Entoloma hygropileum sp. nov. (Entolomataceae, Basidiomycota) from a ‘brejo de altitude’ forest in Brazil and its Western Gondwana counterpart

Entoloma hygropileum sp. nov. 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(Entolomataceae, Basidiomycota) from a ‘brejo de altitude’ forest in Brazil and its Western Gondwana counterpart Leonardo Firmino da Silva, Ricardo Koroiva, Felipe Wartchow This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-5243657/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 3 You are reading this latest preprint version Abstract A new species of Entoloma subg. Entoloma is described from a forest fragment in Northeast Brazil. The species description is based on macro and micro-anatomical characterization of the basidiomata, and molecular phylogenetic analyses of ITS and 28S LSU region of nuclear ribosomal DNA. Entoloma hygropileum sp. nov. is distinguished by the tricholomatoid to somewhat clitocyboid habit with very moist to almost slimy and glabrous brownish pileus, broadly adnate lamellae with subdecurrent tooth, proportionally short stipe, isodiametric to subisodiametric basidiospores 6.8–8.6 × 6.1–7.6 µm, absence of hymenial cystidia and a callidermic pileipellis. The most similar species is E. djaense that primarily differs in the proportionally much longer stipe and the lamellae attachment. They present a pileipellis construction unique among the genus, suggesting a Western Gondwana distribution. Africa Agaricales Agaricomycetes Agaricomycotina palaeogeology Neotropic systematic taxonomy Figures Figure 1 Figure 2 Figure 3 Figure 4 Figure 5 Figure 6 Introduction Entoloma sect. Calliderma was described as a section of Rhodophyllus by Romagnesi ( 1941 : 45) corresponding to species with conical-umbonate pileus and a pileipellis a palisade with continuous fusiform elements and subpellis (i.e., hypoderm with loosely interwonen hyphe), with R. callidermus as type species (Romagnesi 1956 ). Later, Noordeloos ( 1983 ) transferred this section to Entoloma , and then Largent ( 1994 ) erected this section as the genus Calliderma . Finally, Noordeloos and Gates ( 2012 ) and Morgado et al. ( 2013 ) confirmed that Calliderma belongs to an integral clade corresponding to the subgenus Entoloma . Coi mbra (2014) compiled 271 Entolomatace species occurring in Central and South American countries, of which more than 100 only from Brazil and Guyana and five taxa treated under Calliderma . In recent years, new records for the genus have been made in the northeastern region of this country (Wartchow 2006 ; Wartchow et al. 2011 ; Alves and Nascimento 2012 ; Coimbra et al. 2013 ). Regarding to taxa with callidermatoid pileipellis, Dennis ( 1953 ), Horak ( 1977 , 1982 , 1983 ), Karstedt and Capelari (2010, as Calliderma ), Aime et al. ( 2010 , as Calliderma ), Wartchow et al. ( 2011 ), and Coimbra et al. ( 2013 ) reported the following taxa (see new combinations and names in Blanco-Dios 2015 ): Entoloma brasiliensis , E. caeruleosplendens , E. foldatsii , E. ipirangaennsis , E. obtusisporum , E. peculiare , E. pruinatocutis , E. pruinatum , and E. tucuchense from Argentina, Brazil, Guyana, and Venezuela. In this study, a similar species to E. djaense (originally treated as belonging to the callidermatoid group by Largent et al. 2020 ) was found in a highland region with humid forests enclaves surrounding by Caatinga (called ‘brejo de altitude’) in the Borborema Plateau. However, we found it that corresponds to as an undescribed taxon, which suggests a Western Gondwanan (sensu Trompette 1997 ) distribution, and macro- and microscopic description as well phylogenetic analysis data are presented for the new species described herein. Material and Methods Sample Collection The ‘Parque Estadual Mata do Pau-Ferro’ is a ‘brejo de altitude’ forest located in region called Borborema Plateau in Northeast Brazil in an altitude up to 600 meters above sea level with predominantly rain forest vegetation (Andrade et al. 2006 ; Oliveira et al. 2006 ). In the region, the most important families are Rubiaceae , Malvaceae , Asteraceae , Convolvulaceae , Solanaceae , and Fabaceae (Barbosa et al. 2004 ). This forest, although corresponding a Conservation Unit, suffers high degree of edge effect (Campos et al. 2018 ; Marques et al. 2019 ). Morphological studies Descriptions of macromorphological features and photographs were made from fresh material. Colors were described subjectively and coded according to Kramer ( 2004 ), with color codes noted in parentheses. Dried basidiomata were sectioned and rehydrated in KOH 3% and water, and stained with Congo Red. The basidiospores were measures excluding the hilar appendix, and basidia excluding the sterigmata. A Zeiss Primo Star Binocular Biological Microscope with built-in camera and ZEN software 3.1 imaging software was used to obtain the micrographies. Microscopic analysis follow Largent ( 1994 ). In the taxonomic descriptions, “x =” refers to mean dimensions of structures, “Q” refers to the mean quotient of the length of structures divided by their width. The holotype is deposited in JPB herbarium (Herbário Lauro Pires Xavier, Universidade Federal da Paraíba). DNA extraction and sequencing To confirm the identification of our specimens, the total genomic DNA of a specimen was extracted using the DNeasy Plant Mini Kit (Qiagen, Germany). PCR amplifications were done for complete internal transcribed spacers 1 and 2 and the 5.8S rDNA (nuc-ITSrDNA) bounded by primers ITS7 and ITS8 (Dentinger et al. 2010 ) and 28S rDNA gene bounded by primers LR0R and the reverse primer LR7 (Moncalvo et al. 2000 ). PCR conditions for amplification consisted of 1× buffer, dNTP at 0.2 mM, each primer at 0.2 µM, MgCl2 at 2mM, 1U Taq polymerase and 2 µl of template DNA, in a total reaction volume of 25 µl. The PCR cycling program was used for both primer sets: 94°C for 2 min, followed by 35 cycles of 94°C for 30 s, 51°C for 40 s, and 72°C for 1 min and concluding with a 10 min extension at 72°C. PCR products were bidirectional sequenced in ABI 3130 Genetic Analyzer (Applied Biosystems). Data analyses We used BioEdit v. 7.2.5 (Hall 1999 ) to check the sequence quality of the strands by comparison to their respective chromatograms and to assemble and edit if necessary. Sequences were compared with similar and available in the GenBank database ( http://www.ncbi.nlm.nih.gov/genbank/ ) using the BLASTn algorithm. Entoloma sequences present in GenBank were incorporated to analyze for both genetic regions, 29 sequences from LSU and 19 ITS sequences. In our analysis, we included 28S rDNA sequences with more than 400bp. Also, in order to increase robustness in the homology statement and elevate matrix occupancy, long sequences of nuc-ITSrDNA were truncated to cover only this region. In the phylogenetic analysis, were aligned sequences using MUSCLE ( https://www.ebi.ac.uk/Tools/msa/muscle/ ), a module implemented in MEGA v. 7.0.26 software using GTR + G algorithm. Maximum Parsimony (MP) analysis was conducted using PAUP v. 4.0b10 (Swofford 2002 ) with 1000 heuristic bootstrap replicates, using random step-wise addition, and holding one tree at