Chapter 2 Septal Pore Complex Morphology in the Agaricomycotina (Basidiomycota) with Emphasis on the Cantharellales and Hymenochaetales Kenneth G.A. van Driel, Bruno M. Humbel, Arie J. Verkleij, Joost Stalpers, Wally Müller & Teun Boekhout Chapter 2 ABSTRACT The ultrastructure of septa and septum-associated septal pore caps are important taxonomic markers in the Agaricomycotina (Basidiomycota, Fungi). The septal pore caps covering the typical basidiomycetous dolipore septum are distinguished into three main morphotypes: vesicular, imperforate, and perforate. Until recently, the septal pore cap-type relected the higher-order relationships within the Agaricomycotina. However, the new classiication of Fungi resulted in many changes including addition of new orders. Therefore, the septal pore cap ultrastructure of more than 350 species as reported in literature was related to this new classiication. In addition, the septal pore cap ultrastructure of Rickenella ibula and Cantharellus formosus was examined by transmission electron microscopy. Both fungi were shown to have dolipore septa associated with perforate septal pore caps. These results combined with data from the literature show that the septal pore cap type within orders of the Agaricomycotina is generally monomorphic, except for the Cantharellales and Hymenochaetales. INTRODUCTION Morphology of for example fruiting bodies (e.g. Fries, 1874; Patouillard, 1900; Fennel, 1973; Müller & Von Arx, 1973; Jülich, 1981; Berbee & Taylor, 1992), basidia (e.g. Martin, 1957; Donk, 1958; Talbot, 1973), spindle pole bodies (SPB) (e.g. McLaughlin et al., 1995; Celio et al., 2006), and septa (e.g. Moore, 1980, 1985, 1996; Khan & Kimbrough, 1982; Oberwinkler & Bandoni, 1982; Kimbrough, 1994; Wells, 1994; McLaughlin et al., 1995; Bauer et al., 1997; Müller et al., 2000b; Hibbett & Thorn, 2001) as well as physiological and biochemical characteristics (Bartnicki-Garcia, 1968; Van der Walt & Yarrow, 1984; Prillinger et al., 1993; Kurtzman & Fell, 1998; Boekhout & Guého, 2002) have strongly contributed to fungal systematics. The structural and biochemical database for fungi (Celio et al., 2006) aims to capture several of these characters in a comprehensive manner. Next to these morphological and physiological characteristics, sequence data from ribosomal DNA (i.e. nSSU and nLSU rDNA), mitochondrial DNA and protein coding genes (e.g. EF1, RPB1, RPB2) have been instrumental in fungal systematics (e.g. Swann & Taylor, 1993, 1995; Liu, 1999, 2006; Fell et al. 2000; Schüßler et al., 2001; Lutzoni et al., 2004; Tanabe et al., 2004). More recently, complete fungal genomes were used in phylogeny (phylogenomics) and revealed consistency with the molecular studies done so far (Fitzpatrick et al., 2006; Kuramae et al., 2006). Collaborations between fungal systematics (AFTOL/Deep Hyphae) have increased the resolution of the fungal tree of life that resulted in an upgraded classiication of the Fungi (James et al., 2006; Hibbett et al., 2007). 28 Septal pore complex morphology in the Agaricomycotina Since the last overview of septal ultrastructure in relation with fungal phylogeny (Fell et al., 2001; Hibbett & Thorn, 2001; Wells & Bandoni, 2001) many new orders have been proposed in the Agaricomycotina (equivalent to Hymenomycetes; Swann & Taylor, 1995) (Larsson et al., 2004; Binder et al., 2005; Hosaka et al., 2006; Hibbett et al., 2007) and the fundamental distinction between Heterobasidiomycetes and Homobasidiomycetes has disappeared. At present the Agaricomycotina contains three main clades, namely the Tremellomycetes, the Dacrymycetes, and the Agaricomycetes and 21 orders are recognized (Hibbett, 2006; Hibbett et al., 2007). In general, members of the Agaricomycotina have a dolipore septum that is lared towards the pore and may be associated with septal pore caps (SPCs) (Girbardt, 1958; Moore & McAlear, 1962; Bracker & Butler, 1963; Müller et al., 1998a, 2000b). These SPCs are distinguished into three main morphotypes: the vesicular (tubular, saccular), the imperforate (continuous) and the perforate SPC-type. The ultrastructure of the septum and septum-associated subcellular structures relected the higher-order relationships within the Agaricomycotina, and until recently, the orders herein contained only one SPC-type, either vesicular, imperforate, or perforate (e.g. Wells, 1994; Müller et al., 1998b, 2000b; Fell et al. 2001; Hibbett & Thorn, 2001; Wells & Bandoni, 2001). However, the basic changes inferred by molecular data necessitated a reconsideration of the septal ultrastructure in relation with the new classiication. Furthermore, the orders Cantharellales and Hymenochaetales both were considered having only imperforate SPCs (Hibbett & Thorn, 2001), but at present these orders probably include also members with perforate SPCs (Larsson et al., 2006; Moncalvo et al., 2006). Into the Cantharellales the Ceratobasidiales were placed, to which, Thanatephorus, Uthatobasidium and Ceratobasidium belong that all have perforate SPCs (Bracker & Butler, 1963; Lisker et al., 1975; Tu et al., 1977; Langer, 1994; Andersen, 1996; Müller et al., 1998b, 2000a; Moncalvo et al., 2006). Moreover, the position of Cantharellus itself is unclear, as it has been reported to contain perforate SPCs (Keller, 1997) as well as imperforate SPCs (Hibbett & Thorn, 2001; Larsson et al., 2004; Moncalvo et al., 2006). Hyphoderma praetermissum with perforate SPCs (Langer & Oberwinkler, 1993; Keller, 1997) is now classiied in the Hymenochaetales (Larsson et al., 2004, 2006). Finally, the omphalinoid fungi that previously were classiied in the Agaricales (Singer, 1986) revealed to be polyphyletic and a biotrophic group, including Rickenella ibula (Bull.) Raitelhuber (1973), was placed in the Hymenochaetales (Moncalvo et al., 2002; Redhead et al., 2002; Larsson et al., 2004, 2006). Here, SPC ultrastructural data from the literature was related with the recently proposed classiication of the Agaricomycotina. Moreover, the SPC ultrastructure of Cantharellus formosus and R. ibula was examined by transmission electron microscopy. It is concluded that the SPC-type within the orders of the Agaricomycotina is generally monomorphic, except for the Cantharellales and Hymenochaetales. 