Taxon circumscription
- General introduction
- Introduction to generic arrangement
- Arrangement of Australian agaric genera within families
- Comparison with other classifications
- Circumscription and placement of genera
- References
Following an arrangement of Australian agaric genera within families, we discuss the circumscription of each family (in terms of which genera belong within it) and its genera (in terms of any different versions of the limits of the genera).
The placement of agaric genera within families has changed markedly over the last few decades with the use of molecular data in the reconstruction of evolutionary history. The previous systems based on morphological, chemical and cytological data, such as laid out by Kühner (1980) and Singer (1986), have been found to vary considerably from classifications based on molecular phylogeny. Although some families are similar in composition for both morphological and molecular classifications, many are not. In addition, analyses of molecular data have often led to the creation of segregate genera, or the merging of long-established genera. In such rearrangements, the novel delimitations of genera can follow existing groups established at infrageneric taxa, or bring together species that were not suspected to be closely realted on morphological evidence alone.
A species of Melanoleuca. This genus was formerly included in the Tricholomataceae, but molecular data now place it in the Pluteaceae.
Singer (1986) defined the family Pluteaceae to include Amanita, Limacella, Pluteus and Volvariella. Molecular analyses (Moncalvo et al., 2002; Matheny et al., 2007a) originally supported delimitation of genera themselves, but later (Justo et al., 2011) Volvariella was found to be polyphyletic, necessitating erection of the genus Volvopluteus for V. gloiocephala and its close allies. In terms of relationships, based on molecular data Pluteus and Volvopluteus are not related to Amanita and Limacella, and the latter are segregated in the family Amanitaceae, while the Pluteaceae is emended to also include Melanoleuca, a genus with ornamented spores which Singer (1986) had placed in the Tricholomataceae. The placement of Volvariella in the strict sense (exclusive of Volvopluteus) is unresolved, but it belongs in neither Amanitaceae nor Pluteaceae.
While the rapid changes to the classification of agarics are an exciting development, they do make recognition of coherent taxa quite difficult in some families, especially where sequence data are available for few species. Indeed, the full ramifications of molecular data, in terms of the need to redefine generic limits, have still not been addressed in some genera. In choosing taxa to use for identification purposes we have tried to align the limits of the identification units to genera (or subdivisions of genera) that are circumscribed phylogenetically. However, because FunKey utilises morphology, in some cases the identification units are coherent morphologically, but molecular data indicate heterogeneity. In such cases, new combinations in segregate genera are usually not yet available as in Psilocybe sens. lat., and are mentioned in the discussion of the relevant families.
The most recent comprehensive arrangements of agaric genera in families, accompanied by supporting discussion, were those of Kühner (1980) and Singer (1986), and the process of incorporating molecular data into such classifications is ongoing. The most comprehensive molecular data are derived from multilocus studies with extensive taxon sampling, such as that of Matheny et al. (2007a). However, some genera have not been subjected to any molecular investigation, or perhaps only a single DNA region has been sequenced. In addition, even when multilocus data are available, they do not allow unequivocal placement of some genera into established families. Thus, the most up-to-date classification does not yet provide a complete transition from older arrangements, in terms of allocating all genera to named families.
The following arrangement of genera within families is based primarily on the multi-locus molecular phylogenies for Agaricales (Matheny et al., 2007a), Boletales (Binder & Hibbett, 2007), Cantharellales (Moncalvo et al., 2007), Hymenochaetales (Larsson et al., 2007) and Russulales (Miller et al., 2007). Genera not included in these studies are designated either by an asterisk, where the family placement is based on the analysis of the nuclear large ribosomal subunit (nLSU) by Moncalvo et al. (2002), where in some instances there was poor statistical support for some relationships, or a double asterisk, where family placement follows other sources as indicated in the discussion under the particular family (and molecular data can be lacking).
In the Agaricales, some of the major clades recognised by Matheny et al. (2007a) do not have formal names at family level. Consequently, four of the taxa listed below, which we consider to be at family rank, are indicated as un-named, and are designated either by an existing name of a tribe (Cystodermateae and Gymnopileae) or by an informal clade name (/hydropoid clade and /xeromphalinoid clade). Furthermore, five taxa (Macrocybe, Mycenella, Panellus longinquus, Singerocybe and Tricholomopsis) cannot be unambiguously placed in families on the basis of current molecular evidence.
