB-15629-Sp Ceriporia bresadolae Dai24539 Postia lactea Kotiranta 20058 Dacryobolus karstenii Miettinen 18685 Phanerochaete s.l. sp. RLG 13408 Bjerkandera adusta Dai 14516 Amylocorticium cebennense CFMR-HHB-2808 Meruliopsis rhizomorpha Dai 24733 Athelia epiphylla CFMR-FP-100564 Ceriporia mpurii Miettinen 14381 Ganoderma lingzhi Wu 1006/38 Trametes conchifer FP-106793-sp Gloeophyllum sepiarium Wilcox-3BB Ceriporia mellita BR 4865 Radulodon erikssonii CBS 126044 Irpex laceratus Dai 21940 Ceriporia sinospissa Cui 11282 Meruliopsis cystidiata ICN 139059 Meripilus sulphureus Dai 17841 Meruliopsis pseudocystidiata Dai 18405 Trametes ochracea HHB 13445 Irpiciporus sinuosus Dai 12234 Ceriporia septocystidia RLG-9759-sp Irpex lacteus Dai11230 Resiniporus pseudogilvescens Wu 1209-46 Podoscypha venustula LR 40821 Meripilus crystallinus Cui 10475 Cotylidia sp. MB 5 Radulodon yunnanensis Cui 17979 Grifola frondosa AFTOL 701 Meripilus crataegi Dai 15497 Ceriporia subbadia Dai 15062 Trametopsis cervina AJ 185 “Ceriproia reticulata” Dai 27072 Ceriporia sordescens Miettinen 15492.2 Phanerochaete velutina GC 1604/56 Meripilus caesiomarginatus Dai 19793 Flaviporus minutus Dai 16240 Gloeoporus orientalis Cui 17922 Phlebiopsis gigantea FCUG 1417 Meripilus tamilnaduensis MKDM 01 Ceriporia daedaleoides Dai 16779 Phlebiporia bubalina Dai 13168 Meruliopsis parvispora Wu 1209-58 Gloeoporus citrinoalbus Yuan 9654 Meripilus expallescens Dai 21060 Meripilus roseus Dai 19877 Pseudospongipellis litschaueri BRNM 712626 Bondarzewia tibetica Yu 56 Phanerochaetella xerophila HHB 8509 Meripilus crocatus JV 0808/33 Meripilus sumstinei Russell 5913 Meripilus rigidus JV 1704/79 Ceriporia cf. mellita Dai 8168 Meripilus dollingerii Dollinger 886 Spongipellis spumeus Dai 20901 Meruliopsis crystallina Dai 26052 Hypochnicium subrigescens KHL 11968 Meripilus yunnanensis CLZhao 21583 Meripilus neovitreus JV 1009/59 Ceriporia sericea Spirin 4944 Cytidiella albida GB 1833 Meripilus cinereus Cui 3266 Hyphoderma praetermissum AFTOL-ID-518 Meripilus subfurcatus Dai 2544 Panus fragilis HHB-11042-Sp Macrohyporia dictyopora PBU 0051 Meripilus crocatus Dai 15917 Ceriporia manzanitae Ryvarden 21832 Fomitopsis betulinus L 15603 Cymatoderma sp. OMC 1427 Ceriporia macrospora Cui 6740 Terana caerulea FP 104073 Meripilus sanguinolentus JV 1310/11 Meripilus emarginatus Dai 24694A Cytidiella nitidula T 407 Irpex flavus WHC 1381 Pseudospongipellis unicolor CFMRcc-FP-59199-T Meruliopsis crassitunicata Dai 9995 Gloeoporus thelephoroides JV 1808-26 Meripilus sp. TUFC 100564 Meripilus sp. 2 CWU 3874 Antrodia serpens Vampola 1989 Luteochaete subglobosa GC 1605/4 Meripilus tibeticus Cui 9588 Leptosporomyces raunkiaeri CFMR-HHB-7628 Ceriporia crassa Dai 22034 Ceriporia mpurii He 6687 Gloeoporus variiformis Dai 22225 Meruliopsis bambusicola Dai 21944 Spongipellis spumeus BRNM 734877 Ceriporia excelsa Yuan 2747 Ceriporia sp. Dai 26805 Meruliopsis leptocystidiata Wu 1708-43 Pseudospongipellis delectans BRNM 686401 Fomitopsis pinicola AFTOL 770 Grifola sordulenta AFTOL 562 Ceriporia crassiparietata Dai 25079 Meripilus lavendulus Dai 13587A Ceriporia cf. mellita GC 1508-71 Radulodon casearius HHB-9567-sp Gloeoporus septatus Dai 22221 Meripilus stillicidiorum Cui 16620 Ceriporia bubalinomarginata Dai 12113 Ceraceomyces serpens HHB 15692 Schizophyllum radiatum AFTOL-ID516 Hypochnicium cremicolor NH 11149 Ceriporia armeniaca Dai 24678A Gloeoporus citrinoalbus Dai 19547 Spongipellis quercicola Cui 10009 Antrodia heteromorpha Kosolapov 2003 Phanerochaetella sp. HHB 11463 Serpula himantioides MUCL 30528 Meripilus furcatus TAA 150972 24.69/- 67.65/0.99 44/1 48.34/1 37.02/1 17.27/0.94 34.39/1 29.49/- 10.92/- 12.38/0.85 19.83/0.97 18.31/1 49.77/- 25.03/- 16.23/- 66.8/- 45.27/- 115.35/1 7.87/- 5.66/1 55.47/0.98 28.16/1 38.43/- 12.46/- 48.26/1 21.06/0.92 33.79/1 11.92/1 19.88/1 131.7/1 47.99/- 28.94/1 5.92/1 54.13/1 12.91/1 6.47/- 43.58/0.99 62.96/1 57.82/1 10.88/1 66.89/- 25.08/- 3.92/0.89 95.02/1 108.17/0.96 22.62/1 19.07/1 11.71/1 46.59/1 60.13/0.95 22.19/- 11.57/0.84 2.63/1 136.1/- 92.45/1 30.77/0.88 56.98/0.95 6.99/1 83.41/1 148.25/- 32.44/1 7.81/1 66.87/- 102.96/0.82 61.14/1 15.33/1 2.59/1 24.48/1 111.01/1 47.21/- 4.66/1 37.77/- 15.69/1 81.38/1 118/1 100.88/1 75.8/- 9.94/1 46.81/1 14.49/- 25.47/1 9.39/- 71.38/- 61.65/0.98 59.99/1 67.33/- 44.92/- 5.72/1 28.05/1 3.51/- 41.14/0.93 13.23/1 24.82/1 77.67/- 44.07/1 40.53/1 21.73/0..83 165.34/- 64.94/- 17.43/1 41.65/1 52.39/1 98.39/- 32.71/1 10.44/0.99 142.02/- 21.94/- 10.63/1 9.12/- 83.97/1 1.35/- 19.71/1 67.85/0.99 28.89/1 11.48/1 17.32/- 8.81/0.97 11.11/1 31.32/1 83.61/1 6.37/1 47.3/0.98 7.2/1 22.9/1 44.1/1 116.53/1 87.86/0.97 28.93/1 91.17/- 58.54/1 21.37/1 57.76/- 40.88/1 9.9/1 29.07/1 16.44/1 13.67/1 68.45/1 18.76/- 10.54/1 63/- 100.58/0.99 41.22/1 47.09/1 9.99/1 53.85/- 151.5/0.94 27.77/1 74.06/1 14.25/1 124.09/1 67.09/1 61.04/1 18.46/1 72.27/0.93 172.29/1 46.19/1 10.64/1 45.6/- 97.23/0.94 91.55/1 22.86/1 14.01/1 14.05/- 143.21/1 6.44/1 38.22/0.95 11.7/1 11.47/1 34.17/1 47.38/1 74.45/0.98 12.48/1 86.68/1 56.38/0.89 7.46/0.96 24.21/- 5.09/1 89.55/1 40.64/- 78.76/1 148.97/0.89 155.64/- 55.37/0.99 83.69/1 14.63/1 104.9/1 21.01/1 118.27/1 83.39/1 26.78/1 9.94/1 142.680.99 5.06/1 27.04/0.98 5.72/1 79.13/1 7.91/1 4.63/1 32.86/0.9 35.45/0.83 62.82/1 3.07/1 14.22/0.99 17.84/1 42.97/- 61.22/- 4.96/0.93 18.28/1 36.23/0.98 2.43/0.97 47.56/1 124.5/0.98 24/0.99 74.7/- 14.85/1 20.52/1 10.22/0.98 64.22/0.92 39.03/1 108.9/0.98 21.25/1 119.11/- Irpicaceae Meripilaceae Phanerochaetaceae Meruliaceae Hyphodermataceae Steccherinaceae Podoscyphaceae Cerrenaceae Panaceae Postiaceae+Dacryobolaceae Postiaceae Ganodermataceae Trametaceae Grifolaceae Fomitopsidaceae Thelephorales Gloeophyllales Jaapiales Agaricales Boletales Amylocorticiales Atheliales Russulales Hymenochaetales Ceriporia Meruliopsis Gloeoporus Meripilus (Physisporinus) Polyporales Spongipellis A B C D E F G H I* J*
20 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae Description. Basidiomata annual, resupinate, soft, without odor or taste when fresh, consistently soft when fresh and dry, up to 5 cm long, 3 cm wide, and 0.2 mm thick at the center. Pore surface white when fresh, becoming apricot upon drying; sterile margin indistinct to almost lacking; pores round to angular, 5–7 per mm; dissepiments thin, lacerate. Subiculum very thin to almost absent. Tubes concolorous with pore surface, soft when dry, up to 0.2 mm long. Hyphal system monomitic; generative hyphae simple septate, hyaline,IKI−,CB+;tissuesbecomingorange-browninKOH.Subicularhyphae slightly thick-walled with a wide lumen, abundantly covered with small to large hyalinecrystalsandoilysubstances,sometimesencrustedwithfinecrystals and oily substances, frequently branched at more or less a right angle, straight, slightlyinterwoven,5–7µmindiam.Tramalhyphaethin-walledwithawide lumen, abundantly covered with rhombic or irregular hyaline crystals and olive oilysubstances,sometimesencrustedwithfinecrystalsandoilysubstances, frequentlybranched,straighttoslightlyflexuous,subparallelalongthetubes, Table 2. Estimated divergence times of the main nodes. Node Means of stem age (Mya)/95% HPD (Mya)/ posterior probabilities Means of crown age (Mya)/95% HPD (Mya)/ posterior probabilities Period A: Polyporales 148.97/119.64–183.87/0.89 148.25/116.72–186.39/– late Jurassic B: the Irpicaceae main clade 115.35/88.98–147.46/1 108.9/84.92–139.28/0.98 early Cretaceous C: the Meripilaceae main clade 108.17/82.75–140.53/0.96 97.23/74.77–126.72/0.94 late Cretaceous D: the Ceriporia clade 91.17/70.92–116.08/- 83.61/65.25–106.35/1 late Cretaceous E: the Gloeoporus clade 108.9/88.98–147.46/0.98 62.82/41.96–87.11/1 Paleogene F: the Meruliopsis clade 91.17/70.92–116.08/- 57.82/39.69–79.07/1 Paleogene G: the Meripilus clade 97.23/74.77–126.72/0.94 81.38/61.89–105.78/1 late Cretaceous H: the Spongipellis clade 97.23/74.77–126.72/0.94 13.67/7.07–22.73/1 Neogene I*: Hymenochaetales (Calibration point) 165.34/148.75–187.06/– 143.21/135.91–151.74/1 Early Cretaceous J*: Agaricales (Calibration point) 124.5/108.65–144.81/0.98 111.01/102.83–120.44/1 Early Cretaceous Hyphen “–” represents a posterior probability (PP) < 0.8. Figure 4. Basidiomata of Ceriporia armeniaca (holotype, Dai 24678A). Scale bar: 1 cm.