29 Chapter 2 MATERIALS & METHODS Strain, Media, and Culture Conditions Rickenella ibula (CBS 116393) was grown on X-agar medium (110 ml cherry extract, 600 ml pepton-glucose-saccharose, 600 ml oatmeal extract, 480 ml water, and 25 g agar; Gams et al., 1998) at room temperature. After 5 weeks a colony with a diameter of about 1 cm was used for chemical ixation and high pressure freezing. Cantharellus formosus was obtained from a commercial source. The identity of both isolates was checked by sequence analyses of the internal transcribed spacers (ITS) 1 and 2, and the D1/D2 region of the nuclear large subunit (nLSU) ribosomal DNA using standard primers, PCR and sequence conditions (White et al., 1990; Hopple & Vilgalys, 1999). Chemical Fixation Peripheral parts of the R. ibula colony of about 34 mm, and approximately 1 mm tissue blocks from the stipe and the cap of C. formosus were cut. The mycelium was chemically ixed in freshly prepared ice-cold 1% (w/v) aqueous potassium permanganate for 20 min on ice. After rinsing with ice-cold distilled water, the mycelium was dehydrated in a series of 70%, 80%, 90%, 95% and 100% (v/v) ethanol on ice. Subsequently, the ethanol was replaced by 1,2-propylene oxide (Merck KGaA, Darmstadt, Germany) (25%, 50%, 75%, and 100%) and the fungal cells were iniltrated (25%, 50%, 75%, 100%) and embedded in Spurr’s resin (Spurr, 1969), which was polymerized at 65ºC for 2 days. High-pressure Freezing and Freeze-substitution From the periphery of the R. ibula colony, pieces of about 3 mm in diameter were cut and sandwiched between aluminum planchettes (Engineering Oice M. Wohlwend GMbH, Sennwald, Switzerland), which were illed with 1-hexadecene (Müller & Moor, 1984; Studer et al., 1995) and subsequently high-pressure frozen with a Leica EM HPF (Leica Microsystems, Vienna, Austria) according to the supplier’s manual. After freezing the sandwich, it was put into liquid nitrogen and the two aluminum planchettes were separated. The excess of 1-hexadecene was removed by gently scratching the surface of the hyphae with a ine needle in liquid nitrogen (Müller et al., 2002). The fungal cells with the supporting planchette were transferred in liquid nitrogen to a CS auto freezesubstitution apparatus (Reichert-Jung, Vienna, Austria). In the substitution chamber the frozen fungal cells were rapidly put into the freeze-substitution luid, containing 1% OsO4, 3% glutaraldehyde (EM grade, Polysciences Inc, Warrington, PA, USA), and 0.3% uranylacetate (Merck) in anhydrous methanol (Merck) (modiied from Müller et al., 1980). Fungal cells were freeze-substituted for 4.5 days at -85ºC, after which the temperature was gradually raised (3ºC per hr) to 0ºC. Vials containing the freezesubstituted fungal cells were put on ice. After 1 hr the fungal cells were rinsed with anhydrous methanol, followed by anhydrous acetone. After rinsing, they were iniltrated 30 Septal pore complex morphology in the Agaricomycotina and embedded in Spurr’s resin, and polymerized as described above. Transmission Electron Microscopy Sections of 90 nm and 300 nm were post-contrasted with 4% (w/v) aqueous uranylacetate (Merck) and 0.4% (w/v) aqueous lead citrate (Merck) (Venable & Coggeshall, 1965) and viewed in a TECNAI 10 transmission electron microscope (FEI Company, Eindhoven, The Netherlands) at an acceleration voltage of 100 kV. RESULTS & DISCUSSION Septal Pore Cap Ultrastructure of Rickenella ibula and Cantharellus formosus Rickenella ibula is a small gilled mushroom commonly found between moss (Bas et al., 1995) and strongly suspected to be biotrophic (Redhead, 1981; Kost, 1984). It was previously classiied in the family Tricholomatacea within the order Agaricales (Singer, 1986). Sections of chemically ixed R. ibula hyphal cells revealed a dolipore septum associated with perforate septal pore caps (SPCs) (Figure 1A), which corresponds with previous observations in R. aulacomniophila (= R. ibula; Kost, 1984). SPCs had a width of about 300 to 400 nm, a height of about 180 nm, and small perforations of about 50 to 60 nm in diameter. The SPCs of R. ibula were comparable to those observed in Oxyporus latemarginatus (cited as Poria latemarginata; Setlif et al., 1972). The base of the SPC was connected with endoplasmic reticulum (ER) (Figure 1A), supporting previous views that the SPC is a subdomain of the ER (Girbardt, 1961; Bracker & Butler, 1963; Müller et al., 1995, 1998a; Chapter 5.). Sections of high-pressure frozen (HPF) and freeze-substituted hyphal cells of R. ibula conirmed the presence of perforate SPCs at the dolipore septum (Figure 1B). In these hyphal cells the SPC had a width of about 320 to 400 nm at its base, a height of about 200 nm, and perforations of about 50 to 60 nm. In some cells perforations of about 80 nm were found. Cryo-ixation by HPF conirmed the results obtained by chemical ixation, but gave a more detailed view of the SPC membranes and plug morphology. The SPC existed of an inner and an outer membrane enclosing the SPC matrix with an electron-dense layer in the centre (result not shown). Filamentous structures connected the inside of the SPC with the pore-occluding material as was reported previously in Schizophyllum commune (Müller et al., 1998a) and Rhizoctonia solani (Müller et al., 2000a; Van Driel et al., 2007). Sections of chemically ixed mycelium of Cantharellus formosus revealed a dolipore septum associated with perforate SPCs (Figure 2). Tissue from both stipe and hymenophore were analyzed. Stipe tissue revealed few dolipore septa and SPCs were often degenerated, while tissue from the hymenophore gave intact SPCs. Sections 31 Chapter 2 showed that the SPCs were about 630 to 810 nm in diameter with perforations of about 100 to 200 nm (Figure 2). SPCs of C. formosus were comparable to SPCs observed in Ceratobasidium cornigerum (Müller et al., 1998b). ER membrane covering the SPC and forming an outercap region was observed (Figure 2) as previously reported in other fungi (Thielke, 1972; Gull, 1976; Craig et al., 1977; Van der Valk & Marchant, 1978; Desole, 1982). Figure 1 – Transmission electron micrographs of the dolipore-septal pore cap (SPC) complex in Rickenella ibula after chemical ixation (A) and after high-pressure freezing and freeze substitution (B). The dolipore (DP) septum is covered with perforate SPCs. The SPCs in Figure B are near median cut and tangentially cut, the latter showing the surface view. Bars represent 200 nm. Figure 2 – Transmission electron micrographs of the dolipore-septal pore cap (SPC) complex in chemically ixed hyphae of Cantharellus formosus. The dolipore (DP) is covered with SPCs. Arrows indicate the membrane that forms an outer cap region above the SPC, which may be endoplasmic reticulum. Figure B is a magniication of Figure A. Bars represent 250 nm. 32 Septal pore complex morphology in the Agaricomycotina Septal Pore Cap Morphology in the Agaricomycotina According to the current classiication, the Agaricomycotina contains three classes (Tremellomycetes, Dacrymycetes, and Agaricomycetes) and 21 orders (Hibbett, 2006; Hibbett et al., 2007). The SPC ultrastructure of more than 350 species has been published (Appendix, page 42). Table 1 shows a summary of the Appendix by giving the SPC-type per order. The current use of species names was