A re-evaluation of Neotropical Junghuhnia s.lat. (Polyporales, Basidiomycota) based on morphological and multigene analyses.
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Abstract
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A re-evaluation of Neotropical Junghuhnia s.lat. (Polyporales, Basidiomycota) based on morphological and multigene analyses
Abstract
Junghuhnia is a genus of polypores traditionally characterised by a dimitic hyphal system with clamped generative hyphae and presence of encrusted skeletocystidia. However, recent molecular studies revealed that Junghuhnia is polyphyletic and most of the species cluster with Steccherinum, a morphologically similar genus separated only by a hydnoid hymenophore. In the Neotropics, very little is known about the evolutionary relationships of Junghuhnia s.lat. taxa and very few species have been included in molecular studies. In order to test the proper phylogenetic placement of Neotropical species of this group, morphological and molecular analyses were carried out. Specimens were collected in Brazil and used for DNA sequence analyses of the internal transcribed spacer and the large subunit of the nuclear ribosomal RNA gene, the translation elongation factor 1-α gene, and the second largest subunit of RNA polymerase II gene. Herbarium collections, including type specimens, were studied for morphological comparison and to confirm the identity of collections. The molecular data obtained revealed that the studied species are placed in three different genera. Specimens of Junghuhnia carneola represent two distinct species that group in a lineage within the phlebioid clade, separated from Junghuhnia and Steccherinum, which belong to the residual polyporoid clade. Therefore, the new genus Geesterania is proposed including two species, G. carneola comb. nov. and G. davidii sp. nov. Neotropical specimens identified as Junghuhnia nitida represent a different lineage from the European species and are described as Steccherinum neonitidum sp. nov. In addition, the new combinations Steccherinum meridionale, Steccherinum polycystidiferum and Steccherinum undigerum, as well as the new name Flaviporus tenuis, are proposed.
INTRODUCTION
The genus Junghuhnia (Basidiomycota, Polyporales) is characterised by a dimitic hyphal system, clamped generative hyphae, small basidiospores with variable shape and presence of large encrusted skeletocystidia. Traditionally, it has been separated from Steccherinum only by the hymenophore configuration (poroid in Junghuhnia and hydnoid in Steccherinum), but microscopically both genera present almost identical features. Antrodiella is also related but distinguished by the lack of cystidia (Ryvarden 1991). However, recent studies including molecular data (Miettinen et al. 2012) showed that the morphological segregation of Junghuhnia and Steccherinum is not supported by molecular phylogeny. Many species treated as Junghuhnia in the literature group with species with hydnoid hymenophore in the Steccherinum clade, separated from J. crustacea, the type of Junghuhnia. Furthermore, there are no microscopic characters enabling delimitation between Junghuhnia s.str. (including J. crustacea) and Junghuhnia s.lat. (species included in Steccherinum clade) due to convergent micromorphology. In the Neotropics, 11 names were previously reported in Junghuhnia: J. carneola, J. chlamydospora, J. globospora, J. meridionalis, J. minuta, J. neotropica, J. nitida, J. polycystidifera, J. semisupiniformis, J. subundata and J. undigera (Ryvarden 1985, 2007, 2015, Lindblad & Ryvarden 1999, Westphalen et al. 2012). However, molecular data of all of them are unavailable and their phylogenetic position is unknown. Therefore, the current study addresses a knowledge gap, using molecular and morphological data to elucidate the systematics of Neotropical Junghuhnia and related genera.
MATERIAL AND METHODS
Morphological analysis
Specimens were collected in southern Brazil in Santa Catarina, São Paulo and Rio Grande do Sul States. Additional specimens of Neotropical Junghuhnia s.lat. from BPI, FLOR, ICN, O, PRM and S herbaria (Thiers 2016) were studied for morphological revision. For micromorphology observations, hand-cut sections of the basidiomes were prepared on microscope slides with 3 % KOH solution and stained with 1 % aqueous phloxine. When necessary, small pieces of the basidiomes were kept in NaOH 3 % solution under 60 °C for about 12 h in order to obtain a better separation of the hyphae and interpret the hyphal system. The samples were then used for preparing microscope slides. All microscopic structures observed were measured with aid of an eyepiece micrometer and, when possible, a minimum of twenty measurements of each structure were taken. The abbreviations and codes used for the measurements are: Lm × Wm = means of length and width, Q = range of length/width ratios, Qm = length/width mean and n = x/y (x = number of measurements from a given number; y of specimens). Drawings of the microstructures were made with the aid of a drawing tube. Cresyl violet was used for observation on metachromatic reactions.
Culturing, mating tests and nuclear staining
Sporeprints were obtained from fresh basidiomes and used to prepare monosporic and polysporic cultures. Cultures were grown and kept in malt extract agar (MEA) or potato dextrose agar (PDA) at 25 °C. In order to clarify the mating system of the species in question, mating tests were performed according to Hallenberg (1984). At least nine monosporic cultures of each species were used for mating tests. For nuclear behaviour verification, Giemsa stain was used for nuclear staining technique following Boidin (1958). After staining, the mycelia were observed under microscope and nuclear behaviour was classified as Normal, Astatocoenocytic, Heterocytic or Holocoenocytic (Boidin 1971). Cultures obtained were kept at the Botany Institute of São Paulo Culture Collection and Culture Collection of Basidiomycetes (Institute of Microbiology, Czech Academy of Sciences, Prague). Additional cultures form CIEFAP culture collection (CIEFAPcc) were also used in this study.
DNA extraction and amplification
DNeasy Plant Mini Kit (QIAGEN) and Power Soil DNA Isolation Kit (MO BIO) were used for DNA extraction from cultures and basidiomes, respectively. In cases where basidiome extraction was not successful, the 10 first steps of DNeasy Plant Mini Kit were followed and DNA extraction of the samples proceeded using magnetic-bead technology of MagNA Pure compact system (Roche). When necessary, DNA was purified using DNA Clean & Concentration Kit (Zymo Research). DNA amplification of the internal transcribed spacer (ITS) and large subunit (LSU) regions of ribosomal RNA gene were performed using ITS5/ITS4-Basidio (Nikolcheva & Bärlocher 2004) primer combination for ITS and LR0R/LR6 for LSU. The PCR regimes followed Tomšovský et al. (2010a). For translation elongation factor 1-α gene (TEF1-α; primers: 983F/2218R), touchdown PCR with gradually reduced annealing temperature (60–50 °C) was performed (Rehner & Buckley 2005). For the second largest subunit of RNA polymerase II gene (RPB2; primers fRPB25F/bRPB271R), a touch-down protocol modified from Rehner & Buckley (2005) EF-α protocol was performed. In some samples where PCR did not present good results, a nested PCR was performed according to Tomšovský et al. (2010b). Amplification products were sent for purification and sequencing in MacroGen Ltd. (Korea).