21 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae agglutinated,3–4µmindiam.Cystidiaandcystidiolesabsent.Basidiaclavate, withfoursterigmataandasimplebasalseptum,11–15.5×4–6µm;basidioles clavate to pyriform, smaller than basidia. Basidiospores lunate to allantoid, hyaline,thin-walled,smooth,sometimeswithoneortwosmallguttules,IKI−,CB−, 4–4.5(–5)×2–2.2µm,L=4.19µm,W=2.09µm,Q=2.01(n=30/1). Figure 5. Microscopic structures of Ceriporia armeniaca (drawn from the holotype, Dai 24678A) a basidiospores b basidia and basidioles c hyphae from subiculum dhyphaefromtrama.Scalebars:5µm(a);10µm(b–d).
22 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae Notes. Ceriporia armeniaca is closely related to C. arbuscula C.C. Chen & Sheng H. Wu and C. hinnulea Y.C. Dai, Chao G. Wang & Yuan Yuan, but C. arbuscula has a yellowish brown to pale brown pore surface when dry, short and tortuouslybranchedandnarrowersubicularhyphae(2–5μmdiam.vs.5–7µm indiam.,Chenetal.2020),andsmallerbasidiospores(3–3.5×1–1.5μmvs. 4–4.5×2–2.2µm,Chenetal.2020);Ceriporia hinnulea is distinguished from C. armeniaca by a fawn to cinnamon pore surface when dry, narrower subicular hyphaewithoutcrystalsoroilysubstances(3–5μmdiam.vs.5–7µmindiam., Wangetal.2023),andrelativelysmallerbasidiospores(3.5–4×2–2.1µmvs. 4–4.5×2–2.2µm,Wangetal.2023).Inaddition,thesethreespeciesform three independent lineages in the Ceriporia clade (Fig. 1). Ceriporia alba M. Pieri & B. Rivoire, C. camaresiana (Bourdot & Galzin) Bondartsev & Singer and C. rhodella (Fr.) Donk all have white pore surfaces when fresh.However,thefirsttwohavebiggerpores(3–4permminC. alba, 1–3 per mm in C. camaresiana vs. 5–7 per mm, Ryvarden and Gilbertson 1993; Pieri andRivoire1997)andbiggerbasidiospores(5.5–7×2–2.5μminC. alba, 5–6 ×2–3μminC. camaresianavs.4–4.5×2–2.2µm,RyvardenandGilbertson 1993; Pieri and Rivoire 1997); C. rhodellahasnarrowerbasidiospores(3.5–4× 1.5–2μmvs.4–4.5×2–2.2µm,LombardandGilbertson1965). Ceriporia crassiparietata Y.C. Dai, Chao G. Wang & Yuan Yuan, sp. nov. MycoBank No: 856727 Figs 6, 7 Etymology. Crassiparietata (Lat.): refers to the species having a pore surface with thick dissepiments. Diagnosis. Differs from other Ceriporia species by resupinate basidiomata with white to light yellow pore surface when fresh, cream to orange-yellow when dry, angular to irregular or sinuous pores of 4–5 per mm, distinctly thick dissepiments, subicular hyphae distinctly wider than tramal hyphae, lunate to allantoidbasidiosporesmeasuring4–4.4×2.1–2.3µm. Type. CHINA • Zhejiang Province, Wuyi County, Dahongyan Forest Park, on rotten angiosperm wood, 19 June 2023, Dai 25079 (BJFC042632, holotype). Description. Basidiomata annual, resupinate, soft, without odor or taste when fresh, consistently soft when fresh and dry, up to 3 cm long, 1 cm wide, and 0.2 mm thick at the center. Pore surface white to light yellow when fresh, becoming cream to orange-yellow upon drying; sterile margin indistinct to almost lacking; pores angular to irregular or sinuous, 4–5 per mm; dissepiments thick, entire. Subiculum very thin to almost absent. Tubes concolorous with pore surface, soft when dry, up to 0.2 mm long. Hyphal system monomitic;generativehyphaesimpleseptate,hyaline,IKI−,CB+;tissuesunchanged in KOH. Subicular hyphae thin-walled with a wide lumen, abundantly covered with large, rhombic, hyaline crystals and some oily substances, encrusted withfinecrystals,frequentlybranchedatmoreorlessarightangle,slightly flexuous,interwoven,5–6µmindiam.Tramalhyphaethin-walledwithawide lumen, abundantly covered with rhombic hyaline or pale orange crystals and oilysubstances,encrusted with finecrystals, frequently branched,straight toslightlyflexuous,subparallelalongthetubes,agglutinated,3–4.5µmin
23 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae diam. Cystidia and cystidioles absent. Basidia mostly barrel-shaped to short clavate,withfoursterigmataandasimplebasalseptum,9–11×5µm;basidioles of similar shape to basidia, but smaller. Basidiospores lunate to allantoid,hyaline,thin-walled,smooth,IKI−,CB−,(3.8–)4–4.4(–4.6)×(2–)2.1– 2.3(–2.5)µm,L=4.08µm,W=2.19µm,Q=1.85–1.89(n=60/2). Additional specimens examined. CHINA • Liaoning Province, Fengcheng County, Tongyuanbao, on Quercus, 27 August 2006, Dai 7759 (BJFC010215); Xizang Autonomous Region, Linzhi, Chayu County, fallen branch of Rosaceae, 27 October 2023, Dai 26986 (BJFC044538), Dai 26988 (BJFC044540). Notes. Ceriporia crassiparietata is closely related to the Ceriporia viridans group by sharing resupinate basidiomata with a cream, cinnamon buff, pinkish, lilac to apricot orange pore surface; subicular hyphae wider than tramal hyphae; hyphae frequently branched at a right angl; and oblong-ellipsoid, short cylindrical, lunate toallantoidbasidiosporesmostlywiderthan1.5µm(Wangetal.2023).Ceriporia eucalypti Y.C. Dai & Jia J. Chen, C. gossypina Y.C. Dai, Chao G. Wang & Yuan Yuan and C. subviridans Y.C. Dai, Chao G. Wang & Yuan Yuan are similar to C. crassiparietata by having almost the same size of pores (3–5 per mm or 4–5 per mm). However, C. eucalyptihasnarrowerbasidiospores(4–4.4×1.1–1.4µmvs.4–4.4 ×2.1–2.3µm,Chenetal.2022);C. gossypina has a white, buff to deep olive pore surfacewhenfreshandrelativelysmallerbasidiospores(3.5–4×1.8–2µmvs. 4–4.4×2.1–2.3µm,Wangetal.2023);andC. subviridans has a peach to apricot orangeporesurfacewhendry,widersubicularhyphae(4.5–9µmindiam.vs. 5–6µmindiam.,Wangetal.2023),andrelativelysmallerbasidiospores(3.3–3.7 ×1.8–2µmvs.4–4.4×2.1–2.3µm,Wangetal.2023). Ceriporia cystidiata Ryvarden & Iturr., C. microspora I. Lindblad & Ryvarden and C. otakou (G. Cunn.) P.K. Buchanan & Ryvarden share white, cream to isabelline basidiomata with C. crassiparietata. However, C. cystidiata is known from C. crassiparietata by tubular encrusted cystidia and narrower allantoid basidiospores Figure 6. Basidiomata of Ceriporia crassiparietata (holotype, Dai 25079). Scale bar: 1 cm.
24 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae (4–4.5×1µmvs.4–4.4×2.1–2.3µm,RyvardenandIturriaga2003).Ceriporia microspora is easily distinguished from C. crassiparietata by smaller ellipsoid basidiospores(3–3.5×1.5–2µmvs.4–4.4×2.1–2.3µm,LindbladandRyvarden 1999). Ceriporia otakou differs from C. crassiparietata by larger pores (1–3 per mm vs. 4–5 per mm, Cunningham 1947) and bigger ovoid to ellipsoid basidiospores(4.5–6×2–2.5µmvs.4–4.4×2.1–2.3µm,Cunningham1947). Figure 7. Microscopic structures of Ceriporia crassiparietata (drawn from the holotype, Dai 25079). a basidiospores b basidia and basidioles c hyphae from subiculum dhyphaefromtrama.Scalebars:5µm(a);10µm(b–d).