checked in Mycobank (www.mycobank. org; Crous et al., 2004). In the Tremellomycetes, the SPC is absent (Cystoilobasidiales) or has the vesicular morphology (Filobasidiales, Trichosporonales, Tremellales) (Table 1). The Dacrymycetes (Dacrymycetales) contains only species with imperforate SPCs (Table 1). The previously recognized clades that now belong to the Agaricomycetes contained either the imperforate SPC-type (Tulasnellales, Auriculariales, Hymenochaetoid, and Cantharelloid clade) or the perforate SPC-type (Polyporoid, Euagarics, Bolete, Thelephoroid, and Russuloid clade) (Hibbett & Thorn, 2001; Wells & Bandoni, 2001), with the exception of the gomphoid-phalloid clade that contained both perforate and imperforate SPCs Class Subclass Order SPC-type Tremellomycetes Cystoilobasidiales Tremellales Trichosporonales Filobasidiales absent absent or vesicular absent or vesicular absent or vesicular Dacrymycetes Dacrymycetales imperforate Agaricomycetes Sebacinales Cantharellales Auriculariales Geastrales Hysterangiales Phallales Gomphales Trechisporales Hymenochaetales Thelephorales Polyporales Gloeophylalles Corticiales Russulales Agaricales Boletales Atheliales imperforate perforate and imperforate imperforate imperforate * unknown perforate ** imperforate * imperforate * imperforate and perforate perforate perforate perforate ** perforate perforate perforate perforate perforate Phallomycetidae Phallomycetidae Phallomycetidae Phallomycetidae Agaricomycetidae Agaricomycetidae Agaricomycetidae Table 1 – SPC-type per order level in the Agaricomycotina (summary of the Appendix). The SPC-type in Hysterangiales is unknown as no SPC ultrastructure has been published. * SPC-type determined in one species. ** SPC-type determined in two species. 33 Chapter 2 (Hibbett & Thorn, 2001). However, the SPC-type of the latter clade was unclear, as only few taxa were included. Present classiication combined with SPC morphology data shows that the orders in the Agaricomycetes have in general only one SPC-type. The imperforate SPC-type is found in the Geastrales, Gomphales, Trechisporales, Auriculariales, and Sebacinales (Table 1). The perforate SPC-type is found in the Agaricales, Atheliales, Boletales, Phallales, Corticiales, Gloeophyllales, Polyporales, Russulales, and Thelephorales (Table 1). However, both perforate and imperforate SPCs are found in the Cantharellales and Hymenochaetales (Table 1). The SPC-type for members of the Hysterangiales has not been determined yet. Furthermore, the SPC-type in the Trechisporales, Geastrales, and Gomphales was examined only in one species, whereas the SPC-type in Gloeophyllales and Phallales was examined in two species. For these orders, more data on the SPC ultrastructure are required to allow reliable statements concerning their SPC-type. An overview of the SPC-type in relation with the current tree topology of the Agaricomycotina (Hibbett, 2006) is given in Figure 3. The descriptions of the SPC-type of Typhula uncialis, Bolbitius vitellinus, Plicatura nivea, Basidiodendron rimulentum, Phanerochaete sordida, Tremella encephala, Trechispora subsphaerospora, Hydnocristella himantia (Keller, 1997), Auricularia polytricha, A. mesenterica (Patton & Marchant, 1978), and Coltricia perennis (Moore, 1980) were not included in this study as either the images were of suboptimal quality and could be interpreted diferently, or the material was misidentiied. Furthermore, few irregularities on the SPC-type were found in the Agaricales (i.e. Lepista glaucocana, Mycena galopus, and Radulomyces conluens), the Russulales (i.e. Scytinostromella olivaceoalba), and the Tremellales (Ditangiibulae dikaryotae) suggesting that the SPC-type in these orders is not monomorphic (Appendix). However, as misidentiications were made in the past, these anomalies should be conirmed or supported by genetic data (e.g. ITS or nLSU sequence data) and high-quality images of the dolipore-SPC complex, for example, obtained after high-pressure freezing and freeze-substitution. However, a recent study of the SPC ultrastructure in two species of Mycena, showed perforate SPCs in M. hiemalis, while M. galopus has imperforate SPCs (Rexer & Stepanova, 2004). A reversal from perforate to imperforate SPC-type could have taken place in this genus, which would suggest that perforate SPCs might not be morphologically stable. Nevertheless, this is the only reported anomaly within a genus so far. In addition, the authors suggested that Mycena is heterogeneous (Rexer & Stepanova, 2004). Septal Pore Cap Morphology in the Hymenochaetales The Hymenochaetales order has six clades: the Oxyporus, Rickenella, Kneiiella, Hyphodontia, Coltricia, and Hymenochaetaceae clades (Larsson et al., 2006). The SPC ultrastructure is known for many of its members. Imperforate SPCs have been found in Inonotus, Hymenochaete, Hydnochaete, Phellinus, Onnia, Asterodon, Schizopora, Hyphodontia, Coltriciella, Coltricia, and Trichaptum (Appendix). Perforate SPCs were found in the Rickenella clade, i.e. 34 Septal pore complex morphology in the Agaricomycotina perforate SPC imperforate SPC vesicular SPC endoplasmic reticulum Figure 3 – Schematic phylogenetic diagram of the Agaricomycotina adopted from Hibbett (2006). In the Tremellomycetes septal pore caps (SPCs) are absent (Cystoilobasidiales) or have the vesicular morphology (Filobasidiales, Tremellales). In the Dacrymycetes (Dacrymycetales) dolipore septa are associated with imperforate SPCs. In the Agaricomycetes dolipore septa are covered either with imperforate SPCs (Auriculariales, Sebacinales, Gomphales, Trechisporales, and Geastrales) or perforate SPCs (Phallales, Corticiales, Gloeophyllales, Polyporales, Thelephorales, Russulales, Boletales, Atheliales, and Agaricales). Both imperforate and perforate SPCs occur in the Cantharellales and Hymenochaetales. The SPC-type in the Hysterangiales is unknown as no SPC ultrastructure was published. The ER-like strands covering the dolipore in the Cystoilobasidiales seem ancestral to the vesiculate and imperforate SPC-type. It appears that the perforate SPC-type has arisen several times in the Agaricomycetes. Eventually, the perforate SPC was lost in the Cantharellales and Hymenochaetales (grey boxes) and reversed to the imperforate SPC-type. 35 Chapter 2 R. ibula (Figure 1) and Hyphoderma praetermissum (Langer & Oberwinkler, 1993; Keller, 1997). Furthermore, the perforate SPC-type occurs in the Oxyporus clade as Oxyporus latemarginatus (cited as Poria latemarginata) has dolipore septa associated with perforate SPCs (Setlif et al., 1972). Thus the basal clades, viz. the Rickenella and the Oxyporus clade in the Hymenochaetales have perforate SPCs, whereas all the other clades have imperforate SPCs. This suggests that after the perforate