Phylogenetic analysis
All the sequences obtained were initially checked in Chromas 2.6 (http://technelysium.com.au/wp/chromas/) and BioEdit (Hall 1999) software and, when necessary, adjusted manually to correct minor sequencing errors. Two combined dataset of DNA sequences were prepared, one using ITS-LSU and one with four molecular markers (ITS, LSU, TEF1-α, RPB2). Reference sequences for both datasets were chosen based on studies of Miettinen et al. (2012), Floudas & Hibbett (2015), Binder et al. (2013) and through BLAST searches in the NCBI (National Center for Biotechnology http://www.ncbi.nlm.nih.gov), and are summarised in Table 1. The datasets were aligned using MAFFT version 7 (http://mafft.cbrc.jp/alignment/server/). In order to remove highly variable regions with low homology in the 4 gene dataset, Gblocks curation was performed on ITS sequences and intron regions of TEF1-α and RPB2 sequences were deleted manually. The evolutionary models for each gene and for combined datasets were inferred with the jModelTest 2.c1.4 (Darriba et al. 2012) using the Akaike information criterion. The best fit models selected were TrN+I+G for TEF1-α region, TIM2+I+G for RPB2 and GTR+I+ G for ITS, LSU, and for both combined datasets. Bayesian trees and posterior probabilities were estimated with the MrBayes 3.2.6 software (Ronquist et al. 2012). The analysis was run for 10 million generations, sampling every 1 000 generations. Burnin was set to 10 % of the trees. Maximum Likelihood analyses were conducted in RAxML-HPC v. 8 (Stamatakis 2014) with a GTRCAT model of evolution. Number of bootstrap replicates was halted automatically (autoMRE) and 252 replicates were used for the ITS-LSU dataset and the 204 for four marker dataset. All analyses were conducted in CIPRES Science Gateway (Miller et al. 2010).
Table 1
List of sequences used in this study. * Sequences obtained in this study.
| Specimen | ITS | GenBank No. | ||
|---|---|---|---|---|
| LSU | TEF1-α | RPB2 | ||
| Antella americana (KHL 11949) | JN710509 | JN710509 | JN710711 | – |
| Antrodia serialis (KHL 12010) | JX109844 | JX109844 | JX109898 | JX109870 |
| Antrodiella faginea (KHL 11977) | JN710514 | JN710514 | JN710712 | – |
| Antrodiella ichnusana (RS 13892) | JN710516 | JN710516 | – | – |
| Antrodiella onychoides (OM 2312) | JN710517 | JN710517 | – | – |
| Antrodiella pallescens (X1080) | JN710518 | JN710518 | – | – |
| Antrodiella romellii (OM 7429) | JN710520 | JN710520 | – | – |
| Antrodiella semisupina (JL 372) | JN710521 | JN710521 | – | – |
| Aurantioporus fissilis (BRNM 699803) | HQ728292 | HQ729002 | – | – |
| Bjerkandera adusta (BRNM 771948) | KT305935 | KT305935 | KT305938 | – |
| Bjerkandera fumosa (BRNM 771947) | KT305937 | KT305937 | – | – |
| Butyrea japonica (MN 1065) | JN710556 | JN710556 | JN710718 | – |
| Butyrea luteoalba (KHL 13238b) | JN710558 | JN710558 | JN710719 | – |
| Byssomerulius corium (FP-102382) | KP135007 | KP135230 | KP134921 | – |
| Ceriporia viridans (KHL 8765) | AF347109 | AF347109 | – | – |
| Ceriporiopsis gilvescens (TN 5516) | HQ659222 | HQ659222 | – | – |
| Flaviporus brownii (TL 9883) | JN710537 | JN710537 | – | – |
| Flaviporus brownii (X462) | JN710538 | JN710538 | – | – |
| Flaviporus brownii* (MCW 362/12) | KY175008 | KY175008 | KY175022 | – |
| Flaviporus liebmannii (LR 44553) | JN710540 | JN710540 | – | – |
| Flaviporus minutus* (MCW 442/13) | KY175001 | KY175001 | – | – |
| Flaviporus minutus* (MCW 356/12) | KY175002 | KY175002 | – | – |
| Flaviporus minutus* (MCW 386/12) | KY175003 | KY175003 | – | – |
| Flaviporus sp. (PO 60302006) | JN710542 | JN710542 | – | – |
| Flaviporus subundatus* (MCW 367/12) | KY175004 | KY175004 | – | – |
| Flaviporus subundatus* (MCW 457/13) | KY175005 | KY175005 | – | – |
| Flaviporus subundatus* (RP 332) | KY175006 | KY175006 | – | – |
| Frantisekia mentschulensis (AH 1377) | JN710544 | JN710544 | – | – |
| Geesteriana carneola* (MCW 388/12) | KY174999 | KY174999 | KY175013 | KY175011 |
| Geesteriana carneola* (MCW 474/13) | KY175000 | KY175000 | KY175014 | – |
| Geesteriana davidii* (MCW 370/12) | KY174997 | KY174997 | KY175015 | – |
| Geesteriana davidii* (MCW 396/12) | KY174998 | KY174998 | KY175016 | KY175012 |
| Gloeoporus dichrous (BRNM 709971) | EU546097 | FJ496709 | – | – |
| Gloeoporus taxicola (SK 0075) | JX109847 | JX109847 | JX109901 | JX109873 |
| Hyphodermella corrugata (KHL 3663) | EU118630 | EU118630 | – | – |
| Irpex lacteus (DO 421/951208) | JX109852 | JX109852 | JX109911 | JX109882 |
| Junghuhnia crustacea (OM 13852) | JN710554 | JN710554 | – | – |
| Junghuhnia crustacea (OM 2954) | JN710553 | JN710553 | – | – |
| Junghuhnia kotlabae* (PRM 870890) | KY175007 | KY175007 | – | – |
| Junghuhnia micropora (VS 2652) | JN710559 | JN710559 | JN710720 | – |
| Junghuhnia sp1 (OM12860) | JN710563 | JN710563 | JN710723 | – |
| Junghuhnia sp3 (OM 8625) | JN710564 | JN710564 | JN710724 | – |
| Junghuhnia sp4 (X1362) | JN710565 | JN710565 | JN710725 | – |
| Junghuhnia sp5 (OM 13865) | JN710566 | JN710566 | JN710726 | – |
| Luteoporia albomarginata (Dai 15229) | KU598873 | KU598878 | – | – |
| Luteoporia albomarginata (Dai 15240) | KU598874 | KU598879 | – | – |
| Metuloidea rhinocephala (X460) | JN710562 | JN710562 | – | – |
| Mycoacia fuscoatra (KHL13275) | JN649352 | JN649352 | JX109908 | JX109879 |
| Mycoacia nothofagi (KHL13750) | GU480000 | GU480000 | – | – |
| Nigroporus vinosus (BHS2008-100) | JX109857 | JX109857 | – | – |
| Odoria alborubescens (BRNU 627479) | JQ821319 | JQ821318 | – | – |
| Phanerochaete chrysosporium (BKM-F-1767) | HQ188436 | GQ470643 | HQ188379 | – |
| Phanerochaete chrysosporium (HHB-6251) | KP135094 | KP135246 | – | KP134954 |
| Phanerochaete sordida (KHL 12054) | EU118653 | EU118653 | – | – |
| Phlebia chrysocreas (HHB-6333) | KP135358 | KP135263 | – | KP134908 |
| Phlebia uda (FP-101544) | KP135361 | KP135232 | – | KP134909 |
| Phlebia nitidula (N 020830) | EU118655 | EU118655 | – | – |
| Phlebia radiata (FPL 6140) | AY854087 | AF287885 | AY885156 | AY218502 |
| Phlebioporia bubalina (Dai 13168) | KC782526 | KC782528 | – | – |
| Rhizochaete brunnea (MR 11455) | AY219389 | AY219389 | – | – |
| Rhizochaete radicata (FD-123) | KP135407 | KP135279 | – | KP134937 |
| Scopuloides hydnoides (KHL 11916) | EU118665 | EU118665 | – | – |