25 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae Ceriporia wuyiana Y.C. Dai, Chao G. Wang & Yuan Yuan, sp. nov. MycoBank No: 856728 Figs 8, 9 Etymology. Wuyiana (Lat.): refers to the species being found in Wuyi County, Zhengjiang Prov., East China. Diagnosis. Differs from other Ceriporia species by resupinate basidiomata with a white to cream pore surface when fresh, clay pink to pale lavender when dry, round to angular pores of 5–6 per mm, subicular hyphae relatively wider than tramalhyphae,allantoidtolunatebasidiosporesmeasuring4.3–5×1.7–2µm. Type. CHINA • Zhejiang Province, Wuyi County, Guodong Forest Park, on rotten angiosperm wood, 19 June 2023, Dai 24998 (BJFC042551, holotype). Description. Basidiomata annual, resupinate, soft, without odor or taste when fresh, soft when dry, up to 6 cm long, 2.5 cm wide, and 0.2 mm thick at the center. Pore surface white to cream when fresh, becoming clay pink to pale lavender upon drying; sterile margin indistinct to almost lacking; pores round to angular, 5–6 per mm; dissepiments thin, lacerate. Subiculum very thin to almost absent. Tubes concolorous with pore surface, soft when dry, up to 0.2 mm long. Hyphal systemmonomitic;generativehyphaesimpleseptate,hyaline,IKI−,CB+;tissues becoming orange-brown in KOH. Subicular hyphae thinto slightly thick-walled with a wide lumen, abundantly covered with small, rhombic, hyaline crystals and oilysubstances,sometimesencrustedwithfinecrystals,frequentlybranchedat moreorlessarightangle,straight,slightlyinterwoven,4–5µmindiam.Tramal hyphae thin-walled with a wide lumen, abundantly covered with rhombic or irregularpaleorangecrystalsandoilysubstances,sometimesencrustedwithfine crystals,frequentlybranched,straighttoslightlyflexuous,subparallelalongthe tubes,agglutinated,3–4µmindiam.Cystidiaandcystidiolesabsent.Basidiabarrel-shaped to somewhat pyriform, with four sterigmata and a simple basal septum,9–12.5×4–5µm;basidiolesofsimilarshapetobasidia,butsmaller.Basidiosporesallantoidtolunate,hyaline,thin-walled,smooth,IKI−,CB−,(4.1–)4.3–5× (1.5–)1.7–2µm,L=4.58µm,W=1.83µm,Q=2.51(n=30/1). Figure 8. Basidiomata of Ceriporia wuyiana (holotype, Dai 24998). Scale bar: 1 cm.
32 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae when dry, thick-walled and apically encrusted hyphoid cystidia and almost the samesizeasbasidiospores(5–6µminM. lineatus;4.8–5.6×4.5–5.2µminM. sublineatus;5.2–6.2×4.6–5.7µminM. albomarginatus, Ryvarden and Gilbertson 1994; Wang et al. 2024). However, M. lineatus has a pinkish-buff to reddish-brown pileal surface and a bright orange-red pore surface when fresh (Ryvarden and Johansen 1980), and M. sublineatus has a normally azonate pileal surface and ventricosethin-walledhymenialcystidiabearingfinecrystals(Wangetal.2024). Meripilus albomarginatus forms an independent lineage in the Meripilus clade (Fig. 2) and groups with M. lineatus and M. sublineatus in a joint subclade (Fig. 2). However, there are 20 base pair differences in ITS sequences between Meripilus albomarginatus and M. lineatus, which account for a 3% nucleotide difference in the ITS regions. Four recorded synonyms of Meripilus lineatus (Physisporinus lineatus), viz. Polyporus zonalis Berk. (Sri Lanka), P. pusiolus Ces. (Malaysia), P. punctatus Jungh. (Indonesia), and P. epilinteus Berk. & Broome (Sri Lanka), were originally described from Asia. Polyporus zonalis has a pinkish-buff to reddish-brown upper surface, concolorous margin, and white to dingy livid gray pore surface (Pegler and Waterston 1968); P. punctatus has a cloudy-dirty and pale red pore surface and very small pores (Junghuhn 1838); and P. pusiolus and P. epilinteus have resupinate basidiomata (Berkeley and Broome 1873; Cesati 1879). The above four species are different from Meripilus lineatus, M. sublineatus and M. albomarginatus in morphology. Meripilus crystallinus Y.C. Dai, Chao G. Wang & Yuan Yuan, sp. nov. MycoBank No: 856731 Figs 14, 15 Etymology. Crystallinus (Lat.): refers to the species having hyphoid cystidia with crystals. Diagnosis. Differs from other Meripilus species by resupinate basidiomata with buff yellow, salmon to clay pink pore surface when dry, angular pores of 6–8 per mm, thin-walled and apically encrusted hyphae at the dissepiment edge, broadly ellipsoidtoovoidandslightlythick-walledbasidiosporesmeasuring4.3–5×4–4.5µm. Type. CHINA • Yunnan Province, Weixi County, Laojunshan Nature Reserve, on fallen trunk of Picea, 22 September 2011, Cui 10491 (BJFC011386, holotype). Description. Basidiomata annual, resupinate, soft to ceraceous, and without odor or taste when fresh, becoming brittle upon drying, up to 10 cm long, 6 cm wide, and 3.2 mm thick at the center. Pore surface buff yellow, salmon to clay pink when dry; sterile margin almost absent; pores angular, 6–8 per mm; dissepiments thin, entire to slightly lacerate. Subiculum very thin to almost absent, up to 0.2 mm thick. Tubes concolorous with pore surface, brittle when dry, up to 3 mm long. Hyphal system monomitic; generative hyphae simpleseptate,hyalinetoyellowishbrown,smooth,IKI−,CB+;tissuesunchanged in KOH. Subicular hyphae slightly thick-walled with a wide lumen, unbranched, moderatelysimpleseptate,slightlyflexuous,looselyinterwoven,agglutinated, 4–7µmindiam.Tramalhyphaeslightlythick-tothick-walledwithawidelumen, occasionallybranched,frequentlysimpleseptate,slightlyflexuous,subparallelalongthetubes,agglutinated,3–5µmindiam.;somethin-walledhyphae
33 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae at the dissepiment edge bearing crystals at the tips and resembling hyphoid cystidia. Hymenial cystidia absent; cystidioles fusoid, thin-walled, smooth, 12– 18×4–5µm;basidiabarrel-shaped,withfoursterigmataandasimplebasal septum,11–12×5.5–6µm;basidiolesofsimilarshapetobasidia,butsmaller. Basidiospores broadly ellipsoid to ovoid, hyaline, slightly thick-walled, smooth, sometimeswithonemediumguttule,IKI−,weaklyCB+,(4.2–)4.3–5×4–4.5 µm,L=4.75µm,W=4.15µm,Q=1.13–1.15(n=60/2). Additional specimen examined. CHINA • Yunnan Province, Weixi County, Laojunshan Nature Reserve, on fallen trunk of Picea, 22 September 2011, Cui 10475 (BJFC011370). Notes. Meripilus crystallinus is similar and related to M. eminens (Y.C. Dai) Rajchenb. & Westph. by having annual and resupinate basidiomata, the absence of a sterile margin, angular and almost the same size pores (7–8 per mm in M. eminens; 6–8 per mm in M. crystallinus, Dai 1998), and almost the same sizeasbasidiospores(4.2–6×3.9–5.2µminM. eminens;4.3–5×4–4.5µmin M. crystallinus, Dai 1998). However, M. eminens has thinto fairly thick-walled tramal hyphae, and thick-walled and apically encrusted hyphoid cystidia penetrating above the hymenial surface (Dai 1998). Meripilus vitreus (Pers.) Rajchenb. & Westph. is phylogenetically related to M. crystallinus (Fig. 4), but M. vitreus has thick-walled hyphoid cystidia with coarse crystals in most specimens, relatively bigger pores (5–6 per mm vs. 6–8 per mm), and thin-walled basidiospores(5–5.5×4–4.5µmvs.4.3–5×4–4.5µm,Wangetal.2024).In addition, there are 20 base pair differences in ITS sequences between these two species, which accounts for a 3% nucleotide difference in the ITS regions. Meripilus crystallinus and M. stillicidiorum (Speg.) Rajchenb. & Westph. share the pinkish buff, buff yellow, salmon to clay pink pore surface, the absence of sterile margin, and almost the same pore size (5–7 per mm in M. stillicidiorum; 6–8 per mm in M. crystallinus, Wang et al. 2024). However, M. stillicidiorum lacksanykindofcystidia,relativelylargerbasidiospores(5–5.7×4–4.8 µmvs. 4.3–5×4–4.5µm,Wangetal.2024),andsofar,isonlyknownfromthetype locality in Australia. Meripilus revolubilis Westph. & R.M. Silveira and M. robledoi Figure 14. Basidiomata of Meripilus crystallinus (holotype, Cui 10491). Scale bar: 1 cm.
34 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae Rajchenb. & Westph. were described with the resupinate basidiomata, white, cream to isabelline pore surface, thick-walled encrusted cystidia, which are similar to M. crystallinus (Westphalen et al. 2025). Phylogenetically, Meripilus crystallinus, M. eminens, M. revolubilis, M. robledoi, and M. vitreus form a distinct joint clade and share similar morphological characters. Figure 15. Microscopic structures of Meripilus crystallinus (drawn from the holotype, Cui 10491). a basidiospores b basidia and basidioles c cystidioles d hyphoid cystidia-like hyphae at the dissepiment edge e hyphae from subiculum f hyphae fromtrama.Scalebars:5µm(a);10µm(b–f).