each step. Maxtrees were set to 1000, TBR branch swapping algorithm was used to assess branch support. Maximum Likelihood (ML) trees with RAxML v. 8.2.12 (Stamatakis 2014 ) using GTR GAMMA I model with 1.000 bootstrap replicates from MEGA v. 7.0.26. A Bayesian analysis (PP) was constructed with MrBayes v. 3.2.6 (Ronquist et al. 2012 ) launched from TOPALi v. 2.5 (Milne et al. 2004 ). The best available model of evolution was selected with jModel Test v. 3.0.4 (Posada 2008 ) (nuc-ITSrDNA, GTR + G; 28S rDNA, GTR + I + G). Two independent parallel was runs, sampling every 1.000th generations for 10 6 million total generations by TOPALi v. 2.5, with a burning value of 25%. Sequences generated in this study ITS and LSU were deposited in GenBank (NCBI) under accession OR365603 and OR365604 respectively. Results Phylogenetic analysis Based on a BLAST search of NCBIs GenBank nucleotide database using the ITS and LSU sequence, the closest hits were Entoloma djaense , [GenBank MN066547, LSU sequence identity = 869/933 (93,14%); MN069539, ITS sequence identity = 530/650 (81,54%)] and E. olivaceocoloratum [GenBank KT339287, LSU sequence identity = 722/815 (88,59%); KT339257, ITS sequence identity = 168/176 (95%)]. Phylogeny using ITS and LSU barcode (Figs. 1 , 2 ) also showed high similarity in all analyses MP, ML and BI, with well supported bootstrap (BS) with node 97/100/0.99 (Fig. 1 ), with E. hygropileum and E. djaense in the same branch. If considering all branches including E. olivaceocoloratum the node is more supported 100/100/0.99 (Fig. 2 ). Phylogenetic trees were constructed for all analyzes and presented similar topology for the Bayesian analysis, which was chosen for plotting the data. Trees has been deposited at TreeBASE under study number 30942 Taxonomic treatement Entoloma hygropileum L.F. Silva & Wartchow, sp. nov. Figures 3 – 5 MycoBank: MB851355 Type: Brazil. Paraíba: Areia, Parque Estadual Mata do Pau-Ferro, Trilha do Cumbe, 24 Jul 2019, F. Wartchow FW 15/2019 (holo: JPB 65667 ! ) Etimology from Greek hygro (= moist, wet) and pilum (= cap, pileus); due the very moist pileus surface. Basidiome small and stout, tricholomatoid to slightly clitocyboid. Pileus 42 mm, plane-convex, brown (oac722) then darker (oac727) sometime grayish brown (darker than OAC 768) or with yellow tints (oac803) at center; surface smooth, very moist to almost slimy, glabous, shiny; context 4 mm thick, watery white, unchanging; odor slightly sweet. Lamellae broadly adnate with short-decurrent tooth, cream (oac815) to salmon (oac655), distant, 5 mm broad; edge entire, concolorous to side; lamellulae frequent, with diverse lengths, attenuate. Stipe 30 × 5 mm, eccentric, cream, surface, apparently longitudinal, very slightly, appressed squamullose, glabrous; context whitish, unchanging, solid. Basidiospores 6.8–8.6 × 6.1–7.6, (x̅= 7.6 ± 0.5 × 7.1 ± 0.4 µm; Q = 1.00–1.20; Q = 1.10 ± 0.3), isodiametric to sometimes subisodiametric, angular with 5 to 6 blunt angles, pinkish pale, slightly thick-walled; content as large gutule; hilar appendix conic, sublateral. Basidia 34.6–50.9 (–51) × 7.1–9.7 (–10.2) µm ( x̅= 40.9 ± 4.4 × 7.7 ± 0.8 µm; Q = 5.3 ± 0.3) mostly slender clavate to sometimes subcylindric, hyaline, thin-walled, with 4 sterigmata ranging to 3 µm high. Pleurocystidia and cheilocystidia absent. Lamellae trama regular, with narrow hyaline hyphae of variable widths, clamps-connections present, 16.6–61.7 × 3.9–8.8 µm. Pileipellis callidermic, two layered ranging to 55 µm thick; suprapellis to 15 µm thick made of scattered pale colored versiform pileocystidia up to 14–40 × 10–15 µm, lageniform to ventricose or saccate sometimes subcylindrical, thin walled, not encrusted; subpellis to 30 µm thick with more compact inflated short-celled hyphae about 5 cells deep 7.5–12.7 × 6.1–9.2 µm, also pale colorless, and versiforme inflated cells and somewhat radially oriented hyphae, not encrusted. Caulocystidia very occasional 4.9–10.8 × 2.4–5.9 µm, clavate, pale pigmented, thin walled. Stipe trama with longitudinally oriented hyphae of variable lengths 29.4–63.7 × 3.9–10.7 µm, subparallel. Oleiferous hyphae and lipid globules absent. Pigmentation cytoplasmic, not encrusted. Clamp connections commonly found in all examined tissues. Habit and habitat solitary on soil among litter in a ‘brejo de altitude’ forest. Known distribution only known from type locality. Comments about conservation status The new species was collected in a conservation unit with a high degree of illegal human intervention (Barbosa et al. 2017 ) and no management plan exists for this park (Pereira et al. 2019 ; Campos and Lima 2020 ). In addition, there are notices that the edge effects resulting loss of habitat exceed to 50 m in the border-inland gradient (Campos et al. 2018 ; Marques et al. 2019 ). However, data about the population and abundance of E. hygropileum is limited and only future research will show the appropriate threatened classification. Thus, we suggest here the category of data deficient (DD) before facing it against IUCN (2022) criteria of threatened status. Discussion Among the callidermatoid species of Entoloma , the new species is characterized by the tricholomatoid to almost clitocyboid habit with very moist to almost slimy and glabrous brownish pileus, broadly adnate lamellae with subdecurrent tooth, proportionally short stipe, isodiametric to subisodiametric basidiospores 6.8–8.6 × 6.1–7.6 µm, absence of hymenial cystidia, a callidermic pileipellis ( sensu Largent et al. 2020 ), and presence of clamp connections. Entoloma djaense , growing in a monodominant forest containing Gilbertiodendron dewevrei ( Fabacea ) from Cameroon, is the only known species with much similar features. It shares the smooth, moist and shiny pileus, isodiametric basidiospores, presence of clamp connections, and very similar structurally callidermic pileipellis with the subpelis presenting somewhat olive pigmentation and flattener cells. However, the more frequent pileocystididia and the suprapelis with the presence of larger inflated elements 16–32.4 × 9.3–24.2 µm in E. djaense differentiates it from E. hygropileum . Also, the proportionally much longer stipe and the lamellae attachment of African entity, i.e., shallowly sinuate with deccurent tooth, also help in segregate this entity to our new species (Largent et al. 2020 ). These combinations of the pileipellis structure observed in E. djaense and E. hygropileum seem infrequent among Entoloma , and we agree with Largent et al. ( 2020 ) considering both species as having callidermatoid pileipellis. Also, our phylogeny using ITS and LSU barcode also showed high similarity in both analyses with well supported bootstrap. The another species described with viscid when moist and brown colored pileus, isodiametric basidiospores and a ‘agglutinated trichodermial palisade’ pileipellis is the neotropical E. entolomatoides from Martinique and Dominica (Pegler 1983 as Inopilus entolomoides ). However this taxon differs in the account of the larger pileus 40–75 (–90) mm in diam., broader stipe 50–10 mm thick and larger basidiospores 8.5–10.5 × 7–9.5 µm (Pegler 1983 ). Later, Karstedt