| Scopuloides rimosa (HHB-7042) | KP135350 | KP135282 | – | KP134903 |
| Skeletocutis nivea (ES2008-1) | JX109858 | JX109858 | – | – |
| Spongipellis pachyodon (AFTOL-ID 705) | DQ249277 | AY629322 | DQ028599 | DQ408123 |
| Spongipellis spumeus (BRNM 712630) | HQ728288 | HQ728288 | – | – |
| Steccherinum autumnale (VS 2957) | JN710549 | JN710549 | JN710716 | – |
| Steccherinum bourdotii (RS 10195) | JN710584 | JN710584 | – | – |
| Steccherinum collabens (KHL 11848) | JN710552 | JN710552 | JN710717 | – |
| Steccherinum fimbriatellum (OM 2091) | JN710555 | JN710555 | – | – |
| Steccherinum fimbriatum (KHL 11905) | JN710530 | JN710530 | – | – |
| Steccherinum formosanum (TFRI 652) | EU232184 | EU232268 | – | – |
| Steccherinum lacerum (TN 8246) | JN710557 | JN710557 | – | – |
| Steccherinum meridionalis* (MR 284) | KY174992 | KY174992 | KY175019 | KY175009 |
| Steccherinum meridionalis* (MR 11086) | KY174993 | KY174993 | – | – |
| Steccherinum meridionalis* (MR 10466) | KY174994 | KY174994 | – | – |
| Steccherinum neonitidum* (MCW 371/12) | KY174990 | KY174990 | KY175017 | KY175010 |
| Steccherinum neonitidum* (RP 79) | KY174991 | KY174991 | KY175018 | – |
| Steccherinum nitidum (FP-105195) | KP135323 | KP135227 | – | KP134964 |
| Steccherinum nitidum (KHL 11903) | JN710560 | JN710560 | JN710721 | – |
| Steccherinum nitidum* (MT 33/12) | KY174989 | KY174989 | – | – |
| Steccherinum ochraceum (KHL 11902) | JN710590 | JN710590 | JN710730 | JN710738 |
| Steccherinum polycystidiferum* (MCW 419/12) | KY174995 | KY174995 | KY175021 | – |
| Steccherinum polycystidiferum* (RP 140) | KY174996 | KY174996 | – | – |
| Steccherinum pseudozilingianum (MK 1004) | JN710561 | JN710561 | JN710722 | – |
| Steccherinum robustius (G1195) | JN710591 | JN710591 | – | – |
| Steccherinum sp1 (OM 13705) | JN710592 | JN710592 | JN710731 | – |
| Steccherinum sp2 (OM 9300) | JN710593 | JN710593 | – | – |
| Steccherinum sp3 (OM 14391) | JN710594 | JN710594 | JN710732 | – |
| Steccherinum sp4 (OM 13755) | JN710596 | JN710596 | – | – |
| Steccherinum tenue (KHL 12316) | JN710598 | JN710598 | JN710733 | – |
| Steccherinum tenuispinum (OM 8065) | JN710599 | JN710599 | – | – |
| Steccherinum tenuispinum (LE231603) | KM411452 | KM411452 | – | – |
| Steccherinum tenuispinum (VS 2116) | JN710600 | JN710600 | – | – |
| Steccherinum undigerum* (MCW 426/13) | KY174986 | KY174986 | KY175020 | – |
| Steccherinum undigerum* (MCW 472/13) | KY174987 | KY174987 | – | – |
| Steccherinum undigerum* (MCW 496/14) | KY174988 | KY174988 | – | – |
| Trametopsis cervina (TJV 93216T) | JN165020 | JN164796 | JN164882 | JN164877 |
| Trulla dentipora (LR 40822) | JN710512 | JN710512 | – | – |
| Trulla duracina (LR 43589) | JN710513 | JN710513 | – | – |
RESULTS
Molecular data obtained show that the studied Neotropical Junghuhnia spp. are polyphyletic and occur in three different genera (Fig. 1, ,2):2): Steccherinum (supported in ML and BA of both ITS-LSU and combined dataset), Flaviporus (unsupported in ML in ITS-LSU dataset and supported in ML and BA of both ITS-LSU and combined dataset) and a new genus in the phlebioid clade (supported in ML and BA of both datasets). None of the studied species is related to the type of the genus, J. crustacea. The ITS-LSU dataset includes 1 952 positions including 1 070 conserved, 825 variable and 136 singleton. The four gene dataset includes 3 482 positions including 2 191 conserved, 1 287 variable and 317 singleton.
Phylogenetic tree of ITS-LSU regions conducted by Bayesian analysis. Numbers at branches indicate maximum likelihood bootstrap proportion and Bayesian posterior probability values. The asterisk (*) marks different topologies in both analyses. The bar indicates number of expected substitutions per position.
The phylogenetic tree of ITS, LSU, TEF1-α, RPB2 regions conducted by Bayesian analysis. Numbers at branches indicate maximum likelihood bootstrap proportion and Bayesian posterior probability values. The asterisk (*) marks different topologies in both analyses. The bar indicates number of expected substitutions per position.
The description of a new polypore genus in the phlebioid clade, Geesterania, including J. carneola, is supported by both the phylogenetic data and morphological characters. Moreover, J. carneola is in fact a complex of two species. Our data also show that the Neotropical J. nitida represents a different species separated from the European specimens (the species was described from Europe) and here is described as a new species, Steccherinum neonitidum. In addition, three new combinations: Steccherinum meridionale, Steccherinum polycystidiferum and Steccherinum undigerum, and one new name: Flaviporus tenuis, are proposed and the position of J. subundata in Flaviporus (Ginns 1980) is confirmed. Full descriptions and illustrations of the new genus and the two new species are presented below, as well as comments on the new combinations and name proposed. Identification keys for the genera and species of Neotropical Junghuhnia s.lat. are also provided. Further discussion on the four Neotropical species not included in this study (J. chlamydospora, J. globospora, J. neotropica and J. semisupiniformis) is presented.
TAXONOMY
Geesterania Westphalen, Tomšovský & Rajchenb., gen. nov. — MycoBank MB822658; Fig. 3, ,44
a, b. Geesterania carneola; c, d. G. davidii. — a, c. Fresh basidiomes; b, d. detail of pore surface in dried basidiomes. — Scale bars: a, c = 2 cm; b, d = 1 mm. — Photos: a. D.L. Komura; b, d. M.C. Westphalen; c. M.A. Reck.
a–g. Geesterania davidii; h. G. carneola. — a. Basidiospores; b. basidia; c. cystidioles; d. generative hyphae; e. skeletal hyphae; f. cystidium from the trama; g. cystidia from the dissepiments; h. basidiospores. — Scale bar = 10 μm. — Drawn by M.C. Westphalen.
Type. Geesterania carneola (Bres.) Westphalen & Rajchenb.
Etymology. Named in honour of Rudolph Arnold Maas Geesteranus (The Hague 1911–Oegstgeest 2003) for his extensive contribution to the mycology, including studies of the genus Steccherinum and of several phlebioid species.
Diagnosis. Geesterania is characterised by the basidiomes becoming reddish when bruised, the abundant, long, thin- to thick-walled, finely encrusted skeletocystidia, the metachromatic skeletal hyphae and cystidia, a bipolar mating system and astatocoenocytic nuclear behaviour.