35 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae Meripilus emarginatus Y.C. Dai, Chao G. Wang & Yuan Yuan, sp. nov. MycoBank No: 856732 Figs 16, 17 Etymology. Emarginatus (Lat.): refers to the basidiomata of species lacking a sterile margin. Diagnosis. Differs from other Meripilus species by resupinate basidiomata with a white pore surface when fresh, round to angular pores of 6–7 per mm, thick-walled and apically encrusted hyphoid cystidia, thin-walled and smooth hymenial cystidia, subglobose to globose basidiospores measuring 4.8–5.2×4.5–5.2µm. Type. CHINA • Guangdong Province, Guangzhou, Baiyunshan Forest Park, on rotten angiosperm wood, 18 April 2023, Dai 24682A (BJFC042236, holotype). Description. Basidiomata annual, resupinate, soft to ceraceous, and without odor or taste when fresh, becoming fragile upon drying, up to 8 cm long, 5 cm wide, and 0.6 mm thick at the center. Pore surface white when fresh, unchanged after bruising, pinkish buff to clay buff when dry; sterile margin absent; pores round to angular, 6–7 per mm; dissepiments thin, slightly lacerate. Subiculum very thin to almost absent. Tubes concolorous with pore surface, fragile when dry, up to 0.6 mm long. Hyphal system monomitic; generativehyphaesimpleseptate,hyalinetopaleyellowish,smooth,IKI−, moderately CB+; tissues unchanged in KOH. Tramal hyphae slightly thickwalled with a wide lumen, occasionally branched, frequently simple septate,slightlyflexuous,subparallelalongthetubes,agglutinated,4–5µmin diam. Hyphoid cystidia present, arising from tramal hyphae and completely embedded in trama, not projecting from the hymenium, sometimes projecting from the dissepiment edge, thick-walled with swollen tips, apically encrusted,6–12µmindiam.attheapex;hymenialcystidiapresent,fusoid, thin-walled, smooth, 20–25 × 4.5–5.5 µm; cystidioles fusoid, thin-walled, smooth, 13–14 × 5–6 µm; basidia barrel-shaped to capitate, with four sterigmataandasimplebasalseptum,15–17×7–8µm;basidiolesmostly pyriform, smaller. Crystals present among the hymenium and tube trama. Basidiospores subglobose to globose, hyaline, thin-walled, smooth, sometimeswithonesmallorlargeguttule,IKI−,weaklyCB+, (4.5–)4.8–5.2(–5.5) ×4.5–5.2(–5.5)µm,L=5.11µm,W=4.90µm,Q=1.03–1.05(n=90/3). Additional specimens examined. CHINA • Fujian Province, Fuding County, Tailao Mts., on fallen trunk of Cunninghamia, 22 August 2016, Dai 16971 (BJFC023076); • Guangdong Province, Guangzhou, Baiyunshan Forest Park, on rotten angiosperm wood, 18 April 2023, Dai 24683A (BJFC042237), Maofengshan Forest Park, on rotten angiosperm wood, 19 April 2023, Dai 24694A (BJFC042248); • Xizang Autonomous Region, Linzhi, Motuo County, on dead Miscanthus, 24 October 2023, Dai 26696 (BJFC044246). Notes. Meripilus emarginatus is similar to M. albostygius (Berk. & M.A. Curtis) Westph. & Rajchenb., M. eminens, M. rigidus (Y.C. Dai, Chao G. Wang & Vlasák) Westph. & Rajchenb., M. srilankensis, and M. sulphureus (Y.C. Dai, Yuan Yuan & Chao G. Wang) Westph. & Rajchenb. in micromorphology by the thick-walled hyphoid cystidia and subglobose basidiospores. However, M. albostygius has a red to violet pore surface when fresh, smaller pores
36 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae (8–10 per mm vs. 6–7 per mm, Wang et al. 2024), and smaller basidiospores (4–4.7×3.2–4µmvs.4.8–5.2×4.5–5.2µm,Wangetal.2024),andtodate is only known from Central and South America; M. eminens also has ceraceous basidiomata with a white pore surface when fresh, almost the same size angular pores (7–8 per mm vs. 6–7 per mm, Dai 1998) and basidiospores (4.2–6×3.9–5.2µmvs.4.8–5.2×4.5–5.2µm,Dai1998),but,itdiffersfrom M. emarginatus by the absence of thin-walled and smooth hymenial cystidia; M. rigidus differs from M. emarginatus by a brown-red pore surface when fresh, smaller pores (10–12 per mm vs. 6–7 per mm, Wang et al. 2024), smallerbasidiospores(4–4.6×3.2–4µmvs.4.8–5.2×4.5–5.2µm,Wangetal. 2024), and, to date, is only known from Central America; M. srilankensis is distinguished from M. emarginatus by distinctly thick-walled and wider tramal hyphae(3.8–8µmindiam.vs.4–5µmindiam.,Wangetal.2024)andthe absence of hymenial cystidia; M. sulphureus differs from M. emarginatus by a sulphur yellow pore surface when fresh and relatively smaller basidiospores (4–5×3.5–4µmvs.4.8–5.2×4.5–5.2µm,DaiandDai2018).Inmacro-morphology, Meripilus emarginatus has an unchanged pore surface. Phylogenetically, Meripilus emarginatus forms an independent lineage nested in the Meripilus clade (100% ML, 1.00 BPP, Figs 3, 4). However, it grouped with M. dollingerii (Y.C. Dai, Chao G. Wang & Vlasák) Westph. & Rajchenb. and M. malayanus in a joint subclade (90% ML, 1.00 BPP, Fig. 4). Although Meripilus dollingerii and M. emarginus share thin-walled smooth hymenialcystidiaandalmostthesamesizeasbasidiospores(4.5–5.5× 4–5µminM. dollingerii,4.8–5.2×4.5–5.2µminM. emarginatus, Wang et al. 2024), M. dollingerii has a pinkish to red pore surface when fresh and thin-walled apically encrusted hyphoid cystidia. Meripilus malayanus also has thick-walled and apically encrusted hyphoid cystidia, but it differs from M. emarginatus by hard corky basidiomata when dry, a grayish brown pore surface, and the presence of a sterile margin. Figure 16. Basidiomata of Meripilus emarginatus (holotype, Dai 24682A). Scale bar: 1 cm.
37 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae Meripilus malayanus Y.C. Dai, Chao G. Wang & Yuan Yuan, sp. nov. MycoBank No: 856733 Figs 18, 19 Etymology. Malayanus (Lat.): refers to the species being found in Malaysia. Diagnosis. Differs from other Meripilus species by resupinate nodulose basidiomata with a honey buff to grayish brown pore surface when dry, angular and Figure 17. Microscopic structures of Meripilus emarginatus (drawn from the holotype, Dai 24682A). a basidiospores b basidia and basidioles c cystidioles d hymenial cystidia e hyphoid cystidia f hyphaefromtrama.Scalebars:5µm(a); 10µm(b–f).
38 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae sometimes elongated pores of 7–8 per mm, thick-walled and apically encrusted hyphoidcystidia,subglobosebasidiosporesmeasuring5–5.5×4.5–5µm. Type. MALAYSIA • Selangor, Taman Botani Negara Shah Alam, on fallen angiosperm trunk, 12 April 2018, Dai 18529 (BJFC026818, holotype). Description. Basidiomata annual, resupinate, nodulose, soft corky, and without odor or taste when fresh, becoming hard corky upon drying, up to 15 cm long, 6 cm wide, and 0.8 mm thick at the center. Pore surface white to cream when fresh, unchanged after bruising, honey buff to grayish brown when dry; sterile margin thinning out, grayish brown when dry; pores angular, sometimes elongated, 7–8 per mm; dissepiments thin, slightly lacerate. Subiculum pinkish buff, corky, up to 0.3 mm thick. Tubes concolorous with pore surface, hard corky when dry, up to 0.5 mm long. Hyphal system monomitic;generativehyphaesimpleseptate,smooth,IKI−,CB+;tissuesbecoming reddish brown in KOH. Subicular hyphae distinctly thick-walled with a wide lumen,rarelybranchedandsimpleseptate,moreorlessflexuous,interwoven,5–8µmindiam.Tramalhyphaeslightlythick-walledwithawidelumen, occasionally branched, moderately simple septate, straight, subparallel along thetubes,agglutinated,3.5–4.8µmindiam.Hyphoidcystidiapresent,arising from tramal hyphae and completely embedded in trama, not projecting from the hymenium or dissepiment edge, thick-walled with swollen tips, apically encrusted,7–9µmindiam.attheapex.Hymenialcystidiaabsent;cystidiolesfusoid,thin-walled,smooth,14–17×4–5µm;basidiabarrel-shapedtocapitate, withfoursterigmataandasimplebasalseptum,14–17×7–8µm;basidioles of similar shape to basidia, but smaller. Basidiospores subglobose, hyaline, thin-walled,smooth,withonemediumorsmallguttule,IKI−,CB−,5–5.5(–6)× (4.2–)4.5–5(–5.5)µm,L=5.20µm,W=4.68µm,Q=1.11(n=30/1). Notes. Meripilus malayanus is similar to M. rigidus in morphology by having resupinate and hard corky to rigid basidiomata with a honey buff to grayish Figure 18. Basidiomata of Meripilus malayanus (holotype, Dai 18529). Scale bar: 1 cm.
39 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae brown or deep olive pore surface when dry, distinctly thick-walled subicular hyphae, and thick-walled and apically encrusted hyphoid cystidia. However, the latter has round and smaller pores (10–12 per mm vs. 7–8 permm,Wangetal.2024),smallerbasidiospores(4–4.6×3.2–4µmvs. 5–5.5×4.5–5µm,Wangetal.2024),andsofar,itisonlyknownfromthe type locality in Central America. Figure 19. Microscopic structures of Meripilus malayanus (drawn from the holotype, Dai 18529). a basidiospores b basidia and basidioles c cystidioles d hyphoid cystidia e hyphae from subiculum fhyphaefromtrama.Scalebars:5µm(a); 10µm(b–f).
40 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae In the phylogenetic analyses, Meripilus malayanus is closely related to M. dollingerii, but M. dollingerii has thin-walled and apically encrusted hyphoid cystidia and thin-walled smooth hymenial cystidia (Wang et al. 2024). Rigidoporus adnatus Corner and Polyporus pellicula Jungh. occur in Southeast Asia and have resupinate basidiomata. However, Rigidoporus adnatushassmallerbasidiospores(2.5–3.2×1.7–2µmvs.5–5.5×4.5–5µm, Corner 1987), and Polyporus pellicula haslargerpores(1−3permmvs.7–8 per mm, Teixeira 1992). Meripilus niveomarginatus Y.C. Dai, Chao G. Wang & Yuan Yuan, sp. nov. MycoBank No: 856734 Figs 20, 21 Etymology. Niveomarginatus (Lat.): refers to the species having a white sterile margin when fresh. Diagnosis. Differs from other Meripilus species by resupinate basidiomata with orange-yellow to peach when fresh, white sterile margin when fresh, angular to irregular pores of 6–8 per mm, thick-walled and apically encrusted hyphoidcystidia,broadlyellipsoidbasidiosporesmeasuring4.2–5.2×4–4.6µm. Type. CHINA • Guangdong Province, Zhaoqing, Dinghushan Nature Reserve, on rotten wood of Pinus massoniana, 28 April 2018, Dai 18540A (BJFC027008, holotype). Description. Basidiomata annual, resupinate, soft corky, and without odor or taste when fresh, becoming hard corky to somewhat rigid upon drying, up to 15 cm long, 6 cm wide, and 4 mm thick at center. Pore surface brownish vinaceous, orange-yellow to peach when fresh, becoming reddish brown after bruising, pinkish buff, vinaceous gray, dark brown to black when dry; sterile margin distinct, thinning out, white when fresh, cream to olivaceous buff when dry, up to 2 mm wide; pores angular to irregular, 6–8 per mm; dissepiments thin, slightly lacerate. Subiculum cream, corky, up to 1 mm thick. Tubes paler than pore surface, cinnamon buff, hard corky when dry, up to 3 mm long. Hyphal system monomitic; generative hyphae simple septate, hyaline,smooth,IKI−,CB+;tissuesbecomingdarkbrowninKOH.Subicular hyphae distinctly thick-walled with a wide lumen, rarely branched and simpleseptate,moreorlessflexuous,interwoven,4–8µmindiam.Tramal hyphae distinctly thick-walled with a wide lumen, occasionally branched, moderately simple septate, slightly flexuous, subparallel along the tubes, agglutinated,4–6µmindiam.Hyphoidcystidiapresent,arisingfromtramal hyphae and completely embedded in trama, not projecting from the hymenium, sometimes projecting from the dissepiment edge, thick-walled with swollentips,apicallyencrusted,10–15µmindiam.attheapex.Hymenial cystidiaabsent;cystidiolesfusoid,thin-walled,smooth,13–14×4–5µm; basidia barrel-shaped to capitate, with four sterigmata and a simple basal septum,14–17×7–9µm;basidiolesofsimilarshapetobasidia,butsmaller. Basidiospores broadly ellipsoid, hyaline, thin-walled, smooth, with one mediumorsmallguttule,IKI−,weaklyCB+,(4–)4.2–5.2(–5.5)×(3.8–)4– 4.6(–4.8)µm,L=4.78µm,W=4.16µm,Q=1.13–1.17(n=90/3).