and Capelari (2010, as Caliderma ) considered as a synonym of E. pruinatocutis after an accuracy microscopic analysis. The recently described E. dicymbophilum Largent & T.W. Henkel from the Guiana Shield shares in the moist pileus surface and similar basidiospores size 6.9–8.9 × 6.2–8.6 µm, but it is quite different in the account of the off-white to pale cream pileus, broadly sinuate lamellae and the tightly entangled, subpericlinal hyphae 3–5 µm wide (Henkel and Largent 2023 ). Entoloma cantharelluloides also from Paraíba shares with E. hygropillum in the decurrent lamellae and isodiametric basidiospores. However, the subvelutinous to subglabrous pileus without mention of viscidity with grayish-fuligineous tints contrasting the whitish then yellowing stipe, and the pileipellis a cutis with narrow hyphae separate this species from E. hygropileum (Singer 1965 ). Entoloma wednae from Pernambuco shares the decurrent lamellae, isodiametric basidiospores, pileipellis divided in two layers and mountain forest habit. However it differs in the surface, texture and pileus color as well absense of inflated elements in the pileipellis (Coimbra et al. 2013 ). Bayesian analysis of nrLSU clusters E. hygropileum sp. nov. and E. djaense in a bracnch forming a clade with two additional sequences of Neotropical E. subg. Entoloma viz., E. olivaceocoloratum , and E. rugosostriatum . They correspond to species bearing a robust tricholomatoid habit, subisodiametric to isodiametric spores and pileipellis a cutis (Largent et al. 2008). Also, at least in E. rugosostriatum we also observed a moist pileus surface. Indeed ITS analysis also clustered the new species with sequences corresponding to the same species mentioned above, confirming their phylogenetic affinitiy. Brief note on the Western Gondwana The site collection of E. hygropileum (in the Borborema Plateau) and E. djaense (Cameroon) (Fig. 6 ), as well the high morphological similarities and phylogeny, suggest a Gondwanan distribution since their pileipellis construction is unique among Entoloma . The Borborema Plateau, where our new species was found, is a very old formation with a Paleoproterozoic basement that goes back to 600 Ma (Fuck et al. 2008 ; Ngako et al. 2008 ). It represents one of the major Brasiliano–Panafrican belt (Vaughan & Pankhurst 2008 ) that is located between the West Africa-São Luis-Parnaiba (north and northwest) and the São Francisco-Congo (at south) (Brito-Neves et al. 1999 ). The evidence is the similar marginal open basin defining the corner of a new-born South Atlantic margin between Pernambuco-Paraiba/Cameroon, resulted from a crustal rupture process across the Equatorial Branch (Darros de Matos 1999 ) during the Cretaceous (Pletsch et al. 2001 ). Thus, based in the evidences emphasized above, E. hygropileum and E. djaense are sister-taxa with well-defined morphology, which is also well-supported in phylogenetic and geographic data. Declarations Conflict of interest The authors declare that they have no confict of interest. Funding ‘Conselho Nacional de Desenvolvimento Científico e Tecnológico’ (CNPq) is acknowledged for funding the following projects: 'Programa de Pesquisas em Biodiversidade' (PPBio Proc. 60/2009), ‘Fungos agaricoides em áreas de Mata Atlântica e Caatinga no Estado da Paraíba' (Edital Universal Proc. 420.448/2016-0) and ‘Produtividade em Pesquisa’ for FW (Proc. 307922/2014-6, Proc. 307947/2017-3, Proc. 309652/2020-0 and Proc. 307423/2023-9). We also thank the ‘Universidade Federal da Paraíba’ that is recognized for funding this project through the ‘Chamada Interna Produtividade em Pesquisa’ (PROPESQ/UFPB Nº 06/2021 Cód. PVA13212-2020). RK received financial support (BLD-PDRP) nº 2022/2023 from the ‘Fundação de Apoio à Pesquisa do Estado da Paraíba’ (FAPESQ). Acknowledgement The authors wish acknowledge coordinator of the ‘Laboratório de Anatomia Vegetal’ (Dr. Rivete S. Lima) for authorizing the use and staffs for help to take the microscopic photos. 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Mol Biol Evol 25:1253–1256. 10.1093/molbev/msn083 Romagnesi H (1941) Les Rhodophylles de Madagascar ( Entoloma , Nolanea , Leptonia, Eccilia , Claudopus ). ’ (Paris Romagnesi H (1956) Les Rhodophylles du Congo Belge d’apress les recoltes de Mme. Goossens-Fontana. Bull Jard Bot l'État Bruxelles 26:137–182. https://doi.org/10.2307/3666871 Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Syst Biol 61:539–542. 10.1093/sysbio/sys029 Singer R (1965) Interesting and new agarics from Brazil. Atas Inst Micol Univ Recife 2:15–59 Stamatakis A (2014) RAxML version 8: a tool for phylogenetic analysis and post-analysis of large phylogenies. Bioinformatics 30:1312–1313. https://doi.org/10.1093/bioinformatics/btu033 Swofford DL (2002) PAUP*. Phylogenetic analysis using parsimony (*and other methods). Sinauer Associates, Sunderland, MA Trompette R (1997) Neoproterozoic (~ 600 Ma) aggregation of Western Gondwana: a tentative scenario. Precambrian Res 82:101–112. 10.1016/S0301-9268(96)00045-9 Vaughan APM, Pankhurst RJ (2008) Tectonic overview of the West Gondwana margin. Gondwana Res 14:150–162. 10.1016/j.gr.2007.07.004 Wartchow F (2006) The Neotropical Entoloma dragonosporum ( Agaricales , Basidiomycota ): a new record from Northeast Brazil. Biociências 14:93–94 Wartchow F, Maia LC, Cavalcanti MAQ (2011) New records of Agaricales from Atlantic Forest fragments of Pernambuco, Northeast Brazil. Mycotaxon 118:137–146. 10.5248/118.137 Tables Table 1 and 2 are available in the Supplementary Files section. Supplementary Files Table1.doc Table2.doc Cite Share Download PDF Status: Under Review Version 1 posted Reviewers invited by journal 11 Nov, 2024 Editor assigned by journal 28 Oct, 2024 First submitted to journal 24 Oct, 2024 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-5243657","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":376654152,"identity":"8f8d64e6-2477-466a-98c0-67a2c5bfd203","order_by":0,"name":"Leonardo Firmino da Silva","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA4ElEQVRIiWNgGAWjYBACPgjFzMDAzviA4QODHJDDg18LG1wLM7MB4wwGYxK1MPMQpYW9+diDn3us5YBamD/bthnk8TPwHvuAVwvPsXTDnmfpxkAtbNK5bQbFkg18yTPwapHIMZPgOXA4sYGZ/xhzbtufxA0HeIzxO0z+jZnkH7AWoMMs2wwS9xPUIsFjJg2xhZlBmhGoZQMDIS08aWnSMgfSjdmAfpHsOWdQLHGYLxmvFn72w8ck3xywluNnb2b+8KMMGGLtvYfxakFYB6UTQHFEGkggVcMoGAWjYBQMfwAAfV847O5oVSoAAAAASUVORK5CYII=","orcid":"https://orcid.org/0000-0002-0009-7549","institution":"Universidade Federal de Pernambuco Centro de Biociencias","correspondingAuthor":true,"prefix":"","firstName":"Leonardo","middleName":"Firmino da","lastName":"Silva","suffix":""},{"id":376654153,"identity":"3ce08cfa-1384-4483-9f5e-82279fe607af","order_by":1,"name":"Ricardo Koroiva","email":"","orcid":"","institution":"Universidade Federal do Pará: Universidade Federal do Para","correspondingAuthor":false,"prefix":"","firstName":"Ricardo","middleName":"","lastName":"Koroiva","suffix":""},{"id":376654154,"identity":"39eb51c9-6f8c-4ca5-b2fb-68d19ad664e0","order_by":2,"name":"Felipe Wartchow","email":"","orcid":"","institution":"Universidade Federal da Paraíba: Universidade Federal da Paraiba","correspondingAuthor":false,"prefix":"","firstName":"Felipe","middleName":"","lastName":"Wartchow","suffix":""}],"badges":[],"createdAt":"2024-10-11 06:04:55","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-5243657/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-5243657/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":70224162,"identity":"afec41bf-96cb-4a2f-acdf-273c91b5daa0","added_by":"auto","created_at":"2024-11-29 17:53:44","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":1406415,"visible":true,"origin":"","legend":"\u003cp\u003ePhylogenetic tree derived from Bayesian analysis, based on ITS1 and ITS2 region data. Values are added at the nodes (≥75 % for ML bootstrap / ≥75% for MP bootstrap / ≥0.75 for BI posterior probability, respectively). The scale bar represents the number of nucleotide changes per site. Species type in \u003cem\u003e\u003cstrong\u003eT\u003c/strong\u003e\u003c/em\u003e (capitular italics). The new species are in bold italic.