SPC-type appeared in the Rickenella clade and the Oxyporus clade, it was subsequently lost and reversed into the imperforate type in the other clades. Septal Pore Cap Morphology in the Cantharellales The Cantharellales order consists of four clades: a core cantharelloid clade (including Cantharellus, Craterellus, Hydnum, Sistotrema, Clavulina, Multiclavula, and Membranomyces), the Botryobasidium clade, the Ceratobasidiales clade (including Ceratobasidium, Thanatephorus, and Uthatobasidium) and the Tulasnella clade (Moncalvo et al., 2006). The literature on the septal pore morphology in Cantharellus is confusing. Cantharellus cinereus was reported having dolipore septa associated with perforate SPCs (Keller, 1997). On the other hand, others interpreted Cantharellus having imperforate SPCs based on this publication (Hibbett & Thorn, 2001; Larsson et al., 2004; Moncalvo et al., 2006). Our examination of the SPC of C. formosus showed dolipore septa covered with perforate SPCs (Figure 2) and conirmed Keller’s interpretation (Keller, 1997). Next to Cantharellus, Sistotrema brinkmannii also has dolipore septa with perforate SPCs (Dong et al., 1981; Langer, 1994), and thus, members of the core cantharelloid clade have perforate SPCs. The Botryobasidium clade, which is sister to the core cantharelloid clade, has been studied extensively with respect to its SPC ultrastructure (Appendix). It has dolipore septa with imperforate SPCs. Interestingly, the Ceratobasidiales, which is the sister group of the core cantharelloid clade and the Botryobasidium clade, all do have perforate SPCs (Appendix). Finally, members of the Tulasnella clade have dolipore septa that are covered with imperforate SPCs (Appendix). The exact position of Tulasnella remained unclear, but it may be in basal position within the Cantharellales (Moncalvo et al., 2006). After the perforate SPC-type appeared in the Cantharellales it disappeared in the Botryobasidium clade and reversed to the imperforate SPC-type, which is schematically drawn in Figure 4. Trends in the Evolution of Septal Pore Cap Morphology in the Agaricomycotina As the position of certain orders is uncertain, the fungal phylogeny is not inal yet (Hibbett et al., 2007) and future phylogenetic studies may involve changes in the current tree topology. Furthermore, the SPC ultrastructure in certain orders (Geastrales, Gloeophyllales, Gomphales, Phallales, and Trechisporales) has been studied only in few species, and thus, these studies should be extended to get a better-supported SPC-type in these orders. Therefore, we cannot be conclusive on the SPC morphology evolution in 36 Septal pore complex morphology in the Agaricomycotina Figure 4 – Simpliied phylogenetic diagram of the Cantharellales showing the four main clades according to Moncalvo et al. (2006). The core cantharelloid clade and Ceratobasidiales both have dolipore septa associated with perforate septal pore caps (SPCs), whereas the Botryobasidium and Tulasnella clades both have imperforate SPCs. Probably the perforate SPC-type has been lost in the Botryobasidium clade and reversed to the imperforate SPC-type. the Agaricomycotina. Still, certain trends can be inferred from the SPC morphology data combined with the current classiication. As the basal lineage in the Agaricomycotina has dolipore septa without SPCs (Cystoilobasidiales) but covered with ER-like strands (e.g. Itersonilia perplexans; Boekhout, 1991), we assume this might be ancestral to both the vesicular and imperforate SPC-types. Evidence showing that the vesicular SPCtype resembles the ER membrane when stained with zinc-iodine (Müller et al., 1995, 1998a) may support this view of a close relation between ER and the vesicular SPC-type. Eventually, the imperforate has given rise to the perforate SPC-type in the Agaricomycetes, which might have reversed to the imperforate SPC-type (Figure 3). This view difers from the one stated by Moore (1996), who suggested a SPC phylogeny that would progress from imperforate to perforate to vesicular forms. The Cantharellales and Hymenochaetales both have imperforate and perforate SPCs. After perforate SPCs have appeared, they subsequently were lost and reversed to the imperforate SPC-type. However, the presented phylogeny of the Cantharellales is probably not inal, as, for example, the position of the Tulasnella clade is still not clear (Moncalvo et al., 2006). Moreover the classiication of most orders in the Agaricomycetes is still considered uncertain (incertae sedis; Hibbett et al., 2007). Future phylogenetic studies together with ultrastructural studies of the septal pore complex morphology may shed a more deinitive light on SPC morphology evolution. 37 Chapter 2 REFERENCES Andersen, T.F. (1996) A comparative taxonomic study of Rhizoctonia sensu lato employing morphological, ultrastructural and molecular methods. Mycol. 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Academic Press Inc., San Diego, USA. 41 Species cited as Agaricus bisporus Agaricus campestris Agaricus silvicola Agaricus xanthoderma Agrocybe arvalis Agrocybe cylindracea Agrocybe dura Agrocybe praecox Amanita muscaria Amanita rubescens Amanita strobiliformis Armillaria mellea Calocybe chrysenteron Clitocybe martiorum Clitocybula lacerata Coprinopsis cinerea Coprinopsis lagopus Coprinopsis radiata Coprinopsis stercorea Coprinus comatus Cortinarius odorifer Cortinarius orellanus Cortinarius trivialis Cortinarius xanthophyllus Crepidotus amygdalosporus Agaricus essettei Agrocybe aegerita Coprinus cinereus Coprinus lagopus Coprinus radiatus Coprinus stercorarius Order SPC-type Author Agaricales perforate Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales perforate perforate perforate perforate perforate perforate perforate perforate perforate perforate perforate perforate perforate perforate perforate Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales perforate perforate perforate perforate perforate perforate perforate perforate perforate Craig et al., 1977; Patton & Marchant, 1978a; Thielke, 1972 Manocha, 1965 Keller, 1997 Keller, 1997 Keller, 1997 Keller, 1997 Keller, 1997 Gull, 1976 Flegler et al., 1976; Patton & Marchant, 1978a Müller et al., 1998b (reference herein) Keller, 1997 Berliner & Duf, 1965; Moore, 1965 Keller, 1997 Keller, 1997 Keller, 1997 McLaughlin, 1974; Moore et al., 1979; Van der Valk & Marchant, 1978 Giesy & Day, 1965; Waters et al., 1975 Desole, 1982 Ellis et al., 1972 Oberwinkler, 1985 Keller, 1997 Keller, 1997 Keller, 1997 Keller, 1997 Keller, 1997 Chapter 2 42 Appendix Chapter 2 - Septal pore cap type in the Agaricomycotina Appendix Chapter 2 - continued Species Collybia velutipes Galera paludosa Collybia peronata Calvatia gigantea Limacella glioderma Order SPC-type