Basidiomes resupinate, adnate, becoming reddish when bruised and/or upon drying, soft to fleshy when fresh, becoming harder and somewhat waxy when dried. Hymenophore poroid; pores regular to irregular, round to angular. Hyphal system dimitic, generative hyphae clamped, skeletal hyphae metachromatic, hyphae IKI-. Skeletocystidia present in the trama and dissepiments, cylindrical to clavate, encrusted at the apex with thin crystals, usually very long and appearing as skeletal hyphae ends, metachromatic, IKI-. Basidiospores ellipsoid to sub-cylindrical, hyaline, smooth, IKI-.
Notes — Geesterania is characterised by resupinate, somewhat fleshy basidiomes typically becoming reddish when bruised or dried. The colour change is variable, and in some basidiomes only some reddish spots appear while others become pale reddish brown to flesh coloured throughout. Microscopically, the very long finely encrusted (skeleto)cystidia are typical. In the trama, the cystidia are usually markedly thick-walled and long, arising from skeletal hyphae that slowly increase in diameter. For that reason, the distinction between skeletal hyphae and cystidia sometimes is not clear and the cystidia may be interpreted as wide encrusted skeletal hyphae ends. Here we choose to use the term cystidia as these structures can be very conspicuous in the dissepiments and in the trama, sometimes having many crystals covering a widened apex. In addition, the skeletal hyphae and cystidia are metachromatic, while the generative hyphae, hymenium and basidiospores are non metachromatic in Cresyl Violet. Geesterania differs from other polypore genera exactly by this combination of characteristics, which is unique and unknown in other poroid fungi. Even though the type species had been placed in Junghuhnia, morphological, biological and molecular data support it as a different genus. Junghuhnia s.str. and Steccherinum present long encrusted skeletocystidia. However, in both genera the cystidia are encrusted with large crystals and can be easily distinguished from the skeletal hyphae, as they present a more abrupt widening and more evident encrustation. In addition, the skeletal hyphae in Junghuhnia and Steccherinum are non metachromatic in Cresyl Violet and only the cystidia present a variable metachromatic reaction, while in Geesterania the cystidia and skeletal hyphae are metachromatic throughout. Furthermore, culture studies had already shown that G. carneola is bipolar and astatocoenocytic, while species of Junghuhnia s.str. and Steccherinum s.lat. are tetrapolar and have a normal nuclear behaviour (David & Rajchenberg 1985, Rajchenberg 2011). The combination of bipolarity and astatocoenocyty is characteristic in the phlebioid clade (Rajchenberg 2011), and our molecular data confirm the position of Geesterania there (Fig. 1). Phylogenetically, the closest related polypore species known is Phlebioporia bubalina (Chen & Cui 2014) but, morphologically, it substantially differs by presenting a monomitic hyphal system with simple septate dextrinoid hyphae and lack of cystidia. The recently described genus and species Luteoporia albomarginata (Wu et al. 2016) is also phylogenetically related to Geesterania, presenting similar resupinate basidiomes with presence of cystidia-like structures. However, Luteoporia differs in being monomitic, turning red in contact with KOH, and having differently shaped thin-walled cystidia.
Geesterania carneola (Bres.) Westphalen & Rajchenb., comb. nov. — MycoBank MB822659; Fig. 3a–b, 4a–g
≡Poria carneola Bres., Hedwigia 35: 282. 1896, basionym.
≡Junghuhnia carneola (Bres.) Rajchenb., Revista Invest. Agropecu., Ser. 5, 19: 45. 1984.
Specimens examined. Brazil, Paraná, Paranaguá, Prox. a Rodovia PR 508, 2 Sept. 2013, M.C. Westphalen 458/13 (SP 446225); Rio Grande do Sul, São Francisco de Paula, FLONA, 30 Apr. 2012, M.C. Westphalen 388/12 (SP 446186); Santa Catarina, Blumenau, Moller (S F159730), syntype; ibid., Moller 284 (S F15834), syntype; ibid., Moller 465 (S F15836, syntype); Itapoá, RPPN Volta Velha, 17 Nov. 2012, M.C. Westphalen 412/12 (FLOR); Santo Amaro de Imperatriz, Parque Estadual da Serra do Tabuleiro, Hotel Plaza Caldas da Imperatriz, 24 Jan. 2014, M.C. Westphalen 480/14 (SP 446238); São Paulo, Santo André, Reserva Biológica do Alto da Serra de Paranapiacaba, 5 Dec. 2013, M.C. Westphalen 474/13 (SP 446234); São Luiz do Paraitinga, P.E. da Serra do Mar, Núcleo Sta. Virgínea, 17 Apr. 2013, R.M. Pires RP2 (SP 446259).
Notes — This species was described from southern Brazil (Santa Catarina State) by Bresadola (1896) and since then registered from different tropical and subtropical regions in South and Central America (Lowe 1966, Carranza-Velásquez & Ruiz-Boyer 2005, Robledo & Rajchenberg 2007, Westphalen et al. 2012). It is characterised by its yellowish basidiomes that become reddish when bruised and sometimes after drying and the large and very irregular pores (3–5 per mm) that are often split and lacerate, forming a daedaloid surface. Microscopically it presents a dimitic hyphal system, thin to thick-walled cystidia encrusted with fine crystals and narrowly ellipsoid basidiospores (4.0–5.0 × 2.0–2.5 μm, Lm × Wm = 4.38 × 2.38, Q = 1.60–2.25, Qm = 1.86, n = 30/2).
Geesterania davidii Westphalen & Rajchenb., sp. nov. — MycoBank MB822660; Fig. 3c–d, ,4h4h
Holotype. Brazil, Rio Grande do Sul, Morrinhos do Sul, Perdida, 14 May 2012, M.C. Westphalen 396/12 (SP 446193).
Etymology. Named in honour of Alix David (Lyon, France), for her extensive work on the biology of fungi, especially her numerous investigations on mating systems and nuclear behaviour of polypores.
Basidiomes annual, resupinate, thin, soft and somewhat fleshy when fresh, becoming hard and slightly waxy when dried, margin narrow to absent, regular, white to cream-coloured; pore surface pale yellow to pale greenish yellow when fresh, becoming reddish when bruised and straw-coloured to pinkish brown after drying; pores regular, round to angular, with thin and often dentate dissepiments, sometimes partially split, 5–7(–8) per mm, tubes concolour with the pore surface, up to 2 mm deep; subiculum very thin, up to 0.2 mm thick, cream-coloured when fresh, becoming ochraceous when dried. Hyphal system dimitic; generative hyphae with clamps, branched or more rarely unbranched, hyaline and thin-walled, 2.0–4.0 μm wide; skeletal hyphae hyaline to slightly yellowish, thick-walled to almost solid, unbranched, 3.0–5.0 μm wide. Cystidia appearing as thickened skeletal hyphal ends, in the dissepiments slightly thick-walled, in the trama forming a continuous with the skeletal hyphae with very thick walls to almost solid, but usually with a wider lumen at the apex, finely encrusted, 5.0–8.0 μm diam. Cystidioles fusoid to ventricose, 9–11 × 4–4.5 μm. Basidia clavate, 4-sterigmate, 10–12 × 4–5 μm; basidiospores ellipsoid, hyaline, smooth, thin-walled, usually with one or two oil drops, 3.5–4.5 × 2.0–3.0 μm, Lm × Wm = 4.02 × 2.60, Q = 1.33–1.80, Qm = 1.55, n = 30/1.