\u003c/p\u003e","description":"","filename":"Fig.1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-5243657/v1/33f998640f7928e578fc5c22.jpg"},{"id":70224406,"identity":"9022f684-46d4-4051-9dd5-aafbdb848cc1","added_by":"auto","created_at":"2024-11-29 18:01:44","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":2201045,"visible":true,"origin":"","legend":"\u003cp\u003ePhylogenetic tree derived from Bayesian analysis, based on nrLSU region data. Values are added at the nodes (≥75 % for ML bootstrap / ≥75% for MP bootstrap / ≥0.75 for BI posterior probability, respectively). The scale bar represents the number of nucleotide changes per site. Species type in \u003cem\u003e\u003cstrong\u003eT\u003c/strong\u003e\u003c/em\u003e (capitular italics). The new species are in bold italic.\u003c/p\u003e","description":"","filename":"Fig.2.jpg","url":"https://assets-eu.researchsquare.com/files/rs-5243657/v1/6172e348ad042b3bd79c0a85.jpg"},{"id":70224407,"identity":"fc96da3b-7981-4029-b744-f0e4b6a3ef15","added_by":"auto","created_at":"2024-11-29 18:01:45","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":1137605,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eEntoloma hygropileu\u003c/em\u003em (holotype). A-D. Many views of the basidiome. a;b. scales = 20 mm.\u003c/p\u003e","description":"","filename":"Fig.3.jpg","url":"https://assets-eu.researchsquare.com/files/rs-5243657/v1/13a19d81a5300e55e22fbb15.jpg"},{"id":70224165,"identity":"b97ef254-8be8-46a0-a7e2-bad7dcbddf4f","added_by":"auto","created_at":"2024-11-29 17:53:44","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":1038078,"visible":true,"origin":"","legend":"\u003cp\u003eMicroscopy \u003cem\u003eEntoloma hygropileum \u003c/em\u003e(holotype). A. Basidiospores. B. Basidia. C. Pileipellis with inflated elements. Scale bars: A = 10 µm; B-C = 20 µm. Photos Felipe Wartchow; drawings by Leonardo Firmino.\u003c/p\u003e","description":"","filename":"Fig.4.jpg","url":"https://assets-eu.researchsquare.com/files/rs-5243657/v1/f2687baec210dc656b6b3cc4.jpg"},{"id":70224166,"identity":"c5a40294-51d2-44c7-acb2-7af7d9ee40c6","added_by":"auto","created_at":"2024-11-29 17:53:44","extension":"jpg","order_by":5,"title":"Figure 5","display":"","copyAsset":false,"role":"figure","size":3766852,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eEntoloma hygropileum \u003c/em\u003e(holotype). A. Basidiospores. B. Basidia. C. Pileus context hyphae showing clamp connections. D. Pileipellis with inflated elements. E. Suprapellis with cells. Scale bars: A = 10 µm; B-C = 20 µm. Photos by Leonardo Firmino.\u003c/p\u003e","description":"","filename":"Fig.5.jpg","url":"https://assets-eu.researchsquare.com/files/rs-5243657/v1/aaeab344fcb392d6c3144161.jpg"},{"id":70224167,"identity":"e03e5e34-b895-4cce-a531-35feee2295cd","added_by":"auto","created_at":"2024-11-29 17:53:44","extension":"jpg","order_by":6,"title":"Figure 6","display":"","copyAsset":false,"role":"figure","size":1877973,"visible":true,"origin":"","legend":"\u003cp\u003eMap with the distribution of \u003cem\u003eE. hygropileum \u003c/em\u003eand \u003cem\u003eE. djaense\u003c/em\u003e. Map generated with mapchart.net.\u003c/p\u003e","description":"","filename":"Fig.6.jpg","url":"https://assets-eu.researchsquare.com/files/rs-5243657/v1/bcea269c419d6dc652d78baa.jpg"},{"id":70224408,"identity":"1683c785-fd6e-41cd-b3b8-6c364ae91adf","added_by":"auto","created_at":"2024-11-29 18:01:53","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":11854074,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-5243657/v1/f2b87602-8c0f-4497-883a-73e20124ab02.pdf"},{"id":70224164,"identity":"d14a7cdb-6b39-439f-9db4-72fe6f0dd5cd","added_by":"auto","created_at":"2024-11-29 17:53:44","extension":"doc","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":53760,"visible":true,"origin":"","legend":"","description":"","filename":"Table1.doc","url":"https://assets-eu.researchsquare.com/files/rs-5243657/v1/4ff29dc32748c005ade786b8.doc"},{"id":70224168,"identity":"c2b79a38-be50-4c6d-8786-d6bab0572c6a","added_by":"auto","created_at":"2024-11-29 17:53:45","extension":"doc","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":55808,"visible":true,"origin":"","legend":"","description":"","filename":"Table2.doc","url":"https://assets-eu.researchsquare.com/files/rs-5243657/v1/8b9c356ad671e33bc00540cf.doc"}],"financialInterests":"","formattedTitle":"Entoloma hygropileum sp. nov. (Entolomataceae, Basidiomycota) from a ‘brejo de altitude’ forest in Brazil and its Western Gondwana counterpart","fulltext":[{"header":"Introduction","content":"\u003cp\u003e \u003cem\u003eEntoloma\u003c/em\u003e sect. \u003cem\u003eCalliderma\u003c/em\u003e was described as a section of \u003cem\u003eRhodophyllus\u003c/em\u003e by Romagnesi (\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e1941\u003c/span\u003e: 45) corresponding to species with conical-umbonate pileus and a pileipellis a palisade with continuous fusiform elements and subpellis (i.e., hypoderm with loosely interwonen hyphe), with \u003cem\u003eR. callidermus\u003c/em\u003e as type species (Romagnesi \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e1956\u003c/span\u003e). Later, Noordeloos (\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1983\u003c/span\u003e) transferred this section to \u003cem\u003eEntoloma\u003c/em\u003e, and then Largent (\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e1994\u003c/span\u003e) erected this section as the genus \u003cem\u003eCalliderma\u003c/em\u003e. Finally, Noordeloos and Gates (\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2012\u003c/span\u003e) and Morgado et al. (\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) confirmed that \u003cem\u003eCalliderma\u003c/em\u003e belongs to an integral clade corresponding to the subgenus \u003cem\u003eEntoloma\u003c/em\u003e.