Author Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales perforate perforate perforate perforate perforate perforate perforate perforate perforate perforate perforate perforate perforate imperforate perforate perforate perforate perforate perforate perforate perforate perforate perforate imperforate perforate perforate perforate Boekhout et al., 1989 Patrignani & Pellegrini, 1986 Foerster et al., 1965 Besson & Froment, 1968 Keller, 1997 Keller, 1997 Keller, 1997 Keller, 1997 Keller, 1997 Beneke, 1963 Rexer & Stepanova, 2004 (reference herein) Keller, 1997 Keller, 1997 Keller, 1997 Keller, 1997 Flegler et al., 1976 Keller, 1997 Keller, 1997 Keller, 1997 Keller, 1997 Keller, 1997 Keller, 1997 Keller, 1997 Rexer & Stepanova, 2004 Rexer & Stepanova, 2004 Keller, 1997 Khan & Kimbrough, 1979 43 Appendix Disporotrichum dimorphosporum Fistulina hepatica Flammulina velutipes Galerina paludosa Gymnopilus sapineus Gymnopus peronatus Hygrophorus karstenii Laccaria amethystina Lachnella alboviolascens Langermannia gigantea Lentinula edodes Lepiota grangei Lepista glaucocana Lepista luscina Limacella delicata var. glioderma Lycoperdon perlatum Lyophyllum favrei Lyophyllum ulmarium Macrocystidia cucumis Melanoleuca subalpina Melanoleuca subpulverulenta Melanoleuca verrucipes Mucronella calva Mycena galopus Mycena hiemalis Mycena pseudocorticola Nematoloma puiggarii cited as Species Nidularia conluens Omphalotus olearius Panellus stipticus Phaeolepiota aurea Phaeomarasmius erinaceus Pholiota terrestris Pleurotus cystidiosus Pluteus salicinus Psilocybe cubensis Psilocybe mexicana Radulomyces conluens Resupinatus applicatus Rhodocybe popinalis Schizophyllum commune Strobilurus esculentus Stropharia aeruginosa Stropharia rugosoannulata Tephrocybe anthracophila Tephrocybe boudieri Tephrocybe coracina Volvariella bombycina Xeromphalina cornui Xerula caussei cited as Clitocybe olearia Rhodocybe mundula Lyophyllum anthracophilum, Lyophyllum spaerosporum Lyophyllum boudieri Lyophyllum coracinum Order SPC-type Author Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales Agaricales perforate perforate perforate perforate perforate perforate perforate perforate perforate perforate imperforate perforate perforate perforate Patton & Marchant, 1978a Patrignani & Pellegrini, 1986 Lingle, 1989 Keller, 1997 Keller, 1997 Wells, 1978 Moore, 1977; Moore & Patton, 1975 Keller, 1997 Tu & Kimbrough, 1978 Flegler et al., 1976 Keller, 1997 Keller, 1997 Clemencon, 2004 Jersild et al., 1967; Marchant & Wessels, 1973, 1974; Moore & Patton, 1975; Müller et al., 1994, 1995, 1998a, 1999, 2000c; Patton & Marchant, 1978a; Raudaskoski, 1972; Van der Valk & Marchant, 1978; Wells, 1965 Agaricales Agaricales Agaricales Agaricales perforate perforate perforate perforate Keller, 1997 Keller, 1997 Thielke, 1972 Keller, 1997 Agaricales Agaricales Agaricales Agaricales Agaricales perforate perforate perforate perforate perforate Keller, 1997 Keller, 1997 Flegler et al., 1976 Keller, 1997 Keller, 1997 Chapter 2 44 Appendix Chapter 2 - continued Appendix Chapter 2 - continued cited as Order SPC-type Author Athelia rolfsii Athelopsis glaucina Cristinia helvetica Leptosporomyces mutabilis Piloderma bicolor Sclerotium rolfsii Atheliales Atheliales Atheliales Atheliales Atheliales perforate perforate perforate perforate perforate Tu et al., 1977 Keller, 1997 Keller, 1997 Keller, 1997 Keller, 1997 Auriculariales Auriculariales imperforate imperforate Wells, 1994 Lü & McLaughlin, 1991; Moore, 1978b; Oberwinkler, 1985; Tu & Kimbrough, 1978; Wells, 1994; Wells & Bandoni, 2001 Auricularia fuscosuccinea Auricularia mesenterica Basidiodendron cinereum Basidiodendron eyrei Basidiodendron sp. Ductifera sp. Eichleriella sp. Exidia candida Exidia glandulosa Auriculariales Auriculariales Auriculariales Auriculariales Auriculariales Auriculariales Auriculariales Auriculariales Auriculariales imperforate imperforate imperforate imperforate imperforate imperforate imperforate imperforate imperforate Exidia nucleata Exidia sp. Exidia thurentiana Exidia truncata Exidiopsis calcea Auriculariales Auriculariales Auriculariales Auriculariales Auriculariales imperforate imperforate imperforate imperforate imperforate Auriculariales imperforate McLAughlin, 1980; Wells, 1994 Keller, 1997 Wells, 1994 Khan & Kimbrough, 1980 Oberwinkler, 1985 Oberwinkler, 1985 Oberwinkler, 1985 Wells, 1994 Keller, 1997; Moore, 1978b; Patton & Marchant, 1978a; Wells, 1994 Wells, 1964 Oberwinkler, 1985 Keller, 1997 Patton & Marchant, 1978a Andersen, 1996; Wells, 1994; Williams & Thilo, 1989 Keller, 1997 Aporpium caryae Auricularia auricula-judae Exidiopsis efusa Fibulomyces mutabilis Piloderma croceum Elmerina caryae Hirneola auricula-judae, Auricularia auricula Sebacina calcea 45 Appendix Species Species Exidiopsis sp. Exidiopsis sublivida Exidiopsis umbrina Helicomyxa everhartioides Myxarium sp. Patouillardina cinerea Protodontia oligacantha Pseudohydnum gelatinosum Stypella dubia Stypella sp. Stypella vermiformis Tremiscus helvelloides Aureoboletus gentilis Boletus cramesinus Boletus edulis Chalciporus rubinellus Coniophora fusispora Coniophora puteana Leucogyrophana mollusca Pisolithus arhizus cited as Sebacina umbrina Heterochaetella dubia Guepinia rufa Pulveroboletus gentilis Boletus rubinellus Coniophora cerebella Pisolithus tinctorius Serpula lacrymans Xerocomus chrysenteron Aphelaria tuberosa Botryobasidium candicans Tremellodendropsis tuberosa Order SPC-type Author Auriculariales Auriculariales Auriculariales Auriculariales Auriculariales Auriculariales Auriculariales Auriculariales Auriculariales Auriculariales Auriculariales Auriculariales imperforate imperforate imperforate imperforate imperforate imperforate imperforate imperforate imperforate imperforate imperforate imperforate Oberwinkler, 1985 Khan & Kimbrough, 1980 Andersen, 1996; Williams & Thilo, 1989 Kirschner & Chen, 2004 Oberwinkler, 1985 Wells, 1994 Wells, 1994 Keller, 1997; Moore, 1996; Wells, 1994 Wells, 1994 Oberwinkler, 1985 Keller, 1997 Patrignani & Pellegrini, 1986 Boletales Boletales Boletales Boletales Boletales Boletales Boletales Boletales perforate perforate perforate perforate perforate perforate perforate perforate Boletales Boletales perforate perforate Keller, 1997 Patrignani & Pellegrini, 1986 Patton & Marchant, 1978a Becket et al., 1974 Keller, 1997 Langvad, 1971 Keller, 1997 Orlovich & Ashford, 1994; Shepherd et al., 1993 Keller, 1997 Hofmann, 1989 Cantharellales Cantharellales imperforate imperforate Wells, 1994 Langer, 1994 Chapter 2 46 Appendix Chapter 2 - continued Appendix Chapter 2 - continued Species Botryobasidium conspersum Botryobasidium curtisii Botryobasidium grandisporum Botryobasidium lacinisporum Botryobasidium laeve Botryobasidium longisporum Botryobasidium obtusisporum Botryobasidium