Some authors, such as Moncalvo et al. (2002), use informal names for clades that are denoted by a slash '/' before the name, as in '/omphalotus'. Such clade names are a combination of names of formal taxa (such as genera or families, but used with a lower case initial letter) or adjectives derived from generic names, such as '/hydropoid'.
Several families contain a mixture of agaric and non-agaric genera, the latter can have quite different fruit-body forms, such as boletes or polypores. Genera within the following families which only contain non-agaric forms are not listed below. Moreover, a few genera include species with agaric (lamellate) fruit-bodies and species with other fruit-body types, especially truffle-like. It is clear that the agaric fruit-body form has evolved in a variety of unrelated lineages. In other words, agarics do not form a single, uniform group within the modern classification. Nevertheless, the agaric fruit-body form is readily recognisable, and it is practical to deal with the agarics as a morphological unit for the purposes of identification.
The various orders, and families within, are listed alphabetically. All orders are referable to the phylum Basidiomycota, class Agaricomycetes. The orders Agaricales and Boletales belong to the subclass Agaricomycetidae. The other orders (Cantharellales, Gloeophyllales, Hymenochaetales, Polyporales and Russulales) are unplaced as to their position within subclasses of the Agaricomycetes. In the list below, orders and families are in bold, with orders in capitals.
This arrangement does not include Excluded taxa.
*, ** For explanation of asterisks, see above.
Below, and in the Taxon Fact Sheets, an equals sign at the start of a list of one or more names in parentheses (=) indicates that the name/s within the parentheses are synonyms of the name preceding the parentheses.
AGARICALES
Agaricaceae
Agaricus, Chlorophyllum, Coprinus, *Gyrophragmium, Lepiota, Leucoagaricus, *Leucocoprinus, Macrolepiota, *Melanophyllum, *Montagnea.
Amanitaceae
Amanita (= Amanitopsis), *Limacella.
Bolbitiaceae
Bolbitius, Conocybe, Descolea, Pholiotina.
Clavariaceae
Camarophyllopsis (= Hygrotrama).
Cortinariaceae
Cortinarius (= Cuphocybe, Dermocybe,
Rapacea, Rozites).
Crepidotaceae
Crepidotus (= Pleurotellus), Pleuroflammula, Simocybe.
Cyphellaceae
Cheimonophyllum.
Entolomataceae
Clitopilus (= Rhodocybe), Entoloma (= Alboleptonia, Claudopus, Eccilia, Inocephalus, Inopilus, Leptonia, Nolanea, Pouzarella, Pouzaromyces, Rhodophyllus).
*Fayodiaceae
*Conchomyces.
Hydnangiaceae
Laccaria.
Hygrophoraceae
*Arrhenia, Humidicutis, Hygrocybe (= Bertrandia, Camarophyllus, Cuphophyllus, Gliophorus), Hygrophorus, Lichenomphalia (= Botrydina, Phytoconis).
Hymenogastraceae
Galerina, Hebeloma, Phaeocollybia, Psilocybe [in the strict sense, see also Deconica under Strophariaceae].
Inocybaceae
Inocybe (= Astrosporina, Auritella).
Lyophyllaceae
Asterophora (= Nyctalis), Lyophyllum (= Tephrocybe).
Macrocystidiaceae
Macrocystidia.
Marasmiaceae
Campanella, Chaetocalathus, Crinipellis, Marasmius, Tetrapyrgos.
Mycenaceae
Cruentomycena, Mycena sens. strict. [exclusive of section Adonideae], Panellus [exclusive of P. longinquus], Roridomyces.
Omphalotaceae
Anthracophyllum, Gymnopus (= Micromphale, Setulipes), Lentinula, *Marasmiellus [M. affixus only], Omphalotus, Rhodocollybia.
Panaeolaceae
Panaeolina, **Panaeolopsis, Panaeolus (= Anellaria, Copelandia).
Physalacriaceae
Armillaria (= Armillariella), Cyptotrama, Flammulina, Oudemansiella (= Hymenopellis, Protoxerula).
Pleurotaceae
Hohenbuehelia, Pleurotus, Resupinatus.
Pluteaceae
Melanoleuca, Pluteus, Volvariella.
Psathyrellaceae
Coprinellus, Coprinopsis (= Rhacophyllus), *Coprinus cordisporus, Lacrymaria, *Parasola, Psathyrella.