\u003c/p\u003e \u003cp\u003e \u003cstrong\u003eCoi\u003c/strong\u003e \u003cp\u003embra (2014) compiled 271 Entolomatace species occurring in Central and South American countries, of which more than 100 only from Brazil and Guyana and five taxa treated under \u003cem\u003eCalliderma\u003c/em\u003e. In recent years, new records for the genus have been made in the northeastern region of this country (Wartchow \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Wartchow et al. \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Alves and Nascimento \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Coimbra et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Regarding to taxa with callidermatoid pileipellis, Dennis (\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1953\u003c/span\u003e), Horak (\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e1977\u003c/span\u003e, \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e1982\u003c/span\u003e, \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e1983\u003c/span\u003e), Karstedt and Capelari (2010, as \u003cem\u003eCalliderma\u003c/em\u003e), Aime et al. (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2010\u003c/span\u003e, as \u003cem\u003eCalliderma\u003c/em\u003e), Wartchow et al. (\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2011\u003c/span\u003e), and Coimbra et al. (\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) reported the following taxa (see new combinations and names in Blanco-Dios \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2015\u003c/span\u003e): \u003cem\u003eEntoloma brasiliensis\u003c/em\u003e, \u003cem\u003eE. caeruleosplendens\u003c/em\u003e, \u003cem\u003eE. foldatsii\u003c/em\u003e, \u003cem\u003eE. ipirangaennsis\u003c/em\u003e, \u003cem\u003eE. obtusisporum\u003c/em\u003e, \u003cem\u003eE. peculiare\u003c/em\u003e, \u003cem\u003eE. pruinatocutis\u003c/em\u003e, \u003cem\u003eE. pruinatum\u003c/em\u003e, and \u003cem\u003eE. tucuchense\u003c/em\u003e from Argentina, Brazil, Guyana, and Venezuela.\u003c/p\u003e \u003c/p\u003e \u003cp\u003eIn this study, a similar species to \u003cem\u003eE. djaense\u003c/em\u003e (originally treated as belonging to the callidermatoid group by Largent et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) was found in a highland region with humid forests enclaves surrounding by Caatinga (called \u0026lsquo;brejo de altitude\u0026rsquo;) in the Borborema Plateau. However, we found it that corresponds to as an undescribed taxon, which suggests a Western Gondwanan (sensu Trompette \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e1997\u003c/span\u003e) distribution, and macro- and microscopic description as well phylogenetic analysis data are presented for the new species described herein.\u003c/p\u003e"},{"header":"Material and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eSample Collection\u003c/h2\u003e \u003cp\u003eThe \u0026lsquo;Parque Estadual Mata do Pau-Ferro\u0026rsquo; is a \u0026lsquo;brejo de altitude\u0026rsquo; forest located in region called Borborema Plateau in Northeast Brazil in an altitude up to 600 meters above sea level with predominantly rain forest vegetation (Andrade et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Oliveira et al. \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). In the region, the most important families are \u003cem\u003eRubiaceae\u003c/em\u003e, \u003cem\u003eMalvaceae\u003c/em\u003e, \u003cem\u003eAsteraceae\u003c/em\u003e, \u003cem\u003eConvolvulaceae\u003c/em\u003e, \u003cem\u003eSolanaceae\u003c/em\u003e, and \u003cem\u003eFabaceae\u003c/em\u003e (Barbosa et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). This forest, although corresponding a Conservation Unit, suffers high degree of edge effect (Campos et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Marques et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eMorphological studies\u003c/h3\u003e\n\u003cp\u003eDescriptions of macromorphological features and photographs were made from fresh material. Colors were described subjectively and coded according to Kramer (\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2004\u003c/span\u003e), with color codes noted in parentheses. Dried basidiomata were sectioned and rehydrated in KOH 3% and water, and stained with Congo Red. The basidiospores were measures excluding the hilar appendix, and basidia excluding the sterigmata. A Zeiss Primo Star Binocular Biological Microscope with built-in camera and ZEN software 3.1 imaging software was used to obtain the micrographies. Microscopic analysis follow Largent (\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e1994\u003c/span\u003e). In the taxonomic descriptions, \u0026ldquo;x =\u0026rdquo; refers to mean dimensions of structures, \u0026ldquo;Q\u0026rdquo; refers to the mean quotient of the length of structures divided by their width. The holotype is deposited in JPB herbarium (Herb\u0026aacute;rio Lauro Pires Xavier, Universidade Federal da Para\u0026iacute;ba).\u003c/p\u003e\n\u003ch3\u003eDNA extraction and sequencing\u003c/h3\u003e\n\u003cp\u003eTo confirm the identification of our specimens, the total genomic DNA of a specimen was extracted using the DNeasy Plant Mini Kit (Qiagen, Germany). PCR amplifications were done for complete internal transcribed spacers 1 and 2 and the 5.8S rDNA (nuc-ITSrDNA) bounded by primers ITS7 and ITS8 (Dentinger et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2010\u003c/span\u003e) and 28S rDNA gene bounded by primers LR0R and the reverse primer LR7 (Moncalvo et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). PCR conditions for amplification consisted of 1\u0026times; buffer, dNTP at 0.2 mM, each primer at 0.2 \u0026micro;M, MgCl2 at 2mM, 1U Taq polymerase and 2 \u0026micro;l of template DNA, in a total reaction volume of 25 \u0026micro;l. The PCR cycling program was used for both primer sets: 94\u0026deg;C for 2 min, followed by 35 cycles of 94\u0026deg;C for 30 s, 51\u0026deg;C for 40 s, and 72\u0026deg;C for 1 min and concluding with a 10 min extension at 72\u0026deg;C. PCR products were bidirectional sequenced in ABI 3130 Genetic Analyzer (Applied Biosystems).\u003c/p\u003e\n\u003ch3\u003eData analyses\u003c/h3\u003e\n\u003cp\u003eWe used BioEdit v. 7.2.5 (Hall \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1999\u003c/span\u003e) to check the sequence quality of the strands by comparison to their respective chromatograms and to assemble and edit if necessary. Sequences were compared with similar and available in the GenBank database (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttp://www.ncbi.nlm.nih.gov/genbank/\u003c/span\u003e\u003cspan address=\"http://www.ncbi.nlm.nih.gov/genbank/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e) using the BLASTn algorithm. \u003cem\u003eEntoloma\u003c/em\u003e sequences present in GenBank were incorporated to analyze for both genetic regions, 29 sequences from LSU and 19 ITS sequences. In our analysis, we included 28S rDNA sequences with more than 400bp. Also, in order to increase robustness in the homology statement and elevate matrix occupancy, long sequences of nuc-ITSrDNA were truncated to cover only this region.