pruinatum Botryobasidium simile Botryobasidium subcoronatum Botryobasidium vagum Botryohypochnus isabellinus Cantharellus cinereus Cantharellus formosus Ceratobasidium anceps Ceratobasidium calosporum Ceratobasidium cornigerum cited as Botryobasidium botryosum Botryobasidium isabellinum Ceratobasidium ramicola SPC-type Author Cantharellales Cantharellales Cantharellales Cantharellales Cantharellales Cantharellales Cantharellales Cantharellales Cantharellales Cantharellales Cantharellales Cantharellales Cantharellales Cantharellales Cantharellales Cantharellales Cantharellales imperforate imperforate imperforate imperforate imperforate imperforate imperforate imperforate imperforate imperforate imperforate imperforate perforate perforate perforate imperforate perforate Langer, 1994 Langer, 1994 Langer, 1994 Langer, 1994 Langer, 1994 Langer, 1994 Langer, 1994 Keller, 1997 Langer, 1994 Langer, 1994; Oberwinkler, 1985 Keller, 1997; Langer, 1994 Langer, 1994 Keller, 1997 Van Driel et al., Chapter 2 Figure 2 Andersen, 1996 Weiss & Oberwinkler, 2001 Andersen, 1996; Currah & Sherburne, 1992; Keller, 1997; Müller et al., 1998b, 2000c; Patton & Marchant, 1978a; Tu et al., 1977; Wells, 1994; Wells & Bandoni, 2001 Cantharellales Cantharellales Cantharellales Cantharellales Cantharellales Cantharellales perforate perforate perforate perforate perforate perforate / imperforate Andersen, 1996; Currah & Sherburne, 1992 Keller, 1997 Weiss et al., 2004 Andersen, 1996 Andersen, 1996 Keller, 1997 (perforate); Oberwinkler, 1985 (imperforate) 47 Appendix Ceratobasidium obscurum Ceratobasidium pseudocornigerum Ceratobasidium sp. Ceratorhiza cerealis Rhizoctonia cerealis Ceratorhiza fragariae Rhizoctonia fragariae Clavulicium macounii Order Species Epulorhiza anaticula Heteroacanthella acanthophysa Heteroacanthella variabilis Moniliopsis anomala Monosporonella termitophila Paullicorticium pearsonii Rhizoctonia endophytica Rhizoctonia oryzae Rhizoctonia praticola Rhizoctonia ramicola Rhizoctonia repens Rhizoctonia solani Sistotrema brinkmannii Sistotrema sp. Stilbotulasnella conidiophora Thanatephorus cucumeris Thanatephorus pennatus Toispora biapiculata Toispora repetospora Tulasnella araneosa Tulasnella calospora cited as Order SPC-type Author Rhizoctonia anaticula Cantharellales imperforate Cantharellales Cantharellales Cantharellales Cantharellales Cantharellales Cantharellales Cantharellales Cantharellales Cantharellales Cantharellales Cantharellales imperforate imperforate perforate imperforate imperforate perforate perforate perforate perforate imperforate perforate Andersen, 1996; Currah & Sherburne, 1992; Müller et al., 1998b, 2000c Wells, 1994 (reference herein) Wells, 1994 (reference herein) Currah & Sherburne, 1992 Wells, 1994 (reference herein) Oberwinkler, 1985 Andersen, 1996 Andersen, 1996 Andersen, 1996 Andersen, 1996 Andersen, 1996; Currah & Sherburne, 1992 Andersen, 1996; Bracker & Butler, 1963, 1964; Lisker et al., 1975; Müller et al., 1998b, 2000a; Setlif et al., 1972 Cantharellales Cantharellales Cantharellales Cantharellales perforate perforate imperforate perforate Dong et al., 1981; Langer, 1994 Currah & Sherburne, 1992 Wells, 1994 (references herein) Andersen, 1996; Bracker & Butler, 1963; Langer, 1994; Müller et al., 1998b, 2000c; Tu et al., 1977 Cantharellales Cantharellales Cantharellales Cantharellales Cantharellales perforate perforate perforate imperforate imperforate Andersen, 1996; Currah & Sherburne, 1992 Langer, 1994 Langer, 1994 Wells, 1994 Andersen, 1996 Heteroacanthella variabile Epulorhiza repens Rhizoctonia dichotoma Aquathanatephorus pendulus, Thanatephorus praticola, Thanathephorus sasakii, Pellicularia ilamentosa Chapter 2 48 Appendix Chapter 2 - continued Appendix Chapter 2 - continued Species cited as Order SPC-type Author Tulasnella fuscoviolacea Tulasnella irregularis Tulasnella sp. Cantharellales Cantharellales Cantharellales imperforate imperforate imperforate Tulasnella violacea Tulasnella violea Uthatobasidium fusisporum Uthatobasidium sp. Cantharellales Cantharellales Cantharellales Cantharellales imperforate imperforate perforate perforate Moore, 1978b Andersen, 1996 Langer, 1994; Weiss et al., 2004; Wells & Bandoni, 2001 Keller & Job, 1992 Keller & Job, 1992 Keller & Job, 1992; Langer, 1994; Tu et al., 1977 Oberwinkler, 1985 Corticiales Corticiales Corticiales Corticiales Corticiales Corticiales Corticiales Corticiales Corticiales ? Corticiales ? perforate perforate perforate perforate perforate perforate perforate perforate perforate perforate Laeticorticium lundellii Laetocorticium roseum Cystoilobasidium capitatum Cystoilobasidium ferigula Cystoilobasidium inirmominiatum Itersonilia perplexans Mrakia frigida Trichosporon pullulans Leucosporidium lari-marini Corticium fuciforme Galzinia culmigena Keller, 1997 Keller, 1997 Hoch & Howard, 1981 Patton & Marchant, 1978a Wells, 1994 Keller, 1997 Keller, 1997 Diederich et al., 2003 Andersen, 1996 Andersen, 1996; Müller et al., 1998b; Tu et al., 1977 Cystoilobasidiales absent Cystoilobasidiales absent Cystoilobasidiales absent Suh & Sugiyama, 1993 Weiss et al., 2004 Suh et al., 1993 Cystoilobasidiales absent Cystoilobasidiales absent Cystoilobasidiales absent Boekhout, 1991; Fell et al., 2001 Suh et al., 1993 Guého et al., 1992 (inlated non-perf. septum) Appendix 49 Corticium boreoroseum Corticium roseum Laetisaria arvalis Laetisaria fuciformis Limonomyces culmigenus Lindtneria lava Lindtneria trachyspora Marchandiomyces corallinus Rhizoctonia zeae Waitea circinata Species cited as Order SPC-type Author Calocera cornea Dacrymycetales imperforate Calocera viscosa Cerinomyces aculeatus Cerinomyces altaicus Dacrymyces abietinus Dacrymyces chrysocomus Dacrymyces dendrocalami Dacrymyces minor Dacrymycetales Dacrymycetales Dacrymycetales Dacrymycetales Dacrymycetales Dacrymycetales Dacrymycetales imperforate imperforate imperforate imperforate imperforate imperforate imperforate Keller 1992; Tu & Kimbrough, 1978; Wells, 1994 Keller 1992; Patton & Marchant, 1978a Wells, 1994 (reference herein) Wells, 1994 Tu & Kimbrough, 1978 Wells, 1994 Wells, 1994 (reference herein) Moore, 1965 Dacrymycetales imperforate Flegler et al., 1976; Keller & Job, 1992; Moore, 1978b; Mossebo & Amougou, 2001; Wells, 1994. Dacrymycetales Dacrymycetales imperforate imperforate Wells, 1994 (reference herein) Keller & Job, 1992 Guepiniopsis chrysocoma Dacrymyces stillatus Dacrymyces deliquescens var. minor Dacrymyces deliquescens Ditiola haasii Ditiola peziziformis Femsjonia peziziformis Entorrhiza casparyana Filobasidium capsuligenum Filobasidium loriforme Leucosporidium capsuligenum Entorrhizomycetes absent Bauer et al., 1997; Deml & Oberwinkler, 1981; Weiss et al. 2004. Filobasidiales Filobasidiales vesicular absent or vesicular