Distribution — Known only from southern Brazil, in Rio Grande do Sul State.
Substratum — Growing on dead branches and logs of unidentified angiosperms.
Additional specimens examined. Brazil, Rio Grande do Sul, São Francisco de Paula, CPCN Pró-Mata, 14 Nov. 2009, M.C. Westphalen 289/09 (ICN 154468); ibid., 18 Apr. 2012, M.C. Westphalen 370/12 (SP 446173).
Notes — Geesterania davidii is characterised by yellowish basidiomes that become reddish when bruised and after drying, the medium to small pores and the small ellipsoid basidiospores. It differs from G. carneola in smaller and more regular pores, sometimes only partially split (Fig. 3c–d). In addition, G. carneola presents slightly longer and thinner, narrowly ellipsoid spores. Even though there are cases of overlapping in basidiospore size between the two species and only a detailed examination can reveal the differences, the values of Q and Qm show the difference in spore shape.
Flaviporus tenuis Westphalen, Rajchenb. & Tomšovský, nom. nov. — MycoBank MB822514
≡Junghuhnia minuta I. Lindblad & Ryvarden, Mycotaxon 71: 346. 1999, basionym.
Etymology. tenuis refers to the small and fragile pilei characteristic of this species.
Specimens examined. Brazil, Paraná, Piraquara, Morro do Canal, 4 Sept. 2013, M.C. Westphalen 470/13 (SP 446231); Santa Catarina, Florianópolis, UCAD, 12 Mar. 2012, M.C. Westphalen 356/12 (SP 446164); Rio Grande do Sul, São Francisco de Paula, FLONA, 30 Apr. 2012, M.C. Westphalen 386/12 (SP 446184); ibid., 15 Apr. 2013, M.C. Westphalen 427/13 (SP 446207) and 430/13 (SP 446210); ibid., CPCN Pró-Mata, 19 Apr. 2013, M.C. Westphalen 442/13 (SP 446218). –Ecuador, Orellana, Yasuni National park, Yasuni Scientific Research Station, 9–12 Mar. 2002, L. Ryvarden 44687 (O).
Notes — Due to the presence of strongly agglutinated hyphae, Junghuhnia minuta morphologically fits the concept of Flaviporus, which is supported by our molecular data (Fig. 1). Since the name Flaviporus minutus is already taken by a different species (Wu et al. 2017), we propose the new name Flaviporus tenuis. This species can easily be recognised by the very small and brittle basidiomes growing in clusters, that become resinous-hard upon drying. Microscopically, it is very similar to F. brownii, presenting hyphal pegs and thick-walled encrusted cystidia. Macroscopically, F. tenuis is similar to F. liebmannii, presenting whitish basidiomes with darkening of the pileus surface upon drying. However, F. liebmannii differs in solitary to imbricate larger basidiomes and microscopically it lacks encrusted thick-walled cystidia. For a full description see Westphalen et al. (2012; as Junghuhnia minuta).
Steccherinum meridionale (Rajchenb.) Westphalen, Tomšovský & Rajchenb., comb. nov. — MycoBank MB822515
≡Junghuhnia collabens var. meridionalis Rajchenb., Sydowia 40: 236. 1987, basionym.
≡Junghuhnia meridionalis (Rajchenb.) Rajchenb., Austral. Syst. Bot. 16 (4): 477. 2003.
Specimens examined. Argentina, Chubut, P.N. Los Alerces, Río Arrayanes y Menéndez, 8 May 1991, M. Rajchenberg 11086 (CIEFAPcc 54); Neuquén, P.N. Lanín, Lago Lácar, Cascada Chachín, 19 May 1999, M. Rajchenberg 11924 (CIEFAPcc 284). –Brazil, Paraná, Morretes, Serra da Graciosa, 13 Oct. 2009, M.A. Reck 251/09 (ICN 154709); Rio Grande do Sul, São Francisco de Paula, FLONA, 22 June 2009, M.C. Westphalen 238/09 (ICN 154290); ibid., 26 Mar. 2010, M.C. Westphalen 295/10 (ICN 154659); ibid., 26 Mar. 2010, M.C. Westphalen 303/10 (ICN 154660); Derrubadas, Parque Estadual do Turvo, 15 Sept. 2009, M.A. Reck 198/09 (ICN 154340); Santa Catarina, Santo Amaro da Imperatriz, Parque Estadual Serra do Tabuleiro, 18 Sept. 2010, M.A. Reck 559/10 (ICN 154719). –Chile, Palena, Chaitén, ruta a Caleta Gonzalo, km 20, 6 Apr. 1996, M. Rajchenberg 10466 (CIEFAPcc 55); Valdivia, Corral, Thaxter, Aug. 1905 (BAFC 31012), holotype.
Notes — Steccherinum meridionale is characterised by cinnamon to brick red resupinate basidiomes with small pores (7–9 per mm) and waxy soft consistency when fresh, becoming hard upon drying. Microscopically, it presents abundant skeletocystidia projecting above the hymenium and sub-cylindrical basidiospores (3.0–4.0 × 1.5–2.0 μm). This species was first described as a variety of Junghuhnia collabens (Rajchenberg 1987a) but later, using cultural features and intercompatibility tests, Rajchenberg (2003) verified it as an autonomous taxon. Our data shows that S. meridionale and J. collabens are not phylogenetically related (Fig. 1) and that the former groups close to the Neotropical species S. neonitidum. Junghuhnia meridionalis was described from the Patagonian Andes forests in southern Argentina and Chile and later registered from Neotropical regions (Westphalen et al. 2010, 2012). Unfortunately, the DNA amplification and sequencing of Brazilian specimens was unsuccessful, but we decided to keep the Neotropical specimens under the name S. meridionale and include it in the identification key. Further work can clarify if the Neotropical and Patagonian specimens belong to the same widely distributed species or if the Neotropical collections represent a morphologically similar but molecularly separated species. For a full description see Westphalen et al. (2012; as J. meridionalis).
Steccherinum neonitidum Westphalen & Tomšovský, sp. nov. — MycoBank MB822516; Fig. 5, ,66
Steccherinum neonitidum. — Scale bar = 2 cm. — Photo: M.A. Reck.
Steccherinum neonitidum. a. Basidiospores; b. basidia; c. cystidioles; d. generative hyphae; e. skeletal hyphae; f. skeletocystidia. — Scale bar = 10 μm. — Drawn by M.C. Westphalen.
Holotype. Brazil, Rio Grande do Sul, São Francisco de Paula, CPCN Pró-Mata, 18 Apr. 2012, M.C. Westphalen 371/12 (SP 446174).
Etymology. Neo- means new in Latin. The name refers to morphological similarity of the species to Steccherinum nitidum (Pers.) Vesterh. (syn. Junghuhnia nitida).