\u003c/p\u003e \u003cp\u003eIn the phylogenetic analysis, were aligned sequences using MUSCLE (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://www.ebi.ac.uk/Tools/msa/muscle/\u003c/span\u003e\u003cspan address=\"https://www.ebi.ac.uk/Tools/msa/muscle/\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e), a module implemented in MEGA v. 7.0.26 software using GTR\u0026thinsp;+\u0026thinsp;G algorithm. Maximum Parsimony (MP) analysis was conducted using PAUP v. 4.0b10 (Swofford \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2002\u003c/span\u003e) with 1000 heuristic bootstrap replicates, using random step-wise addition, and holding one tree at each step. Maxtrees were set to 1000, TBR branch swapping algorithm was used to assess branch support. Maximum Likelihood (ML) trees with RAxML v. 8.2.12 (Stamatakis \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2014\u003c/span\u003e) using GTR GAMMA I model with 1.000 bootstrap replicates from MEGA v. 7.0.26. A Bayesian analysis (PP) was constructed with MrBayes v. 3.2.6 (Ronquist et al. \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2012\u003c/span\u003e) launched from TOPALi v. 2.5 (Milne et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). The best available model of evolution was selected with jModel Test v. 3.0.4 (Posada \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2008\u003c/span\u003e) (nuc-ITSrDNA, GTR\u0026thinsp;+\u0026thinsp;G; 28S rDNA, GTR\u0026thinsp;+\u0026thinsp;I\u0026thinsp;+\u0026thinsp;G). Two independent parallel was runs, sampling every 1.000th generations for 10\u003csup\u003e6\u003c/sup\u003e million total generations by TOPALi v. 2.5, with a burning value of 25%. Sequences generated in this study ITS and LSU were deposited in GenBank (NCBI) under accession OR365603 and OR365604 respectively.\u003c/p\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003ePhylogenetic analysis\u003c/h2\u003e \u003cp\u003eBased on a BLAST search of NCBIs GenBank nucleotide database using the ITS and LSU sequence, the closest hits were \u003cem\u003eEntoloma djaense\u003c/em\u003e, [GenBank MN066547, LSU sequence identity\u0026thinsp;=\u0026thinsp;869/933 (93,14%); MN069539, ITS sequence identity\u0026thinsp;=\u0026thinsp;530/650 (81,54%)] and \u003cem\u003eE. olivaceocoloratum\u003c/em\u003e [GenBank KT339287, LSU sequence identity\u0026thinsp;=\u0026thinsp;722/815 (88,59%); KT339257, ITS sequence identity\u0026thinsp;=\u0026thinsp;168/176 (95%)]. Phylogeny using ITS and LSU barcode (Figs.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, \u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) also showed high similarity in all analyses MP, ML and BI, with well supported bootstrap (BS) with node 97/100/0.99 (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e), with \u003cem\u003eE. hygropileum\u003c/em\u003e and \u003cem\u003eE. djaense\u003c/em\u003e in the same branch. If considering all branches including \u003cem\u003eE. olivaceocoloratum\u003c/em\u003e the node is more supported 100/100/0.99 (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Phylogenetic trees were constructed for all analyzes and presented similar topology for the Bayesian analysis, which was chosen for plotting the data. Trees has been deposited at TreeBASE under study number 30942\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eTaxonomic treatement\u003c/h3\u003e\n\u003cp\u003e \u003cb\u003eEntoloma hygropileum\u003c/b\u003e L.F. Silva \u0026amp; Wartchow, \u003cb\u003esp. nov.\u003c/b\u003e Figures\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e\u0026ndash;\u003cspan refid=\"Fig5\" class=\"InternalRef\"\u003e5\u003c/span\u003e\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eMycoBank: MB851355\u003c/p\u003e \u003cp\u003eType: Brazil. Para\u0026iacute;ba: Areia, Parque Estadual Mata do Pau-Ferro, Trilha do Cumbe, 24 Jul 2019, F. Wartchow FW 15/2019 (holo: JPB 65667\u003cb\u003e!\u003c/b\u003e)\u003c/p\u003e \u003cp\u003e \u003cstrong\u003eEtimology\u003c/strong\u003e \u003cp\u003efrom Greek \u003cem\u003ehygro\u003c/em\u003e (=\u0026thinsp;moist, wet) and \u003cem\u003epilum\u003c/em\u003e (=\u0026thinsp;cap, pileus); due the very moist pileus surface.\u003c/p\u003e \u003c/p\u003e \u003cp\u003eBasidiome small and stout, tricholomatoid to slightly clitocyboid. Pileus 42 mm, plane-convex, brown (oac722) then darker (oac727) sometime grayish brown (darker than OAC 768) or with yellow tints (oac803) at center; surface smooth, very moist to almost slimy, glabous, shiny; context 4 mm thick, watery white, unchanging; odor slightly sweet. Lamellae broadly adnate with short-decurrent tooth, cream (oac815) to salmon (oac655), distant, 5 mm broad; edge entire, concolorous to side; lamellulae frequent, with diverse lengths, attenuate. Stipe 30 \u0026times; 5 mm, eccentric, cream, surface, apparently longitudinal, very slightly, appressed squamullose, glabrous; context whitish, unchanging, solid.\u003c/p\u003e \u003cp\u003eBasidiospores 6.8\u0026ndash;8.6 \u0026times; 6.1\u0026ndash;7.6, (x̅= 7.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.5 \u0026times; 7.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4 \u0026micro;m; Q\u0026thinsp;=\u0026thinsp;1.00\u0026ndash;1.20; Q\u0026thinsp;=\u0026thinsp;1.10\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3), isodiametric to sometimes subisodiametric, angular with 5 to 6 blunt angles, pinkish pale, slightly thick-walled; content as large gutule; hilar appendix conic, sublateral. Basidia 34.6\u0026ndash;50.9 (\u0026ndash;51) \u0026times; 7.1\u0026ndash;9.7 (\u0026ndash;10.2) \u0026micro;m ( x̅= 40.9\u0026thinsp;\u0026plusmn;\u0026thinsp;4.4 \u0026times; 7.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.8 \u0026micro;m; Q\u0026thinsp;=\u0026thinsp;5.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3) mostly slender clavate to sometimes subcylindric, hyaline, thin-walled, with 4 sterigmata ranging to 3 \u0026micro;m high. Pleurocystidia and cheilocystidia absent. Lamellae trama regular, with narrow hyaline hyphae of variable widths, clamps-connections present, 16.6\u0026ndash;61.7 \u0026times; 3.9\u0026ndash;8.8 \u0026micro;m. Pileipellis callidermic, two layered ranging to 55 \u0026micro;m thick; suprapellis to 15 \u0026micro;m thick made of scattered pale colored versiform pileocystidia up to 14\u0026ndash;40 \u0026times; 10\u0026ndash;15 \u0026micro;m, lageniform to ventricose or saccate sometimes subcylindrical, thin walled, not encrusted; subpellis to 30 \u0026micro;m thick with more compact inflated short-celled hyphae about 5 cells deep 7.5\u0026ndash;12.7 \u0026times; 6.1\u0026ndash;9.2 \u0026micro;m, also pale colorless, and versiforme inflated cells and somewhat radially oriented hyphae, not encrusted. Caulocystidia very occasional 4.9\u0026ndash;10.8 \u0026times; 2.4\u0026ndash;5.9 \u0026micro;m, clavate, pale pigmented, thin walled. Stipe trama with longitudinally oriented hyphae of variable lengths 29.4\u0026ndash;63.7 \u0026times; 3.9\u0026ndash;10.7 \u0026micro;m, subparallel. Oleiferous hyphae and lipid globules absent. Pigmentation cytoplasmic, not encrusted. Clamp connections commonly found in all examined tissues.\u003c/p\u003e \u003cp\u003e \u003cstrong\u003eHabit and habitat\u003c/strong\u003e \u003cp\u003esolitary on soil among litter in a \u0026lsquo;brejo de altitude\u0026rsquo; forest.\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eKnown distribution\u003c/strong\u003e \u003cp\u003eonly known from type locality.