41 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae Additional specimens examined. CHINA • Fujian Province, Quanzhou, Qingyuanshan, on fallen trunk of Pinus massoniana, 23 September 2017, Dai 18268 (BJFC025793); • Hainan Province, Baisha County, Yinggeling Nature Reserve, on rotten wood of Pinus latteri, 10 June 2017, Dai 17695 (BJFC025227); • Zhejiang Province, Wuyi County, on rotten wood of Pinus massoniana, 13 October 2023, Dai 26373 (BJFC043923). Notes. Meripilus niveomarginatus is similar and related to M. rigidus by the resupinate and rigid basidiomata with a pinkish buff to vinaceous gray or dark brown pore surface when dry, distinctly thick-walled generative hyphae, and thick-walled and apically encrusted hyphoid cystidia. However, the latter has round and smaller pores (10–12 per mm vs. 6–8 per mm, Wang etal.2024),relativelysmallerbasidiospores(4–4.6×3.2–4µmvs.4.2–5.2 ×4–4.6µm,Wangetal.2024),andsofar,itisonlyknownfromCentral and South America. Similarly, Meripilus albostygius and M. sulphureus also have rigid basidiomata and thick-walled and apically encrusted hyphoid, but M. albostygiushassmallerbasidiospores(4–4.7×3.2–4µmvs.4.2–5.2× 4–4.6µm,Wangetal.2024)and,todate,onlyoccursinCentralAmerica; M. sulphureus has a sulphurous pore surface when fresh and narrower basidiospores(4–5×3.5–4µmvs.4.2–5.2×4–4.6µm,DaiandDai2018). Meripilus roseus (Jia J. Chen & Y.C. Dai) Westph. & Rajchenb. and M. niveomarginatus share a rose or brownish vinaceous pore surface when fresh. However, the former has an almost lacking sterile margin and smaller basidiospores(3.5–4.1×3.1–3.8µmvs.4.2–5.2×4–4.6µm,ChenandDai2021). Poria endoxantha Petch was originally described from Sri Lanka, and it is characterized by a rose-pink to salmon-pink pore surface when fresh, becoming brown or blackish brown upon bruising, thick-walled hyphoid cystidia embedded in trama, globosebasidiosporesof5–7µmindiam.,andgrowthonangiospermwoodinthe Figure 20. Basidiomata of Meripilus niveomarginatus (holotype, Dai 18540A). Scale bar: 1 cm.
48 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae 35 Hyphoid cystidia only thick-walled or thickand thin-walled or absent ......36 – Hyphoid cystidia only thin-walled ...............................................................45 36 Pores 10–12 per mm .....................................................................M. rigidus – Pores 5–9 per mm ......................................................................................37 37 Pore surface sulphur yellow when fresh ................................M. sulphureus – Pore surface white to isabelline or buff yellow, pale brown to peach when fresh .............................................................................................................38 38 Pore surface often nodulose ......................................................... M. vitreus – Pores surface not nodulose........................................................................39 39 Thin-walled and smooth hymenial cystidia present ............ M. emarginatus – Thin-walled and smooth hymenial cystidia absent ...................................40 40 Pore surface becoming reddish brown or pale brown upon bruising ......41 – Pore surface unchanged upon bruising .....................................................42 41 Pore surface vinaceous gray, dark brown to black when dry ....................... .........................................................................................M. niveomarginatus – Pore surface cream to buff when dry......................................... M. eminens 42 Basidiospores<4.5µminlength ............................................ M. revolubilis – Basidiospores≥4.5µminlength ...............................................................43 43 South American species .............................................................. M. robledoi – Asian species ..............................................................................................44 44 Pores round ........................................................................... M. srilankensis – Pores angular ................................................. .......................M. noncontusus 45 Hymenial cystidia present ..........................................................................46 – Hymenial cystidia absent ............................................................................ 47 46 Basidiospores ovoid, Q=1.22–1.28 ................................... M. castanopsidis – Basidiospores broadly ellipsoid to subglobose, Q=1.15–1.20 .....M. dollingerii 47 Basidiospores thick-walled .................................................... M. crystallinus – Basidiospores thin-walled ...........................................................................48 48 Basidiomata juicy when fresh, easily separated from substrate ... M. tibeticus – Basidiomata waxy when fresh, not easily separated from substrate .......... ............................................................................................... M. expallescens Discussion Ceriporia and Meripilus belong to Irpicaceae and Meripilaceae, respectively, and they are distantly related in phylogeny but very similar in morphology, with mostly light-colored hymenophores, a monomitic hyphal system, and cyanophilous generative hyphae bearing simple septa. In the phylogenetic analyses of Irpicaceae (Fig. 1), four distinct groups are nested in the Ceriporia clade: the C. mellita group, the C. pierii group, the C. purpurea group, and the C. viridans group; they are coincident with the previous study (Wang et al. 2023). Ceriporia armeniaca and C. wuyiana are not included in any groups. Ceriporia crassiparietata nested in the C. viridans group, which involved nine species, and the group has resupinate basidiomata with a white to light yellow pore surface when fresh, subicular hyphae wider than tramal hyphae, hyphae frequently branched at a right angle, and lunate to allantoid basidiospores wider than1.5µm.Meripilusisdividedintofivegroups,asmentionedintheprevious study by Wang et al. (2024). Four new species, M. emarginatus, M. malayanus, M. niveomarginatus, and M. noncontusus, nested in Subclade I; among them,
49 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae M. emarginatus and M. noncontusus have soft to ceraceous basidiomata with a white pore surface when fresh but without reddening when bruised, while M. malayanus and M. niveomarginatus have hard, corky to rigid basidiomata with a fuscous pore surface when dry and thick-walled, encrusted hyphoid cystidia. Meripilus albomarginatus nested in Subclade II, and it has resupinate to effused-reflexedbasidiomata,awhitesterilemargin,distinctlythick-walledto almost solid contextual hyphae, and thick-walled, encrusted hyphoid cystidia, whichfitthecommoncharacteristicsoftheotherspeciesinSubcladeII.Meripilus crystallinus is nested in Subclade V, and it has resupinate, soft to ceraceous basidiomata with a white to cream pore surface when fresh, thin-walled and encrusted hyphoid cystidia, and slightly thick-walled basidiospores. Leptoporus has effused-reflexed to pileate basidiomata, a poroid hymenophore, cylindric to allantoid basidiospores, and causes brown rot in gymnosperm wood (Ryvarden and Melo 2017; Liu et al. 2023). However, Ceriporia has resupinate basidiomata with a poroid or smooth hymenophore, cylindrical to allantoid basidiospores, and is associated with white rot. In addition, Leptoporus is only distantly related to any other taxa of Ceriporia (Fig. 1). Thus, it is also regarded as a separate genus in our study. In phylogeny, although four distinct groups are nested in Ceriporia with similar characters, separating them into smaller genera isstillnotjustified.SomespeciesaddressedintheCeriporia clade lack distinct morphological characteristics and scatter across the clade without high support in the phylogeny. In our previous study (Wang et al. 2024), Physisporinus was dividedintofivegroupsinthephylogeneticanalyses,andMeripilus, including two species sequences, was considered a separate genus nested in Physisporinus. Morphologically, species of Physisporinus contain various types of basidiomata, viz.,resupinate,effused-reflexed,andpileate,ellipsoidtoglobosebasidiospores, and hyphoid cystidia that may be present or absent, while Meripilus only has large,multi-pileate,fleshybasidiomata.Wedidnotfurtherprocesstherelationships between Physisporinus and Meripilus due to the differences in macromorphology in Wang et al. (2024). However, Meripilus nested in the Physisporinus clade in all previous studies (Wang and Dai 2022; Wang et al. 2024; Westphalen et al. 2025). In addition, Westphalen et al. (2025) also mentioned that species of thefivecladesinPhysisporinus are overlapping in morphology and transferred all species of Physisporinus to Meripilus. At present, a proposal to merge the two genera (Physisporinus and Meripilus) is a reasonable approach. Karsten (1889) established the genus Physisporinus and segregated it from Poria based on the “fruit-layer separated from basal layer” (Donk 1966). The type species of the genus, Poria vitrea Pers. sensu P. Karst., seems to be an undetermined species because of confusing labels by Karsten on herbarium specimens (Donk 1960, 1966). Accordingly, Donk (1967) transferred Poria vitrea and other typical species of Physisporinus to Rigidoporus Murrill, which is not yet acceptable. Donk (1967) also indicated that the type specimen of Physisporinus vitreus, Poria vitrea Pers. 1796 sensu Pers., may not exist, and mycologists have often confused the species with Polyporus undatus Pers. 1825. Donk (1967)wrotethatthequalificationsare“undulata,subinterrupta;porisobliquis,” and this conception appears to be the same as Poria vitrea described in Lowe (1966) and Poria undata described in Bourdot and Galzin (1928), and the critical feature, thick-walled, encrusted cystidia, was not mentioned. We studied many specimens of Physisporinus undatus and concluded that the cystidia feature is
50 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae physiologically determined—thinor thick-walled, or sometimes not present at all. All of these are possible, and globose, smaller basidiospores and a somewhat nodulose character of older basidiomata are important, which correspond to Poria vitrea. Moreover, Karsten (1883) described Caloporus expallescens P. Karst., which Lowe (1956) determined to be aberrant (not changing color) Physisporinus sanguinolentus, but Pilát (1938) regarded it as a good species similar to P. vitreus. He studied the type and added a new feature that C. expallescens does not detach from the substrate. We observed this feature in three samples: MJ 642/93 (Czechia), MJ 332/94 (Czechia), and Dai 21060 (Belarus). Thus, we still follow the concept of Wang et al. (2024) that P. undatus is a synonym of P. vitreus and P. expallescens is a good species in this study. Meripilus lineatus (Physisporinus lineatus) has effused-reflexed to pileate or stipitate basidiomata, a concentrically zonate-sulcate upper surface, thickwalled hyphoid cystidia with or without crystals, and subglobose to globose basidiospores (Ryvarden 1972; Ryvarden 2015; Ryvarden and Melo 2017; Wang et al. 2024). Meripilus albomarginatus, M. lineatus, M. sublineatus, and M. vinctus form a sister clade in our phylogeny, and they share thick-walled hyphoid cystidia with crystals. In Wang et al. (2024), eight specimens from the subtropical and tropical zones formed a well-supported distinct lineage and are characterized by pileate basidiomata, thick-walled, encrusted hyphoid cystidia, and thin-walled hymenial cystidia with or without crystals; these characters are consistentwiththedefinitionofM. lineatus. Meripilus vinctus (Physisporinus vinctus) is similar to M. lineatus in micromorphology by having thick-walled, encrusted hyphoid cystidia and thin-walled, mammillate hymenial cystidia. However, M. vinctus has mostly resupinate basidiomata and a cinnamon-orange to buff pore surface, while M. lineatus has mostly pileate basidiomata with rich coloration and a distinct concentrically zonate-sulcate pileal surface in macromorphology. Thus, the samples JV 0610/B5, JV 1407/36, JV 0610/A31B, Kout 1807/3, and Cui 16903 were considered as M. vinctus (Physisporinus vinctus) in Wang et al. (2024). Up to now, the thin-walled hymenial cystidia remain an uncertain character distinguishing M. lineatus. Species of Ceriporia and Meripilus grow on different angiosperm and gymnosperm trees, causing a white rot ecology. The host trees of these fungi are summarized in Table 3. Up to now, 67 species are accepted under Ceriporia, including three new species in the present study: 53 species grow on angiosperm wood, four species grow on gymnosperm wood, and nine species grow on both angiosperm and gymnosperm wood. Moreover, the substrate of Ceriporia albobrunnea Ryvarden & Iturr. was recorded as wood without indicating angiosperm or gymnosperm (Ryvarden and Iturriaga 2003). Forty-nine taxa are accepted in Meripilus, including six new species in the present study: 28 grow on angiosperm wood or bamboo, six taxa grow on gymnosperm wood, 14 taxa grow on both angiosperm and gymnosperm wood or Miscanthus, and one species, M. rigidus, grows on ferns. Species in both genera exhibit distinct growth advantages on angiosperm wood, which is largely related to the evolution of plants. In studies of plant origins, the Paleocene epoch of the Paleogene period is typically recognized as a critical phase for the emergence of angiosperm-endemic genera. During this interval, the climate remained warm, continuing Cretaceous patterns, but became more humid, thus creating favorable conditions for biological development (Wang 1989). In addition, most species of both genera evolved from the Paleogene in our dating analyses.