absent or vesicular Moore & Kreger-Van Rij, 1972 Moore & Kreger-Van Rij, 1972 (SPC absent, ER-vesicles); Wells, 1994 (sacculate) Wells, 1994 (sacculate, poorly deined; reference herein) imperforate Hibbett & Thorn, 2001 Filobasidium uniguttulatum Filobasidiales Geastrum sp. Geastrales Chapter 2 50 Appendix Chapter 2 - continued Appendix Chapter 2 - continued Species Order SPC-type Author Gloeophyllum sepiarium Neolentinus sufrutescens Lenzites sepiaria Lentinus lepideus Gloeophyllales Gloeophyllales perforate perforate Hyde & Walkinshaw, 1966 Keller, 1997 Ramaria ignicolor Clavaria ignicolor Gomphales imperforate Patrignani & Pellegrini, 1986 Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales imperforate imperforate imperforate imperforate imperforate imperforate perforate imperforate imperforate imperforate imperforate imperforate imperforate imperforate imperforate imperforate imperforate imperforate imperforate imperforate imperforate imperforate Müller et al., 2000b Müller et al., 2000b Müller et al., 2000b Müller et al., 2000b Müller et al., 2000b Oberwinkler, 1985 Keller, 1997; Langer & Oberwinkler, 1993 Langer & Oberwinkler, 1993 Keller, 1997 Greslebin et al., 2000 Keller, 1997 Langer & Oberwinkler, 1993 Keller, 1997 Keller, 1997; Langer & Oberwinkler, 1993 Langer & Oberwinkler, 1993 Keller, 1997 Wu & Huang, 1997 Langer & Oberwinkler, 1993 Langer & Oberwinkler, 1993 Keller, 1997 Keller, 1997; Langer & Oberwinkler, 1993 Wu & Huang, 1997 Asterodon ferruginosum Coltricia perennis Coltriciella dependens Hydnochaete japonica Hymenochaete cyclolamellata Hymenochaete rubiginosa Hyphoderma praetermissum Hyphodontia alutaria Hyphodontia arguta Hyphodontia australis Hyphodontia barba-jovis Hyphodontia cineracea Hyphodontia crustosa Hyphodontia loccosa Hyphodontia gossypina Hyphodontia hastate Hyphodontia mollis Hyphodontia pallidula Hyphodontia radula Hyphodontia rimosissima Hyphodontia sambuci Hyphodontia subglobosa Cyclomyces fuscus Basidioradulum radula Hyphodontia verruculosa Hyphoderma sambuci 51 Appendix cited as Species Inonotus hispidus Inonotus leporinus Inonotus weirii Onnia circinata Onnia tomentosa Oxyporus latemarginatus Phellinus igniarius var. igniarius Phellinus torulosus Phellinus tuberculosus Porodaedalea chrysoloma Rickenella ibula Schizopora paradoxa Trichaptum abietinum Trichaptum biforme cited as Onnia leporina Polyporus tomentosus Poria latemarginata Fomes igniarius Phellinus chrysoloma Rickenella aulacomniophila Hirschioporus abietinus Hirschioporus pargamenus Clathrus cancellatus Phallus impudicus Order SPC-type Author Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales imperforate imperforate imperforate imperforate imperforate Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales Hymenochaetales perforate imperforate imperforate imperforate imperforate perforate imperforate imperforate imperforate Moore, 1980 Moore, 1980 Müller et al., 2000b Moore, 1980 Moore, 1980; Müller et al., 2000b; Setlif et al., 1972 Setlif et al., 1972 Shukla, 1975 Moore, 1980 Keller, 1997 Keller, 1997 Kost, 1984; Van Driel et al., Chapter 2 Figure 1 Langer & Oberwinkler, 1993 Moore, 1985 Traquair & McKeen, 1978 Phallales Phallales perforate perforate Rexer & Stepanova, 2004 (reference herein) Patton & Marchant, 1978a Keller, 1997; Moore & Marchant, 1972; Moore 1975, 1985; Patton & Marchant, 1978a, b Keller, 1997 Bianchinotti et al., 2005 Patton & Marchant, 1978a Keller, 1997 Keller, 1997 Moore, 1980 Abortiporus biennis Polyporis biennis Polyporales perforate Bulbillomyces farinosus Ceraceomyces americanus Climacodon septentrionalis Conohypha terricola Flavophlebia sulfureoisabellinum Fomes fomentarius Polyporales Polyporales Rhizochaete americana Polyporales Hydnum septentrionale Polyporales Cerocorticium sulfureoisabellinum Polyporales Polyporales perforate perforate perforate perforate perforate perforate Chapter 2 52 Appendix Chapter 2 - continued Appendix Chapter 2 - continued Species Ganoderma lucidum Grifola frondosa Gyrophanopsis polonensis Hyphoderma mutatum Hyphoderma setigerum Hyphoderma subdeinitum Hypochnicium bombycinum Hypochnicium eichleri Hypochnicium lundellii Hypochnicium punctulatum Meruliopsis taxicola Merulius tremellosus Mycoacia fuscoatra Mycoacia uda Osteina obducta Phaeolus schweinitzii Phanerochaete chrysosporium Hypochnicium polonense Hypochnicium sphaerosporum Sporotrichum pruinosum, Chrysosporium xerophilum Rhizochaete ilamentosa Rhizochaete radicata Favolus alveolaris Order SPC-type Author Polyporales Polyporales Polyporales Polyporales Polyporales Polyporales Polyporales Polyporales Polyporales Polyporales Polyporales Polyporales Polyporales Polyporales Polyporales Polyporales Polyporales perforate perforate perforate perforate perforate perforate perforate perforate perforate perforate perforate perforate perforate perforate perforate perforate perforate Keller, 1997 Keller, 1997 Keller, 1997; Langer & Oberwinkler, 1993 Keller, 1997 Langer & Oberwinkler, 1993 Keller, 1997 Langer & Oberwinkler, 1993 Keller, 1997 Keller, 1997 Keller, 1997 Keller, 1997 Moore & McAlear, 1962 Keller, 1997 Keller, 1997 Keller, 1997 Moore, 1980 Boekhout et al., 1989 Polyporales Polyporales Polyporales Polyporales Polyporales Polyporales Polyporales Polyporales Polyporales perforate perforate perforate perforate perforate perforate perforate perforate perforate Bianchinotti et al., 2005 Bianchinotti et al., 2005 Bianchinotti et al., 2005 Keller, 1997 Bianchinotti et al., 2005; Tsuneda et al., 1993 Bianchinotti et al., 2005 Flegler et al., 1976 Wilsenach & Kessel, 1965 Keller, 1997 53 Appendix Phanerochaete ilamentosa Phanerochaete radicata Phanerochaete velutina Phlebia ochraceofulva Phlebia radiata Phlebia rufa Polyporus alveolaris Polyporus rugulosus Polyporus squamosus cited as Species Polyporus tuberaster Rhizochaete brunnea Rhodonia placenta Scutiger oregonensis Sparassis crispa Sporotrichum aurantiacum Steccherinum bourdotii Trametes versicolor Albatrellus ovinus Albatrellus subrubescens Aleurodiscus aurantius Asterostroma medium Auriscalpium vulgare Gloeocystidiellum lactescens Gloeocystidiellum porosum Gloiothele citrina Hericium coralloides Laxitextum bicolor Peniophora laeta Scytinostroma duriusculum Scytinostromella olivaceoalba Spiniger meineckellus Stereum hirsutum Zelleromyces stephensii cited as Poria monticola Albatrellus pes-caprae Steccherinum robustius Coriolus versicolor, Polystictus versicolor Megalocystidium lactescens Vesiculomyces citrinus Confertobasidium olivaceoalbum Order SPC-type Author Polyporales Polyporales Polyporales Polyporales Polyporales Polyporales Polyporales Polyporales perforate perforate perforate perforate perforate perforate perforate perforate Moore, 1980 Bianchinotti et al., 2005 Wilsenach & Kessel, 1965 