\u003c/p\u003e \u003c/p\u003e \u003cp\u003e \u003cstrong\u003eComments about conservation status\u003c/strong\u003e \u003cp\u003eThe new species was collected in a conservation unit with a high degree of illegal human intervention (Barbosa et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) and no management plan exists for this park (Pereira et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Campos and Lima \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In addition, there are notices that the edge effects resulting loss of habitat exceed to 50 m in the border-inland gradient (Campos et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Marques et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). However, data about the population and abundance of \u003cem\u003eE. hygropileum\u003c/em\u003e is limited and only future research will show the appropriate threatened classification. Thus, we suggest here the category of data deficient (DD) before facing it against IUCN (2022) criteria of threatened status.\u003c/p\u003e \u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eAmong the callidermatoid species of \u003cem\u003eEntoloma\u003c/em\u003e, the new species is characterized by the tricholomatoid to almost clitocyboid habit with very moist to almost slimy and glabrous brownish pileus, broadly adnate lamellae with subdecurrent tooth, proportionally short stipe, isodiametric to subisodiametric basidiospores 6.8\u0026ndash;8.6 \u0026times; 6.1\u0026ndash;7.6 \u0026micro;m, absence of hymenial cystidia, a callidermic pileipellis (\u003cem\u003esensu\u003c/em\u003e Largent et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), and presence of clamp connections.\u003c/p\u003e \u003cp\u003e \u003cem\u003eEntoloma djaense\u003c/em\u003e, growing in a monodominant forest containing \u003cem\u003eGilbertiodendron dewevrei\u003c/em\u003e (\u003cem\u003eFabacea\u003c/em\u003e) from Cameroon, is the only known species with much similar features. It shares the smooth, moist and shiny pileus, isodiametric basidiospores, presence of clamp connections, and very similar structurally callidermic pileipellis with the subpelis presenting somewhat olive pigmentation and flattener cells. However, the more frequent pileocystididia and the suprapelis with the presence of larger inflated elements 16\u0026ndash;32.4 \u0026times; 9.3\u0026ndash;24.2 \u0026micro;m in \u003cem\u003eE. djaense\u003c/em\u003e differentiates it from \u003cem\u003eE. hygropileum\u003c/em\u003e. Also, the proportionally much longer stipe and the lamellae attachment of African entity, i.e., shallowly sinuate with deccurent tooth, also help in segregate this entity to our new species (Largent et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). These combinations of the pileipellis structure observed in \u003cem\u003eE. djaense\u003c/em\u003e and \u003cem\u003eE. hygropileum\u003c/em\u003e seem infrequent among \u003cem\u003eEntoloma\u003c/em\u003e, and we agree with Largent et al. (\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) considering both species as having callidermatoid pileipellis. Also, our phylogeny using ITS and LSU barcode also showed high similarity in both analyses with well supported bootstrap.\u003c/p\u003e \u003cp\u003eThe another species described with viscid when moist and brown colored pileus, isodiametric basidiospores and a \u0026lsquo;agglutinated trichodermial palisade\u0026rsquo; pileipellis is the neotropical \u003cem\u003eE. entolomatoides\u003c/em\u003e from Martinique and Dominica (Pegler \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e1983\u003c/span\u003e as \u003cem\u003eInopilus entolomoides\u003c/em\u003e). However this taxon differs in the account of the larger pileus 40\u0026ndash;75 (\u0026ndash;90) mm in diam., broader stipe 50\u0026ndash;10 mm thick and larger basidiospores 8.5\u0026ndash;10.5 \u0026times; 7\u0026ndash;9.5 \u0026micro;m (Pegler \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e1983\u003c/span\u003e). Later, Karstedt and Capelari (2010, as \u003cem\u003eCaliderma\u003c/em\u003e) considered as a synonym of \u003cem\u003eE. pruinatocutis\u003c/em\u003e after an accuracy microscopic analysis.\u003c/p\u003e \u003cp\u003eThe recently described \u003cem\u003eE. dicymbophilum\u003c/em\u003e Largent \u0026amp; T.W. Henkel from the Guiana Shield shares in the moist pileus surface and similar basidiospores size 6.9\u0026ndash;8.9 \u0026times; 6.2\u0026ndash;8.6 \u0026micro;m, but it is quite different in the account of the off-white to pale cream pileus, broadly sinuate lamellae and the tightly entangled, subpericlinal hyphae 3\u0026ndash;5 \u0026micro;m wide (Henkel and Largent \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cem\u003eEntoloma cantharelluloides\u003c/em\u003e also from Para\u0026iacute;ba shares with \u003cem\u003eE. hygropillum\u003c/em\u003e in the decurrent lamellae and isodiametric basidiospores. However, the subvelutinous to subglabrous pileus without mention of viscidity with grayish-fuligineous tints contrasting the whitish then yellowing stipe, and the pileipellis a cutis with narrow hyphae separate this species from \u003cem\u003eE. hygropileum\u003c/em\u003e (Singer \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e1965\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cem\u003eEntoloma wednae\u003c/em\u003e from Pernambuco shares the decurrent lamellae, isodiametric basidiospores, pileipellis divided in two layers and mountain forest habit. However it differs in the surface, texture and pileus color as well absense of inflated elements in the pileipellis (Coimbra et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eBayesian analysis of nrLSU clusters \u003cem\u003eE. hygropileum\u003c/em\u003e sp. nov. and \u003cem\u003eE. djaense\u003c/em\u003e in a bracnch forming a clade with two additional sequences of Neotropical \u003cem\u003eE.\u003c/em\u003e subg. \u003cem\u003eEntoloma\u003c/em\u003e viz., \u003cem\u003eE. olivaceocoloratum\u003c/em\u003e, and \u003cem\u003eE. rugosostriatum\u003c/em\u003e. They correspond to species bearing a robust tricholomatoid habit, subisodiametric to isodiametric spores and pileipellis a cutis (Largent et al. 2008). Also, at least in \u003cem\u003eE. rugosostriatum\u003c/em\u003e we also observed a moist pileus surface. Indeed ITS analysis also clustered the new species with sequences corresponding to the same species mentioned above, confirming their phylogenetic affinitiy.\u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eBrief note on the Western Gondwana\u003c/h2\u003e \u003cp\u003eThe site collection of \u003cem\u003eE. hygropileum\u003c/em\u003e (in the Borborema Plateau) and \u003cem\u003eE. djaense\u003c/em\u003e (Cameroon) (Fig.\u0026nbsp;\u003cspan refid=\"Fig6\" class=\"InternalRef\"\u003e6\u003c/span\u003e), as well the high morphological similarities and phylogeny, suggest a Gondwanan distribution since their pileipellis construction is unique among \u003cem\u003eEntoloma\u003c/em\u003e.