Basidiomes annual, resupinate, very thin, easily separable from the substrate and often detaching upon drying, soft when fresh, becoming corky and somewhat brittle when dried; margin irregular, white, narrow to wide, very thin, cottony and fimbriate, up to 5 mm wide; pore surface pinkish cream to very pale pinkish cinnamon; pores regular, round to angular, sometimes partially lacerate, 8–10 per mm; tubes up to 1.0 mm deep, concolour with the pore surface; subiculum white to cream-coloured, up to 0.5 mm thick. Hyphal system dimitic; generative hyphae clamped, thin to slightly thick-walled, hyaline, 2.0–3.5 μm diam; skeletal hyphae unbranched to branched, thick-walled to solid, hyaline to yellowish, 2.0–5.0 μm diam. Skeletocystidia, abundant, clavate, thick-walled and heavily encrusted, metachromatic, immersed in the trama or projecting above the hymenium, 6.0–10.0 μm diam. Basidia clavate, 4-sterigmate; basidiospores ovoid to ellipsoid, smooth and hyaline, 3.0–4.0 × 2.0–3.0 μm, Lm × Wm = 3.35 × 2.53 μm, Q = 1.2–1.5, Qm = 1.34, n = 40/2.
Mating system — Tetrapolar. Ten monosporic cultures (SP 446174) were confronted and the mating types distributed as follows: A1B1: 1, 2, 4; A2B2: 6, 10; A1B2: 3, 5, 8, 9; A2B1: 7.
Nuclear behaviour — Normal (monosporic culture monokaryotic, polysporic culture dikaryotic).
Distribution — Southern and south-eastern Brazil.
Substratum — Growing on dead branches of unidentified angiosperms.
Additional specimens examined. Brazil, Rio Grande do Sul, Cambará do Sul, PARNA da Serra Geral, Itaimbezinho, R.M. Silveira & R.T. Guerrero 048, 19 Nov. 1987 (ICN); ibid., Fortaleza, 29 Apr. 2012, M.C. Westphalen 380/12 (SP 446179); São Francisco de Paula, CPCN Pró-Mata, 30 May 2009, M.C. Westphalen 222/09 (ICN 154296); ibid., FLONA, 22 June 2009, M.C. Westphalen 236/09 (ICN 154294) and M.C. Westphalen 241/09 (ICN 154295); São Paulo, São Luiz do Paraitinga, P.E. da Serra do Mar, Núcleo Sta. Virgínea, 12 June 2013, R.M. Pires & C.M. Ishida RP79 (SP 445975).
Notes — Steccherinum neonitidum is characterised by pinkish, thin resupinate basidiomes with small pores and fimbriate white margins. Microscopically, it can be recognised by the small ovoid to ellipsoid basidiospores and the abundant encrusted skeletocystidia. This species has been traditionally identified in Brazil as J. nitida due to its morphological similarities. However, Steccherinum nitidum (= Junghuhnia nitida) can be distinguished by larger pores (4–7 per mm; in margins of basidiomes pores can split to 3 per mm) and the somewhat longer basidiospores (3.7–4.5 × 2.0–2.7 μm, Lm × Wm = 4.26 × 2.41 μm, Q = 1.65–1.88, Qm = 1.77, n = 187/9). A careful analysis of herbarium specimens identified as J. nitida in other Neotropical areas may greatly increase the distribution of this species and confirm if S. nitidum is restricted to temperate regions in the Northern Hemisphere, while S. neonitidum is distributed throughout Neotropical regions. Steccherinum meridionale is also morphologically similar, but can be separated by the darker, cinnamon to brick-red basidiomes and the sub-cylindrical basidiospores.
Steccherinum polycystidiferum (Rick) Westphalen, Tomšovský & Rajchenb., comb. nov. — MycoBank MB822517
≡Poria polycystidifera Rick, Iheringia, Bot. 7: 281. 1960, basionym.
≡Junghuhnia polycystidifera (Rick) Rajchenb., Nordic J. Bot. 7 (5): 566. 1987.
Specimens examined. Brazil, Rio Grande do Sul, Caxias do Sul, Terceira Légua, 10 Dec. 2012, M.C. Westphalen 419/12 (SP 446201); São Salvador, 4 Apr. 1944, J. Rick s.n. (PACA FR 18603), holotype; Porto Alegre, Morro Santana, 14 Dec. 2007, M.C. Westphalen 070/07 (ICN 154122); Santa Catarina, Blumenau, Parque Nacional da Serra do Itajaí, 12 June 2012, M.A.B. Silva et al. 253 (FLOR 49252); São Paulo, São Luiz do Paraitinga, P.E. da Serra do Mar, Núcleo Sta. Virgínea, 29 Oct. 2013, R.M. Pires 140 (SP 446271).
Notes — Steccherinum polycystidiferum is characterised by cream-coloured to beige resupinate basidiomes, becoming slightly darker upon drying, small pores (6–8 per mm) and ellipsoid to sub-cylindrical small basidiospores (3.0–4.0 × 1.5–2.0 μm). Steccherinum meridionale presents very similar micromorphology and can be distinguished only by the basidiomes of cinnamon to brick-red colour. Despite the morphological similarities, S. polycystidiferum presents a phylogenetic isolate position, but close to the S. nitidum group. This seems to be a rare species, first described from northern Argentina as Junghuhnia microspora (Rajchenberg 1983), and later synonymised with J. polycystidifera (Rajchenberg 1987b), a species originally described from southern Brazil as Poria polycysidifera (Rick 1960).
Steccherinum undigerum (Berk & M.A. Curtis) Westphalen & Tomšovský, comb. nov. — MycoBank MB822518
≡Polyporus undigerus Berk. & M.A. Curtis, Bot. J. Linn. Soc. 10: 317. 1869, basionym.
≡Junghuhnia undigera (Berk. & M.A. Curtis) Ryvarden, Mycotaxon 20 (2): 359. 1984.
= Junghuhnia kotlabae Pouzar, Czech Mycol. 55 (1–2): 2. 2003.
= Junghuhnia complicata Blumenf. & J.E. Wright, Mycotaxon 19: 472. 1984.
Mating system — Tetrapolar. Two specimens were studied, each with nine monosporic cultures confronted, and the mating types distributed as follows:
(SP 446233) A1B1: 1, 2, 5, 8; A2B2: 4, 9; A1B2: 3, 6; A2B1: 7.
(SP 446206) A1B1: 1, 2, 8; A2B2: 3; A1B2: 4, 6; A2B1: 5, 7, 9.
Nuclear behaviour — Normal (monosporic culture monokaryotic, polysporic culture dikaryotic).
Specimens examined. Argentina, Misiones, Puerto Libertad, ad culturam arbum coniferatum Alto Paraná, S. Blumenfeld, 25 Mar. 1981, ad Pinus taeda vivo (BAFC 28947), holotype of J. complicata. –Brazil, Paraná, Curitiba, Parque Barigui, 26 Jan. 2014, M.C. Westphalen 485/14 (SP 446242); Rio Grande do Sul, São Francisco de Paula, CPCN Pró-Mata, 18 Apr. 2012, M.C. Westphalen 374/12 (SP 446176); ibid., 19 Apr. 2013, M.C. Westphalen 443/13 (SP 446219); FLONA, 30 Apr. 2012, M.C. Westphalen 387/12 (SP 446185); Dom Pedro de Alcântara, Mata da Cova Funda, 14 May 2012, M.C. Westphalen 392/12 (SP 446189); Morrinhos do Sul, Perdida, 14 May 2012, M.C. Westphalen 395/12 (SP 446192); Caxias do Sul, Terceira Légua, 10 Dec. 2012, M.C. Westphalen 418/12 (SP 446200); São Paulo, Santo André, P.E. do Alto da Serra de Paranapiacaba, 5 Dec. 2013, M.C. Westphalen 472/13 (SP 446233); São Luiz do Paraitinga, PE da Serra do Mar, Núcleo Santa Virgínea, Trilha Ipiranga, 7 Nov. 2012, M.C. Westphalen 406/12 (SP 446199); ibid., 26 Apr. 2014, M.C. Westphalen & R.M. Pires 496/14 (SP 446249); ibid., 27 Apr. 2014, M.C. Westphalen & R.M. Pires 501/14 (SP 446254); P.E. Fontes do Ipiranga, 20 May 2013, M.C. Westphalen 426/13 (SP 446206); ibid., P.E. da Cantareira, 20 June 2013, M.C. Westphalen 446/13 (SP 446221). –Cuba, sine loc., C. Wright 457 (K 181077), holotype; Pinar del Rio, Soroa near San Cristóbal, 13 Jan. 1967, F. Kotlaba (PRM 878650), holotype of J. kotlabae; Pinar del Rio, Soroa, 3 Feb. 1967, F. Kotlaba (PRM 870890), paratype of J. kotlabae.