51 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae Table 3. The type locality, host trees and main morphological characteristics of accepted species in Ceriporia and Meripilus. Species DNA sequences Type locality Basidiomata Shape of basdiospores Size of basidiospores (μm) Host trees References Ceriporia allantoidea ＋China: Hunan Resupinate Allantoid 4.5–5×1.1–1.5 Angiosperm (undetermined) Wang et al. (2023) C. allantospora ＋USA Resupinate Allantoid 10–11.5×2.5–3 Angiosperm (Platanus, Sycamore) Gilbertson et al. (1976) C. arbuscula ＋China: Taiwan Resupinate Cylindrical to slightly curved 3–3.5×1–1.5 Gymnosperm (Pinus) Chen et al. (2020) C. armeniaca ＋China: Guangdong Resupinate Lunate to allantoid 4–4.5 × 2–2.2 Angiosperm (undetermined) Present study C. aurantiocarnescens ＋Germany Resupinate Short cylindrical and moderately curved 3.2–4×1.8–2.2 Angiosperm (Populus) Wang et al. (2023) C. bresadolae ＋France Resupinate to effused-reflexed Allantoid 5.9–8.2×1.8–2.3 Angiosperm (Quercus, Ulmus, Rosaceae) and gymnosperm (Pinus, Picea, Juniperus) Spirin et al. (2016), Wang et al. 2023 C. bubalinomarginata ＋China: Henan Resupinate Allantoid 3.5–4.3×1–1.2 Angiosperm (undetermined) Jia et al. (2014) C. crassa ＋China: Hainan Resupinate Allantoid 3.8–4.1×1.2–1.6 Gymnosperm (Pinus) Wang et al. (2023) C. crassiparietata ＋China: Zhejiang Resupinate Lunate to allantoid 4–4.4 × 2.1–2.3 Angiosperm (Quercus, Rosaceae) Present study C. daedaleoides ＋Thailand Resupinate Ellipsoid to slightly curved 3.7–4.1×2–2.3 Angiosperm (undetermined) Wang et al. (2023) C. eucalypti ＋Australia: Victoria Resupinate Allantoid 4–4.4×1.1–1.4 Angiosperm (Eucalyptus) Chen et al. (2022) C. excelsa ＋Sweden Resupinate Lunate to short cylindrical, moderately curved 3.6–4.2×2–2.2 Angiosperm (Betula, Fagus, Fraxinus, Salix, Viburnum) and gymnosperm (Picea) Wang et al. (2023) C. gossypina ＋China: Xizang Resupinate Allantoid 3.5–4×1.8–2 Angiosperm (undetermined) Wang et al. (2023) C. griseoviolascens ＋France Resupinate Bean-shaped 5–6.1×2.5–3.1 Angiosperm (Prunus, Arbutus, Quercus, Populus) Spirin et al. (2016) C. hinnulea ＋China: Fujian Resupinate Lunate to allantoid 3.5–4×2–2.1 Angiosperm (undetermined) Wang et al. (2023) C. humilis ＋Russia Resupinate Narrowly ellipsoid to cylindrical 3.2–4.2×1.9–2.2 Angiosperm (Quercus) Miettinen et al. (2016) C. langloisii ＋USA: Virginia Resupinate Ellipsoid to broadly allantoid 7–9.5×3–4 Angiosperm (undetermined) Burdsall (1984) C. macrospora ＋China: Hainan Resupinate Allantoid 5–7.2×1.6–2 Angiosperm (undetermined) Wang et al. (2023) C. manzanitae ＋USA: California Resupinate Cylindrical to allantoid 5.1–6.2×2.2–2.7 Angiosperm (Arctostaphylos) Spirin et al. (2016) C. mellita ＋France Resupinate Allantoid 5–6×1.5–2 Angiosperm (Quercus, Populus) Wang et al. (2023) C. mpurii ＋Indonesia Resupinate Ellipsoid to narrowly ellipsoid 2.8–3.9×2–2.3 Angiosperm (Spondias, Prunus, Populus) Spirin et al. (2016) C. occidentalis ＋USA: Washington Resupinate Allantoid 5.1–7.1×1.8–2.2 Angiosperm (Platanus, Umbellularia, Corylus) Spirin et al. (2016) C. orientalis ＋China: Zhejiang Resupinate Lunate to allantoid 5.4–6.5×2.8–3.1 Angiosperm (undetermined) Wang et al. (2023) C. pierii ＋France Resupinate Ellipsoid to rarely cylindrical 4.1–5.4×2.4–3.1 Angiosperm (Populus) Miettinen et al. (2016) C. pseudospissa ＋China: Beijing Resupinate Allantoid 5–7.2×1.6–2.1 Angiosperm (undetermined) Wang et al. (2023) C. punctata ＋China: Xinjiang Resupinate Allantoid 4–5×1.7–2.1 Angiosperm (Populus) Wang et al. (2023) C. punicans ＋USA: Pennsylvania Resupinate Short cylindrical, slightly curved 4.1–5.3×1.7–2.1 Angiosperm (undetermined) Spirin et al. (2016) C. purpurea ＋France Resupinate Allantoid 5–8.4×1.7–2.3 Angiosperm (Corylus, Populus, Alnus, Salix, Quercus, Fraxinus, Acer, Tilia, Pyracantha) Spirin et al. (2016) C. reticulata ＋Germany Resupinate Allantoid 7.5–9×3–3.5 Angiosperm (Acacia, Alnus, Acer, Betula etc.) and gymnosperm (Picea) Domanski (1963) C. septocystidia ＋Jamaica Resupinate Allantoid 4.5–6.5×1.5–2 Angiosperm (Populus) and gymnosperm (undetermined) Burdsall (1984)
52 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae Species DNA sequences Type locality Basidiomata Shape of basdiospores Size of basidiospores (μm) Host trees References C. sericea ＋Russia: Khabarovsk Resupinate Cylindrical to slightly curved 3.9–4.8×2.2–2.7 Angiosperm (Tilia) and gymnosperm (Picea, Pinus) Miettinen et al. (2016) C. sinospissa ＋China: Fujian Resupinate Allantoid 5–5.8×1.5–2 Angiosperm (undetermined) Wang et al. (2023) C. sino-viridans ＋China: Hainan Resupinate Lunate to allantoid 3–3.5×1.7–2.2 Angiosperm (undetermined) Chen et al. (2022) C. sordescens ＋USA: New York Resupinate Ellipsoid to narrowly ellipsoid 3.3–4.2×2.1–2.5 Angiosperm (Acer) Miettinen et al. (2016) C. spissa ＋USA: Carolina Resupinate Allantoid 4.2–4.8×1.3–1.6 (Dai 19164) Angiosperm (undetermined) and gymnosperm (Pinus) Rajchenberg (1983), present study C. subbadia ＋USA: Alabama Resupinate Cylindrical to allantoid 4.5–5.8×2–2.6 Angiosperm (Fagus) Wang et al. (2023) C. subviridans ＋China: Yunnan Resupinate Lunate to allantoid 3.3–3.7×1.8–2 Angiosperm (undetermined) Wang et al. (2023) C. torpida ＋Finland Resupinate Short cylindrical, slightly to moderately curved 4.3–5.7×1.9–2.3 Angiosperm (Fagus, Salix) Spirin et al. (2016) C. triumphalis ＋Spain: Canary Island Resupinate Short cylindrical to allantoid 4.1–5×1.8–2.1 Angiosperm (undetermined) Spirin et al. (2016) C. wuyiana ＋China: Zhejiang Resupinate Lunate to allantoid 4.3–5 × 1.7–2 Angiosperm (undetermined) Present study C. viridans (type species) ＋UK Resupinate Cylindrical to allantoid 4–4.6×1.7–2.1(Dai 17003) Angiosperm (Padus, Populus) Ryvarden and Gilbertson (1993), Wang et al. (2023) Ceriporia alania – USA: Hawaiian Island Resupinate Narrowly allantoid 7.5–10×2–2.5 Angiosperm (Metrosideros) Gilbertson and Hemmes (2004) Ceriporia alba – France Resupinate Cylindrical to allantoid 5.5–7×2–2.5 Angiosperm (undetermined) Pieri and Rivoire (1997) Ceriporia albobrunnea – Venezuela Resupinate Cylindrical 4–4.5×1.5 Unknown Ryvarden and Iturriaga (2003) Ceriporia amazonica – Brazil: Amapá Resupinate Ellipsoid 3×2 Angiosperm (undetermined) Soares et al. (2014) Ceriporia angulata – Brazil: Amazonas Resupinate Oblong-ellipsoid 4–4.5×1.7–2.2 Angiosperm (undetermined) Gomes-Silva et al. (2012) Ceriporia aurea – Venezuela Resupinate Cylindrical to allantoid 4–5×2 Angiosperm (undetermined) Ryvarden (2014) Ceriporia camaresiana – France Resupinate Cylindrical to sub-allantoid 5–6×2–3 Angiosperm (Betula. Eucalyptus, Prunus) and gymnosperm (Picea) Ryvarden and Gilbertson (1993) Ceriporia citrina – Costa Rica Resupinate Oblong-ellipsoid to subcylindrical 7–8×3.2–3.5 Angiosperm (undetermined) Mata and Ryvarden (2010) Ceriporia cystidiata – Venezuela Resupinate Allantoid 4–4.5×1 Angiosperm (undetermined) Ryvarden and Iturriaga (2003) Ceriporia dentipora – Ecuador Resupinate Oblong-ellipsoid to cylindrical 5–6×2.5–3 Angiosperm (undetermined) Læssøe and Ryvarden (2010) Ceriporia ellipsospora –Seychelles Resupinate Ellipsoid 3–4×2.5–2.8 Angiosperm (undetermined) Ryvarden (2018) Ceriporia ferrugineocincta – USA: Florida Resupinate Subcylindrical 3.5–5×2–3 Angiosperm (Quercus) and gymnosperm (undetermined) Ryvarden and Johansen (1980) Ceriporia incrustata – Costa Rica Resupinate Ellipsoid 3–3.5×1.8–2 Angiosperm (undetermined) Mata and Ryvarden (2010) Ceriporia kenyensis – Kenya Resupinate Cylindrical 3–4×1–1.2 Angiosperm (undetermined) Decock et al. (2021) Ceriporia leptoderma – Sri Lanka Resupinate Broadly ellipsoid 5–6×3–4 Angiosperm (undetermined) Ryvarden and Johansen (1980) Ceriporia microspora – Costa Rica Resupinate Ellipsoid 3–3.5×1.5–2 Angiosperm (Quercus, Manilkara) Lindblad and Ryvarden (1999)