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe Borborema Plateau, where our new species was found, is a very old formation with a Paleoproterozoic basement that goes back to 600 Ma (Fuck et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Ngako et al. \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). It represents one of the major Brasiliano\u0026ndash;Panafrican belt (Vaughan \u0026amp; Pankhurst \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2008\u003c/span\u003e) that is located between the West Africa-S\u0026atilde;o Luis-Parnaiba (north and northwest) and the S\u0026atilde;o Francisco-Congo (at south) (Brito-Neves et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e1999\u003c/span\u003e). The evidence is the similar marginal open basin defining the corner of a new-born South Atlantic margin between Pernambuco-Paraiba/Cameroon, resulted from a crustal rupture process across the Equatorial Branch (Darros de Matos \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1999\u003c/span\u003e) during the Cretaceous (Pletsch et al. \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2001\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThus, based in the evidences emphasized above, \u003cem\u003eE. hygropileum\u003c/em\u003e and \u003cem\u003eE. djaense\u003c/em\u003e are sister-taxa with well-defined morphology, which is also well-supported in phylogenetic and geographic data.\u003c/p\u003e \u003c/div\u003e"},{"header":"Declarations","content":"\u003cp\u003e \u003ch2\u003eConflict of interest\u003c/h2\u003e \u003cp\u003eThe authors declare that they have no confict of interest.\u003c/p\u003e \u003c/p\u003e\u003ch2\u003eFunding\u003c/h2\u003e \u003cp\u003e\u0026lsquo;Conselho Nacional de Desenvolvimento Cient\u0026iacute;fico e Tecnol\u0026oacute;gico\u0026rsquo; (CNPq) is acknowledged for funding the following projects: 'Programa de Pesquisas em Biodiversidade' (PPBio Proc. 60/2009), \u0026lsquo;Fungos agaricoides em \u0026aacute;reas de Mata Atl\u0026acirc;ntica e Caatinga no Estado da Para\u0026iacute;ba' (Edital Universal Proc. 420.448/2016-0) and \u0026lsquo;Produtividade em Pesquisa\u0026rsquo; for FW (Proc. 307922/2014-6, Proc. 307947/2017-3, Proc. 309652/2020-0 and Proc. 307423/2023-9). We also thank the \u0026lsquo;Universidade Federal da Para\u0026iacute;ba\u0026rsquo; that is recognized for funding this project through the \u0026lsquo;Chamada Interna Produtividade em Pesquisa\u0026rsquo; (PROPESQ/UFPB N\u0026ordm; 06/2021 C\u0026oacute;d. PVA13212-2020). RK received financial support (BLD-PDRP) n\u0026ordm; 2022/2023 from the \u0026lsquo;Funda\u0026ccedil;\u0026atilde;o de Apoio \u0026agrave; Pesquisa do Estado da Para\u0026iacute;ba\u0026rsquo; (FAPESQ).\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e \u003cp\u003eThe authors wish acknowledge coordinator of the \u0026lsquo;Laborat\u0026oacute;rio de Anatomia Vegetal\u0026rsquo; (Dr. Rivete S. Lima) for authorizing the use and staffs for help to take the microscopic photos.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAime MC, Largent DL, Henkel TW, Baroni TJ (2010) The Entolomataceae of the Pakaraima Mountains of Guyana IV: new species of \u003cem\u003eCalliderma\u003c/em\u003e. Paraeccilia Trichopilus Mycologia 102:633\u0026ndash;649. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.3852/09-162\u003c/span\u003e\u003cspan address=\"10.3852/09-162\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAlves MH, Nascimento CC (2012) \u003cem\u003eEntoloma virescens\u003c/em\u003e (Sacc.) E. Horak ex Courtec., 1986, (Agaricales: Entolomataceae): the first record for the Caatinga Biome, Cear\u0026aacute;, Brazil. 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Bioci\u0026ecirc;ncias 14:93\u0026ndash;94\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWartchow F, Maia LC, Cavalcanti MAQ (2011) New records of Agaricales from Atlantic Forest fragments of Pernambuco, Northeast Brazil. Mycotaxon 118:137\u0026ndash;146. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e10.5248/118.137\u003c/span\u003e\u003cspan address=\"10.5248/118.137\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003eTable 1 and 2 are available in the Supplementary Files section.\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"biologia","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"biol","sideBox":"Learn more about [Biologia](http://link.springer.com/journal/11756)","snPcode":"11756","submissionUrl":"https://www.editorialmanager.com/biol/default2.aspx","title":"Biologia","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Africa, Agaricales, Agaricomycetes, Agaricomycotina, palaeogeology, Neotropic, systematic, taxonomy","lastPublishedDoi":"10.21203/rs.3.rs-5243657/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-5243657/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eA new species of \u003cem\u003eEntoloma\u003c/em\u003e subg. \u003cem\u003eEntoloma\u003c/em\u003e is described from a forest fragment in Northeast Brazil. The species description is based on macro and micro-anatomical characterization of the basidiomata, and molecular phylogenetic analyses of ITS and 28S LSU region of nuclear ribosomal DNA. \u003cem\u003eEntoloma hygropileum\u003c/em\u003e sp. nov. is distinguished by the tricholomatoid to somewhat clitocyboid habit with very moist to almost slimy and glabrous brownish pileus, broadly adnate lamellae with subdecurrent tooth, proportionally short stipe, isodiametric to subisodiametric basidiospores 6.8\u0026ndash;8.6 \u0026times; 6.1\u0026ndash;7.6 \u0026micro;m, absence of hymenial cystidia and a callidermic pileipellis. The most similar species is \u003cem\u003eE. djaense\u003c/em\u003e that primarily differs in the proportionally much longer stipe and the lamellae attachment. They present a pileipellis construction unique among the genus, suggesting a Western Gondwana distribution.\u003c/p\u003e","manuscriptTitle":"Entoloma hygropileum sp. nov. (Entolomataceae, Basidiomycota) from a ‘brejo de altitude’ forest in Brazil and its Western Gondwana counterpart","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-11-29 17:53:39","doi":"10.21203/rs.3.rs-5243657/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewersInvited","content":"","date":"2024-11-11T12:46:12+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2024-10-28T04:18:12+00:00","index":"","fulltext":""},{"type":"submitted","content":"Biologia","date":"2024-10-24T04:25:29+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"biologia","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"biol","sideBox":"Learn more about [Biologia](http://link.springer.com/journal/11756)","snPcode":"11756","submissionUrl":"https://www.editorialmanager.com/biol/default2.aspx","title":"Biologia","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"55f37c24-b4a1-4912-a84c-0535e20b1ebd","owner":[],"postedDate":"November 29th, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2024-11-29T17:53:40+00:00","versionOfRecord":[],"versionCreatedAt":"2024-11-29 17:53:39","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-5243657","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-5243657","identity":"rs-5243657","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}