Notes — Steccherinum undigerum is characterised by pileate, thin and flexible effused-reflexed or more rarely sessile basidiomes that become corky and somewhat brittle upon drying. Even though it presents a poroid hymenophore, the pores (4–6 per mm) present dentate dissepiments and are often split and somewhat irregular. Features of the pore surface is what distinguishes this species from others in the Steccherinum clade. Microscopically, it presents typical skeletocystidia of Steccherinum, more easily observed in the dissepiments edges but also found in the trama, a dimitic hyphal system and ellipsoid to subglobose basidiospores (4–5 × 3.5–4.5 μm), larger than other species in the group. For a full description see Westphalen et al. (2012; as J. undigera). Phylogenetically, S. undigerum is the only studied species that groups very close to S. ochraceum (Fig. 1, ,2),2), type of the genus (for further comments see Discussion). Our materials present identical ITS and LSU sequences to a specimen from Cuba, type locality of this species, available at GenBank as Junghuhnia sp. 4. (Miettinen et al. 2012). Steccherinum undigerum is a very common species in southern Brazil and is often found in different areas of the Atlantic Rainforest.
Junghuhnia kotlabae is another species described from Cuba (Pouzar 2003) that presents morphological features almost identical to J. undigera and in its original description was only separated by the presence of thin-walled cystidia (absent in J. undigera). The basidiospore dimensions are 3.7–5.5 × 3–4.2 μm (Pouzar 2003) or 3.7–4.9 × 3–3.9 μm (our observation). In addition, the ITS sequence we obtained from the paratype of J. kotlabae presents only two different bases from specimens of J. undigera from Brazil. Since both the morphological and molecular similarities are high and cystidia present variation within a species, we consider J. kotlabae a synonym of J. undigera.
Additional specimens examined.Flaviporus subundatus –Brazil, Paraná, Paranaguá, Prox. a Rodovia PR 508, 2 Sept. 2013, M.C. Westphalen 457/13 (SP 446224); Curitiba, Parque Barigui, 26 Jan. 2014, M.C. Westphalen 483/14 (SP 446240); Rio Grande do Sul, São Francisco de Paula, FLONA, 16 Apr. 2012, M.C. Westphalen 367/12 (SP 446171); ibid., 30 Apr. 2012, M.C. Westphalen 381/12 (SP 446180); ibid., 15 Apr. 2013, M.C. Westphalen 445/13 (SP 446220); Santa Catarina, Itapoá, RPPN Volta Velha, 17 Nov. 2012, M.C. Westphalen 411/12 (FLOR); São Paulo, São Paulo, Parque Estadual da Cantareira, 27 June 2012, M.C. Westphalen 398/12 (SP 446195); P.E. da Serra do Mar, Núcleo Santa Virgínea, 27 Apr. 2014, R.M. Pires RP332 (SP 446276); Peruíbe, Estação Ecológica Juréia-Itatins, 25 Mar. 2014, M.C. Westphalen 494/14 (SP 446247). –Cuba, Guantánamo, Baracoa, El Yunque Mountain, Mar. 1903, L.M. Underwood & F.S. Earle 1168 (BPI 0243504), isotype. Steccherinum nitidum –Austria, Styria, Graz, Grazer Bergland, Schöckel Mountain, 31 Sept. 1975, P. Döbbeler & J. Poelt (PRM 801862). –Czech Republic, Brno, Žebětín, 30 Oct. 2015, M. Tomšovský 52/2015 (BRNM 781249); Znojmo, Vranov nad Dyjí, Braitava nature monument, 29 Aug. 2012, M. Tomšovský 33/2012 (BRNM 781248). –Germany, Rhineland, Bonn, Bad Godesberg, Alnus glutinosa, 8 Feb. 1968, H. Gorholt (PRM 690499). –Hungary, Tapolca, Lesence valley, Carpinus betulus, 15 Sept. 1978, V. Holubová (PRM 818395); Mátra Mts, Kékes Mt, Fagus sylvatica, 22 Sept. 1978, J. Klán (PRM 818365). –Slovakia, Zvolen, Hriňová, Fagus sylvatica, 28 Aug. 1982, F. Kotlaba, (PRM 828459), Hajnáčka, Pohanský hrad nature reserve, Quercus sp., 22 July 1993, J. Holec (PRM 886250), Humenné, Svetlice, Fagus sylvatica, 23 Oct. 1987, F. Kotlaba (PRM 854505). Junghuhnia chlamydospora –Belize, Stan Creek Distr., Corkscomb basin wildlife sanctuary, 16 Nov. 2001, leg. L. Ryvarden 44241 (O), holotype. Junghuhnia globospora –Venezuela, Merida Prov., Monte Zerpa by Merida 2000 m, 29 Jan. 2001, L. Ryvarden 45503 (O), holotype. Junghuhnia semisupiniformis –Mexico, Japala, 1909, Murrill (O), holotype. –Puerto Rico, Rio Grande, Big tree trail, 18 June 1996, L. Ryvarden 38921 (O 911195).