53 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae Species DNA sequences Type locality Basidiomata Shape of basdiospores Size of basidiospores (μm) Host trees References Ceriporia otakou – New Zealand Resupinate Ovoid to ellipsoid or pip-shaped 4.5–6×2–2.5 Angiosperm (Nothofagus) Cunningham (1947) Ceriporia retamoana – Argentina: Chubut Resupinate Cylindrical, slightly curved 4.5–5×1.2–1.5 Angiosperm (Diostea) Rajchenberg (2000) Ceriporia rhodella – Sweden Resupinate Short cylindrical, slightly curved 3.5–4×1.5–2 Angiosperm (Alnus, Populus, Fagus) and gymnosperm (undetermined) Lombard and Gilbertson (1965) Ceriporia rubescens – Sri Lanka Resupinate Cylindrical 4–5×1.5–2 Angiosperm (undetermined) Ryvarden (2015) Ceriporia straminea – Bolivia Resupinate Ellipsoid 4.5–5.5×2.5–2.8 Angiosperm (undetermined) Ryvarden (2014) Ceriporia subpudorina – — Resupinate Ovoid to ellipsoid, slightly narrowed 5–6×3–3.5 Angiosperm (Salicis) Bondartsev (1953) Ceriporia subspissa – Guyana Resupinate Ellipsoid 5.5–6.5×3–3.5 Angiosperm (undetermined) Aime et al. (2007) Ceriporia totara – New Zealand: Auckland Resupinate Ovoid to subglobose 2.5–3×1.7–2.5 Gymnosperm (Podocarpus, Dacrydium) Buchanan and Ryvarden (1988) Ceriporia vermicularis – France: Réunion Resupinate Cylindrical to allantoid 6–7×0.8–1 Angiosperm (undetermined) Pieri and Rivoire (1997) Ceriporia violacea – Sweden Resupinate Oblong-ellipsoid 3.5–4.5×2–2.4 Gymnosperm (Pinus) Ryvarden and Melo (2017) Meripilus albomarginatus ＋China: Yunnan Annual, effusedreflexed to pileate Subglobose 5.2–6.2 × 4.6–5.7 Angiosperm (undetermined and bamboo) Present study M. albostygius ＋USA: Puerto Rico Annual, resupinate Boadly ellipsoid to subglobose 4–4.7×3.2–4 Angiosperm (undetermined) Wang et al. (2024) M. brasiliensis ＋Brazil Annual, pileate Subglobose 3–5×3–4 Angiosperm (undetermined) Westphalen et al. 2025 M. caesiomarginatus ＋China: Yunnan Annual, resupinate to effused-reflexed Broadly ovoid 4–4.8×3.3–4.1 Angiosperm (undetermined) Wang et al. (2024) M. castanopsidis ＋China: Yunann Annual, resupinate Ovoid 4.8–5.6×3.8–4.3 Angiosperm (Castanopsis) and gymnosperm (Dacrydium) Chen and Dai (2021) M. cinereus ＋Japan Annual, effusedreflexedto pileate Globose 5–6 Angiosperm (Fagus, bamboo, Lithocarpus) and gymnosperm (Pinus) Núñez and Ryvarden (1999) M. crataegi ＋China: Tianjin Annual, effusedreflexed Broadly ellipsoid to subglobose 4.2–5×3.2–4.2 Angiosperm (Crataegus, Lonicera) Wu et al. (2017) M. crocatus ＋Tunisia Annual to perennial, resupinate Ovoid to subglobose 3.5–5.5×3.5–5 Angiosperm (Betula) and gymnosperm (Abies, Picea) Ryvarden (1983), Ryvarden and Melo (2017) M. crystallinus ＋China: Yunnan Annual, resupinate Broadly ellipsoid to ovoid 4.3–5 × 4–4.5Gymnosperm (Picea) Present study M. dollingerii ＋USA: Florida Annual, resupinate Broadly ellipsoid to subglobose 4.5–5.5×4–5 Angiosperm (undetermined) and gymnosperm (Pinus) Wang et al. (2024) M. emarginatus ＋China: Guangdong Annual, resupinate Subglobose to globose 4.8–5.2 × 4.5–5.2 Angiosperm (Miscanthus and other undetermined wood) and gymnosperm (Cunninghamia) Present study M. eminens ＋China: Jilin Annual, resupinate Globose 4.2–6×3.9–5.2 Angiosperm (Quercus, Populus) Dai (1998) M. expallescens ＋Finland Annual, resupinate Ovoid to globose 5.5–6.5×4.5–5.2 Gymnosperm (Picea, Pinus) Wang et al. (2024)
54 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae Species DNA sequences Type locality Basidiomata Shape of basdiospores Size of basidiospores (μm) Host trees References M. furcatus ＋Russia Annual, resupinate Globose 4.5–5.5 Angiosperm (Alnus) Núñez et al. (2001) M. giganteus (type species) ＋Sweden Annual, multipileate Subglobose, broadly ovoid to broadly ellipsoid 6–6.5×5.5–6 Angiosperm (Quercus, Fagus) Larsen and Lombard (1988) M. lavendulus ＋China: Hainan Annual, pileate Globose 4.2–5×4–5 Angiosperm (undetermined) Wu et al. (2017) M. lineatus ＋Hungary Annual, resupinate, effused-reflexed to pileate or with a stipe Globose 5–6 Angiosperm (Khaya, Ginkgo, Magnolia, Quercus, Cocos, bamboo) and gymnosperm (Metasequoia) Ryvarden (1972), Ryvarden and Johansen (1980), Ryvarden and Gilbertson (1994) M. longicystidius ＋New Zealand Annual, resupinate to effused-reflexed Globose 4.2–5.7 Angiosperm (Eucalyptus, Nothofagus) Buchanan and Ryvarden (2000) M. malayanus ＋Malaysia Annual, resupinate Subglobose 5–5.5 × 4.5–5Angiosperm (undetermined) Present study M. minutissimus ＋Costa Rica Annual, resupinate to effused-reflexed Subglobose to globose 4.1–4.6×4–4.6 Angiosperm (undetermined) Wang et al. (2024) M. neovitreus ＋USA: New Jersey Annual, resupinate to effused-reflexed Broadly ellipsoid to subglobose 4.9–6×4–4.8 Angiosperm (undetermined) Wang et al. (2024) M. niveomarginatus ＋China: Guangdong Annual, resupinate Broadly ellipsoid 4.2–5.2 × 4–4.6 Gymnosperm (Pinus massoniana, P. latteri) Present study M. noncontusus ＋China: Guangdong Annual, resupinate Broadly ellipsoid to subglobose 5–5.5 × 4.2–4.7 Angiosperm (undetermined) Present study M. obscurus ＋Brazil Annual, resupinate Subglobose 3.9–4.8×3.4–4.2 Angiosperm (undetermined) and gymnosperm (Araucaria angustifolia) Westphalen et al. 2025 M. pouzarii ＋Slovakia Annual, resupinate Broadly ellipsoid to ovoid 4.7–6.3×4–4.6 Angiosperm (Alnus, Fagus, Ulmus, Salix) and gymnosperm (Picea) Vampola and Vlasák (2012) M. revolubilis ＋Brazil Annual, resupinate Subglobose to globose 3.4–4.4×3.2–4 Angiosperm (undetermined) Westphalen et al. 2025 M. rhododendri ＋China: Guizhou Annual, resupinate Subglobose 5.3–6.3×5–5.5 Angiosperm (Rhododendron) Wang et al. (2024) M. rigidus ＋Costa Rica Annual, resupinate Broadly ellipsoid to subglobose 4–4.6×3.2–4 Fern (Tectaria) Wang et al. (2024) M. robledoi ＋Argentina Annual, resupinate Subglobose to globose 4.5–5.2×3.2–4 Angiosperm (Nothofagus) and gymnosperm (Araucaria araucana) Westphalen et al. 2025 M. roseus ＋China: Yunnan Perennial, resupinate Subglobose 3.5–4.1×3.1–3.8 Angiosperm (undetermined) Chen and Dai (2021) M. sanguinolentus ＋Germany Annual or biennial, resupinate Ovoid to subglobose 6–7×5–6 Angiosperm (Quercus, Alnus) and gymnosperm (Picea, Abies) Wang et al. (2024) M. srilankensis ＋Sri Lanka Annual, resupinate Broadly ellipsoid to subglobose 5–5.8×4–4.8 Angiosperm (undetermined) Wang et al. (2024)
55 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae Species DNA sequences Type locality Basidiomata Shape of basdiospores Size of basidiospores (μm) Host trees References M. stillicidiorum ＋Australia Annual, resupinate Broadly ellipsoid to subglobose 5–5.7×4–4.8 Angiosperm (Eucalyptus) Wang et al. (2024) M. subfurcatus ＋China: Jilin Annual, resupinate Broadly ellipsoid to subglobose 5.3–6.4×4.5–5.6 Angiosperm (Larix) and gymnosperm (Picea) Wang et al. (2024) M. sublineatus ＋China: Yunnan Annual, resupinate to effused-reflexed Subglobose to globose 4.8–5.6×4.5–5.2 Angiosperm (bamboo and other undetermined wood) Wang et al. (2024) M. sulphureus ＋Singapore Annual, resupinate Subglobose to globose 4–5×3.5–4 Angiosperm (undetermined) Dai and Dai (2018) M. sumstinei ＋USA Annual, multipileate Subglobose to broadly ellipsoid 5–5.5×4–4.5 Angiosperm (undetermined) Larsen and Lombard (1988) M. tamilnaduensis ＋India Annual, Pileate broadly ellipsoid to subglobose 4.5–5.8×4–5 Angiosperm (Azadirachta) Crous et al. (2023) M. tibeticus ＋China: Xizang Annual, resupinate Broadly ellipsoid 4.8–5.5×3.7–4.3 Gymnosperm (Abies, Pinus, Picea) Wu et al. (2017) M. vinctus ＋Dominican Republic Perennial, resupinate to effused-reflexed Ovoid to subglobose 4–5.5×3–4 Angiosperm (undetermined) and gymnosperm (Picea) Ryvarden (1972), Setliff (1972), Ryvarden and Johansen (1980) M. vitreosanguineus ＋Czechia Annual, resupinate Broadly ellipsoid to subglobose 5–6.1×4–5 Gymnosperm (Pinus, Picea, Abies) Wang et al. (2024) M. vitreus ＋Finland Annual, resupinate Ovoid to globose 5–5.5×4–4.5 Angiosperm (Fagus, Ulmus) and gymnosperm (Pinus, Picea, Abies) Wang et al. (2024) M. yunnanensis ＋China: Yunnan Annual, resupinate subglobose 4–5.5×3.5–5 Angiosperm (undetermined) Cai et al. (2023) Meripilus sp. 1 ＋USA: California Annual, resupinate Ellipsoid to broadly ellipsoid 5–6×3.8–4.5 Gymnosperm (Picea, Thuja) Wang et al. (2024) Meripilus sp. 2 ＋Czechia Annual, resupinate Broadly ellipsoid 4.6–5.2×3.8–4.2 Angiosperm (Alnus) Wang et al. (2024) Meripilus sp. 3 ＋Czechia Annual, resupinate Broadly ellipsoid or ovoid 5.5–6.4×4.7–5.2 Angiosperm (Alnus) and gymnosperm (Picea) Wang et al. (2024) Meripilus africanus –São Tomé Annual, effusedreflexedto pileate Subglobose 3–3.5×2.5–3 Angiosperm (Olea capensis) Decock and Ryvarden (2021) M. cataractus – Zimbabwe Annual, resupinate Globose 4–5 Angiosperm (undetermined) Ryvarden (2019) M. resinosus – Uganda Annual, resupinate Globose 2.5–3 Angiosperm (undetermined) Ipulet and Ryvarden (2005) Abbreviations used: + = Present; – = Absent. Bold = new taxa.