Keller, 1997 Patrignani & Pellegrini, 1986 Boekhout et al., 1989 Keller, 1997 Aylmore et al., 1984; Girbardt, 1958,1961 Russulales Russulales Russulales Russulales Russulales Russulales Russulales Russulales Russulales Russulales Russulales Russulales Russulales Russulales Russulales Russulales perforate perforate perforate perforate perforate perforate perforate perforate perforate perforate perforate perforate imperforate perforate perforate perforate Keller, 1997 Keller, 1997 Keller, 1997 Müller et al., 2000b Keller, 1997 Keller, 1997 Keller, 1997 Keller, 1997 Flegler et al., 1976 Keller, 1997 Keller, 1997 Besson & Fremont, 1964 Keller, 1997 Hanlin, 1978 Patrignani & Pellegrini, 1986 Keller, 1997 Chapter 2 54 Appendix Chapter 2 - continued Appendix Chapter 2 - continued Species SPC-type Author Sebacinales Sebacinales Sebacinales imperforate imperforate imperforate Keller, 1997 Wells & Oberwinkler, 1982 Khan & Kimbrough, 1980 (perforate); Berbee & Wells, 1988 (imperforate; reference herein); Wells, 1994 (imperforate; reference herein) Sebacinales Sebacinales Sebacinales imperforate imperforate imperforate Verma et al., 1998 Keller, 1997 Andersen, 1996; Williams & Thilo, 1989 Sebacinales Sebacinales Sebacinales imperforate imperforate imperforate Keller, 1997 Khan & Kimbrough, 1980 Currah & Sherburne, 1992; Oberwinkler, 1985; Williams & Thilo, 1989 Sebacinales imperforate Müller et al., 1998b; Williams & Thilo, 1989 Tremellodendron candidum Sebacinales imperforate Tremelloscypha australiensis Tremelloscypha gelatinosa Tremelloscypha sp. Sebacinales Sebacinales Sebacinales imperforate imperforate imperforate Khan & Kimbrough, 1980; Wells & Oberwinkler, 1982 Wells & Oberwinkler, 1982 Wells & Oberwinkler, 1982 Oberwinkler, 1985 Bankera violascens Hydnellum concrescens Sarcodon versipellis Thelephora anthocephala Thelephora terrestris Thelephorales Thelephorales Thelephorales Thelephorales Thelephorales perforate perforate perforate perforate perforate Keller, 1997 Keller, 1997 Keller, 1997 Patrignani & Pellegrini, 1986 Keller, 1997; Langer, 1994 Piriformospora indica Sebacina epigaea Sebacina grisea Exidiopsis fugacissima, Sebacina fugacissima Exidiopsis grisea, Exidiopsis plumbescens Sebacina helvelloides Sebacina incrustans Sebacina sp. Serendipita vermifera Sebacina vermifera, Exidiopsis vermifera Thelephora palmata 55 Appendix Order Craterocolla cerasi Eibulobasidium rolleyi Microsebacina fugacissima cited as Species Order SPC-type Author Thelephorales Thelephorales Thelephorales Thelephorales perforate perforate perforate perforate Keller, 1997 Calonge, 1969 Keller, 1997 Calonge, 1969; Keller, 1997 Tomentellopsis echinospora Tomentellopsis submollis Thelephorales Thelephorales perforate perforate Keller, 1997 Keller, 1997 Subulicystidium longisporum Trechisporales imperforate Keller, 1997 Bullera variabilis Bulleromyces albus Cryptococcus laurentii var. laurentii Ditangiibulae dikaryotae Fibulobasidium inconspicuum Filobasidiella depauperata Filobasidiella neoformans Phragmoxenidium mycophilum Rhynchogastrema coronatum Sirobasidium magnum Syzygospora alba Syzygospora eibulata Carcinomyces eibulatus Syzygospora pallida Christiansenia pallida Tremella brasiliensis Tremella foliacea Tremella fuciformis Tremellales Tremellales Tremellales Tremellales Tremellales Tremellales Tremellales Tremellales Tremellales Tremellales Tremellales Tremellales Tremellales Tremellales Tremellales Tremellales vesicular vesicular absent reticulate vesicular vesicular absent absent vesicular vesicular absent absent absent vesicular vesicular vesicular Boekhout et al., 1991 (cupulate) Boekhout et al., 1991 (cupulate) Rhodes et al., 1981 Adams et al., 1995 Wells, 1994 (reference herein) Kwon-Chung et al., 1995 (cupulate) Kwon-Chung & Popkin, 1976 Wells, 1994 (reference herein) Wells, 1994 (reference herein) Moore, 1978a (ampulliform vesicles) Wells, 1994 (references herein) Wells, 1994 (reference herein) Wells, 1994 (references herein) Moore, 1978b Wells, 1994 (reference herein) Moore, 1978b (reference herein) Tomentella crinalis Tomentella fuscoferruginosa Tomentella pilosa Tomentellina ibrosa cited as Tomentella ibrosa, Tomentella bombycina Chapter 2 56 Appendix Chapter 2 - continued Appendix Chapter 2 - continued Species cited as Order SPC-type Author Tremella globospora Tremellales vesicular Tremella mesenterica Tremella rhytidhysterii Tremella sp. Tremella uliginosa Tremellales Tremellales Tremellales Tremellales vesicular vesicular vesicular vesicular Trimorphomyces papilionaceus Tremellales vesicular Berbee & Wells, 1988; Oberwinkler, 1985 (sacculate) Moore, 1978b; Wells, 1994 Wells, 1994 (reference herein) Weiss et al., 2004 (sacculate) Berbee & Wells, 1988 (references herein); Wells, 1994 (reference herein) Berbee & Wells, 1988 (references herein); Wells, 1994 (reference herein) Trichosporon asahii Trichosporon brassicae Trichosporon coremiiforme Trichosporon cutaneum Trichosporon inkin Trichosporon laibachii Trichosporon moniliiforme Trichosporon mucoides Trichosporon sporotrichoides Trichosporonales Trichosporonales Trichosporonales Trichosporonales Trichosporonales Trichosporonales Trichosporonales Trichosporonales Trichosporonales vesicular absent vesicular vesicular absent vesicular vesicular absent vesicular Guého et al., 1992 (tubular/vesicular) Guého et al., 1992 (non-perforate septum) Guého et al., 1992 (tubular) Guého et al., 1992 Guého et al., 1992; Fell et al., 2001 Guého et al., 1992; Fell et al., 2001 Guého et al., 1992 Guého et al., 1992 Guého et al., 1992; Müller et al., 1995, 1998a, 2000c (tubular, globular) Wallemia sebi Wallemiomycetes vesicular Moore, 1986 Tetragoniomyces uliginosa Table 1 – Septal pore cap type in the Agaricomycotina. Orders according to Hibbett et al. (2007) plus addition of the order Trichosporonales (Fell et al., 2000). Entorrhizomycetes and Wallemiomycetes are two unplaced classes in the Agaricomycotina (Hibbett et al., 2007). Current use of names was veriied in Mycobank (www. mycobank.org; Crous et al., 2004). The placement of Waitea circinata and Rhizoctoina zeae in Corticiales is still uncertain. Appendix 57 Chapter 2 REFERENCES APPENDIX Adams, G., Klomparens, K. & Hennon, P. (1995) Unusual reticulated parenthesomes surround the dolipore of a hyphomycete with clamp connections, Ditangiibulae dikaryotae gen. et sp. nov. Mycologia 87, 909 – 921. Andersen, T.F. (1996) A comparative taxonomic study of Rhizoctonia sensu lato employing morphological, ultrastructural and molecular methods. Mycol. Res. 100, 1117 – 1128. Aylmore, R.C., Wakley, G.E. & Todd, N.K. (1984) Septal sealing in the Basidiomycete Coriolus versicolor. J. Gen. Microbiol. 130, 2975 – 2982. Bauer, R., Oberwinkler, F. & Vanky, K. 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