Key to Neotropical Junghuhnia s.lat. genera
1. Basidiomes becoming reddish or blackish when bruised and/or after drying, cystidia appearing as elongated finely encrusted skeletal hyphae ends and sometimes hard to differentiate from them, skeletal hyphae metachromatic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Geesterania
1. Basidiomes not changing colour when bruised, sometimes darker after drying, cystidia easily distinguished from the skeletal hyphae, smooth or covered with large crystals, skeletal hyphae non metachromatic. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
2. Basidiomes strongly shrinking when dried, resinous-hard, cystidia variable, thin to thick-walled, smooth or heavily encrusted, hyphae strongly agglutinated, hyphal system pseudodimitic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Flaviporus
2. Basidiomes with no remarkable shrinking when dried, corky to waxy, cystidia thick-walled and heavily encrusted, hyphae not agglutinated, hyphal system strictly dimitic. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Steccherinum
Key to Neotropical Junghuhnia s.lat. species
1. Basidiomes strictly resupinate. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
1. Basidiomes pileate, effused-reflexed to sessile. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9
2. Basidiomes changing colour when bruised or dried, cystidia finely encrusted, skeletal hyphae metachromatic. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
2. Basidiomes with no remarkable colour change when bruised or dried, cystidia smooth or heavily encrusted. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5
3. Basidiomes becoming blackish in parts when dried, dextrinoid chlamydospores present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .J. chlamydospora
3. Basidiomes becoming reddish when bruised and often when dried, dextrinoid chlamydospores absent. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
4. Pore surface irregular, pores 3–5 per mm, strongly split, basidiospores narrowly ellipsoid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .G. carneola
4. Pore surface regular, pores 5–8 per mm, entire or only slightly split, basidiospores ellipsoid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .G. davidii
5. Generative hyphae simple-septate, hyphae strongly agglutinated, cystidia hard to observe and sometimes only large crystals seen in the trama . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .F. subundatus
5. Generative hyphae clamped, hyphae not agglutinated, cystidia easily observed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
6. Basidiomes cinnamon to brick-red or cream to straw coloured, tough and somewhat waxy, basidiospores sub-cylindrical.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7
6. Basidiomes white to cream or pinkish, thin, papery to corky, basidiospores globose to widely-ellipsoid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .8
7. Pore surface brick-red, pale cinnamon or dull red. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . S. meridionale
7. Pore surface cream to beige. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. polycystidiferum
8. Basidiomes pale pinkish cream to pale orange-pink, basidiospores broadly ellipsoid to ovoid (3.0–4.0 × 2.0–3.0 μm), cystidia heavily encrusted. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. neonitidum
8. Basidiomes white to cream, basidiospores globose to subglobose, cystidia smooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .J. globospora
9. Basidiomes resinous-hard when dried, hyphae strongly agglutinated and hard to observe. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .10
9. Basidiomes corky to tough, hyphae not agglutinated and easily observed. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .11
10. Basidiomes growing in clusters, usually dorsally attached, cystidia clavate, heavily encrusted. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .F. tenuis
10. Basidiomes solitary, cystidia elongated with a ventricose apex, smooth. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .J. neotropica
11. Basidiomes flexible when fresh becoming papery to corky upon drying, pores 5–6 per mm, basidiospores 4.5–5.0 × 3.5–4.5 μm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .S. undigerum
11. Basidiomes tougher, somewhat dense when died, pores 8–10 per mm, basidiospores 2.5–3.5 × 2.0–3.0 μm. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .J. semisupiniformis
DISCUSSION
Our results reveal that none of the studied Neotropical species previously placed in Junghuhnia are related to the type of the genus, J. crustacea. Instead, they are placed in different clades representing three genera:
Steccherinum clade
Type. Steccherinum ochraceum.
Steccherinum meridionale, S. neonitidum, S. polycystidiferum and S. undigerum are part of this group. Even though Steccherinum is well-supported within the residual polyporoid clade (Miettinen et al. 2012, Binder et. al 2013), the Steccherinum clade is not yet resolved and more data are needed to better understand the evolutionary relations of its species. Under current knowledge, we decided to consider Steccherinum in a wide sense and to transfer the studied species into this genus, agreeing with Miettinen & Ryvarden (2016). The current wide delimitation of Steccherinum has a high phylogenetic support and can be defined morphologically by dimitic species with encrusted skeletocystidia and small subglobose to subcylindrical spores. Currently there are no morphological characters that would support the segregation of Steccherinum in different genera. However, further studies including more molecular and morphological data may elucidate if Steccherinum is a wide genus or if it encompasses several smaller genera. If the genus is further split, the Neotropical species could be transferred to Chaetoporus, a genus name available with C. nitidus (= Junghuhnia nitida) as type (Donk 1967, Miettinen et al. 2012). The only exception is S. undigerum, which groups close to S. ochraceum, type of Steccherinum.
Flaviporus
Type. Flaviporus brownii.
Our molecular data shows that Junghuhnia minuta and J. subundata group with Flaviporus, which is also supported by morphology. Species in Flaviporus are separated phylogenetically by rather long branches (Fig. 1), which may indicate that there are missing data that could clarify the relations among species of this group. Morphologically, Flaviporus species present basidiomes that become very hard and somewhat resinous after drying, cystidia with variable shape and strongly agglutinated hyphae, which support the genus as monophyletic. The hyphal system in Flaviporus is difficult to interpret due to the hyphae agglutination, especially in dried specimens. Our observations of F. tenuis and F. brownii treated in NaOH solution showed that many thick-walled hyphae present clamps, but some long aseptate segments can also be observed. In addition, there is no clear distinction between the hyphae present, so we classify the hyphal system as monomitic to pseudo-dimitic instead of strictly dimitic, as defined by Ginns (1980). Flaviporus subundatus differs by lacking clamp connections in the generative hyphae, but our molecular data support keeping this species under Flaviporus. Future studies including a wider sampling may clarify if the presence of clamp connections is variable in the genus or if F. subundatus belongs to a separate lineage. For full descriptions of F. subundatus see Lowe (1966), Ginns (1980) and Westphalen & Silveira (2012).
Geesterania
Type. Geesterania carneola.
This third group recognised in this study comprises two species previously identified as Junghuhnia carneola that belong to the phlebiod clade. The phylogenetic position of these species is supported by biological data, since J. carneola was characterised by having a bipolar mating system and an astatocoenocytic nuclear behaviour (David & Rajchenberg 1985), a combination typical of the phlebiod species, while species of Steccherinum are tetrapolar and present a normal nuclear behaviour (Rajchenberg 2011). Morphological and molecular data obtained support the description of a new genus and a new species. Geesterania is unique among polypores by the combination of basidiomes changing colour when bruised or after drying, the presence of long, finely encrusted cystidia and by the dimitic hyphal system with clamped generative hyphae and metachromatic skeletal hyphae. The two species identified in the genus are very similar but can be distinguished by the pore surface (larger, irregular and strongly split pores in G. carneola and smaller regular pores in G. davidii) and by a small difference in spore shape (Fig. 4).
The data presented in this study expand the knowledge of Junghuhnia s.lat. in the Neotropics and are in agreement with Miettinen et al. (2012), showing that the genus is highly polyphyletic and currently only J. crustacea has been confirmed as Junghuhnia s.str. During a survey of polypores in southern Brazil, seven species considered morphologically to belong to Junghuhnia s.lat. were collected and studied. The data obtained support the description of the new genus Geesterania, including one new species, as well as a new Steccherinum species and five new combinations of species previously treated in Junghuhnia. We present here full descriptions and illustrations of the new species, comments on the newly combined species and identification keys. However, more studies are required to clarify the evolutionary relationships of species previously treated as Junghuhnia, especially the ones included in the Steccherinum clade. Furthermore, the phylogenetic positions of J. semisupiniformis, J. chlamydospora, J. globospora and J. neotropica are still unknown. Here we segregated these four species by morphological groups and presented them in the identification key, but molecular data are needed to verify their proper generic position. Ryvarden (2015) also included J. sobria in his study of Neotropical Junghuhnia taxa. Junghuhnia sobria was originally characterised by the presence of gloeocystidia (Lowe 1977), a character absent in all other species of the group, therefore we consider to exclude it from Junghuhnia s.lat.
Acknowledgments
The authors would like to thank BPI, FLOR, O, PRM and S herbaria for the loan of collections and Dr. Mateus A. Reck and MSc. Dirce L. Komura for the images of fresh basidiomes. FAPESP (Fundação de Amparo a Pesquisa do Estado de São Paulo, Brazil; grant number: 2011/17219-0), European Social Fund and the state budget of the Czech Republic (Project Indicators of trees vitality Reg. No. CZ.1.07/2.3.00/20.0265) are acknowledged for financial support.
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