56 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae In the dating analyses, we estimated the divergence times of studied taxa using crown ages, as stated in Varga et al. (2019). The divergence time of the Polyporales, including Irpicaceae and Meripilaceae, with a mean crown age of 148.25 Myr, occurred during the late Jurassic, which aligns with the divergence times of ordinal-level taxonomic units in Agaricomycetes during the Jurassic period (Varga et al. 2019). Our result is consistent with the divergence time ranges of orders proposed by Zhao et al. (2017) and Ji et al. (2022). Irpicaceae and Meripilaceae are estimated to have emerged at the junction of the early and late Cretaceous, with mean crown ages of 108.9 Myr and 97.23 Myr, respectively. Ceriporia and Meripilus are estimated at 83.61 Myr and 81.38 Myr, respectively, emerging in the late Cretaceous. Therapiddiversificationofangiosperms,alongwithnumerousanimalandinsect groups during the midto late Cretaceous, combined with the successful proliferationoffloweringplantsaround100millionyearsago,ledtoterrestrial biodiversity exceeding marine biodiversity for the first time in Earth’s geological history. Following their emergence in the late Jurassic, angiosperms underwent explosive radiation during the early Cretaceous, ultimately displacing gymnosperm-dominated forests primarily composed of conifers and cycads (Ramírez-Barahona et al. 2020). The late Cretaceous mass extinction event caused catastrophic devastation to plant communities while simultaneously triggering a surge in saprotrophic organisms such as fungi—non-photosynthetic life forms that derived nutrients by decomposing dead plant matter—though this flourishingofdecomposersprovedshort-livedinthepost-apocalypticecosystem. Ceriporia and Meripilus likely emerged after this mass extinction. Species in Ceriporia occurred between around 63 Myr (C. griseoviolascens) and 5 Myr (C. occidentalis and C. manzanitae), and they all evolved from the Paleogene, becoming more prevalent in the Neogene. Similarly, species in Meripilus occurred between around 54 Myr (M. srilankensis) and 6 Myr (M. yunnanensis and Meripilus sp. 3), and they also evolved from the Paleogene, becoming more prevalent in the Neogene. In addition, the hosts of both genera are angiosperm wood. In studies of plant evolution, the Paleocene epoch of the Paleogene period stands as a critical phase for the origination of angiosperm-endemic genera. During this interval, the climate maintained Cretaceous-like warmth while becoming more humid than the Cretaceous baseline, creating favorable climatic conditions that drove biotic proliferation (Wang 1989). Thus, we speculate that the timing and evolutionary process of the two genera are somewhat coincident. Ceriporia is estimated to have emerged in the early Cretaceous, and species in it occurred between 6.2 and 103.33 Myr, and they all evolved from the early Cretaceous, becoming more prevalent in the Neogene (Wang et al. 2023). Among them, C. griseoviolascens and Leptoporus are estimated at 103.33 Myr and 88.4 Myr, respectively. In our study, the divergence time of the Ceriporia clade evolved at 83.61 Myr in the late Cretaceous, posterior to the results of Wang et al. (2023). In addition, species in Ceriporia occurred between around 63 Myr (C. griseoviolascens) and 5 Myr (C. occidentalis and C. manzanitae). The phylogenies of C. griseoviolascens and Leptoporus are unstable in Ceriporia, which makes a difference for their divergence times. In the dating analyses, the divergence times of Polyporales, including taxonomic classes, were estimated using more comprehensive species sequences. Climatic records across geological ages reveal that the Mesozoic Era—namely the Triassic, Jurassic, and Cretaceous periods—was characterized by globally
57 IMA Fungus 16: e161336 (2025), DOI: 10.3897/imafungus.16.161336 Chao-Ge Wang et al.: Studies in phylogeny and divergence times of Irpicaceae and Meripilaceae warm yet predominantly arid conditions. The Cenozoic Era witnessed a climatic shift, transitioning into a more humid and progressively warmer environment, culminating in the climatically frigid conditions of the Quaternary Ice Age. From the late Cretaceous to the Eocene of the Paleogene period, the Qinghai-Tibet Plateau began experiencing crustal shortening, thickening, and progressive uplift, driven by the ongoing collision and accretion between the Indian and Eurasian plates, which facilitated the region’s sustained elevation during this orogenic phase (Li 1995). Most species of Ceriporia and Meripilus emerged during this period. Dynamically shifting climatic regimes and evolving topography exert both directandindirectcontrolsonfloristicdistributionpatterns,acceleratingdiversification. The Hengduan Mountains and eastern Himalayan floristic regions constitutetheprimaryconvergencezonefornortherntemperateflora,serving as the modern distribution center for numerous critical alpine plant lineages; significantly,theSino-HimalayanFloristicSubkingdomisrecognizedastheevolutionarycradleofNorthernHemispheretemperateflora(Sun2002).Thereason for this phenomenon also lies in the change of the climate in the Arctic region during the Cenozoic Tertiary. From the late Cretaceous to the Paleogene, the Arcticsustainedatemperateclimatethatsupportedthermophilictemperateflora; as the climate cooled, these plants underwent progressive southward migration (Zachos et al. 2001). Studies revealed that during the Eocene of the Paleogene period,EastAsiahostedaluxuriantthermophilicflora,whichwassubsequently supplanted by temperate thermophilic coniferous-broadleaved mixed forests by the Miocene. Intriguingly, the most representative extant refugia of Paleogene floralelementsarenowlocalizedinSoutheast Asia (Tiffney1985;Budantsev 1992, 1994). However, the species of Ceriporia and Meripilus distributed in Southeast Asia account for only a minority at present. In addition to species diversity, the origin, dispersal, and evolution of Ceriporia and Meripilus will be the focus of further studies by collecting more ecological information on the species. Spongipellis is a genus that usually has pileate basidiomata, and although it is addressed in the Meripilaceae main clade, it occurred during the Neogene period with a mean crown age of 13.67 Myr. Nevertheless, the divergence time of Meripilus, with a mean crown age of 81.38 Myr, occurred during the late Cretaceous. Perhaps Spongipellis will be treated as an independent family, but this is outside the scope of our current study. Additional information Conflict of interest The authors have declared that no competing interests exist. Ethical statement No ethical statement was reported. Use of AI No use of AI was reported. Adherence to national and international regulations All the fungal strains used in this study have been legally obtained, respecting the Convention on Biological Diversity (Rio Convention).
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