Schizophyllaceae
Schizophyllum.
Strophariaceae
Agrocybe, Deconica [keyed out under Psilocybe], Hypholoma (= Naematoloma), Leratiomyces (= Stropholoma), *Melanotus, Pholiota, Stropharia.
Tricholomataceae
Clitocybe [see also Singerocybe under Unplaced within Agaricales. 5], Lepista, Leucopaxillus, *Porpoloma, Tricholoma.
Tubariaceae
Tubaria.
Un-named family (tribe: Cystodermateae)
Cystoderma.
Un-named family (tribe: Gymnopileae)
Gymnopilus (= Pyrrhoglossum).
Un-named family (/hydropoid clade)
Mycena section Adonideae.
*Un-named family (/xeromphalinoid clade)
*Heimiomyces, Xeromphalina.
Unplaced within Agaricales 1.
*Macrocybe.
Unplaced within Agaricales 2.
**Mycenella.
Unplaced within Agaricales 3.
Panellus longinquus (as Pleurotopsis).
Unplaced within Agaricales 4.
Tricholomopsis.
Unplaced within Agaricales 5.
Singerocybe [keyed out under Clitocybe].
BOLETALES
Boletaceae
Phylloporus.
Hygrophoropsidaceae
Hygrophoropsis.
Paxillaceae
**Meiorganum, Paxillus.
Serpulaceae
Austropaxillus.
Tapinellaceae
Tapinella.
CANTHARELLALES
Cantharellaceae
Cantharellus, Craterellus.
GLOEOPHYLLALES
Gloeophyllaceae
Gloeophyllum, Neolentinus.
HYMENOCHAETALES
Repetobasidiaceae
Loreleia, Rickenella.
POLYPORALES
Polyporaceae
Austrolentinus, Lentinus, Lenzites, Panus, Trametes (= Artolenzites).
RUSSULALES
Auriscalpiaceae
Lentinellus.
Russulaceae
Lactarius (= Lactifluus, Multifurca), Russula.
The classification adopted by the 10th edition of the Dictionary of the Fungi (Kirk et al., 2008) is available on-line, with some modifications. It incorporates many of the changes in family circumscription arising from recent studies, such as Matheny et al. (2007a). However, it differs from the arrangement we adopt as follows (in terms of family placement only). Firstly, several families are combined: Crepidotaceae is merged with Inocybaceae, Hymenogastraceae with Strophariaceae, and Omphalotaceae with Marasmiaceae. These three pairs of families are each sister taxa in the phylogenetic reconstruction (Matheny et al., 2007a), so the question of whether to merge or keep separate is a matter of opinion. Secondly, some individual genera have family placements in the Dictionary of the Fungi 10th edn at variance with Matheny et al. (2007a) or other recent molecular analyses, not as a result of merging of sister taxa. These genera are: Arrhenia (Tricholomataceae in the Dictionary not Hygrophoraceae), Camarophyllopsis (unplaced, not Clavariaceae), Conchomyces (Tricholomataceae not Fayodiaceae), Cystoderma (Agaricaceae rather than unplaced), Descolea (Cortinariaceae not Bolbitiaceae), Gymnopilus (Strophariaceae rather than unplaced), Heimiomyces and Xeromphalina (both Mycenaceae rather than unplaced), Loreleia (unplaced, rather than Repetobasidiaceae), Macrocybe (Tricholomataceae rather than unplaced), Macrocystidia (Marasmiaceae not Macrocystidiaceae), Melanoleuca (Tricholomataceae not Pluteaceae), Mycenella (Tricholomataceae rather than unplaced), Neolentinus (Polyporaceae not Gloeophyllaceae), Panaeolus and Panaeolina (both unplaced not Panaeolaceae), Panaeolopsis (Agaricaceae not Panaeolaceae), Phaeocollybia (Cortinariaceae not Hymenogastraceae), Resupinatus (Tricholomataceae not Pleurotaceae), Tubaria (Inocybaceae rather than unplaced) and Tricholomopsis (Tricholomataceae rather than unplaced).
The classification adopted by Funga Nordica (Knudsen & Vesterholt, 2008) differs from that followed here for family circumscription in that Crepidotaceae is merged with Inocybaceae; Mycenaceae is called Favolaschiaceae; Porotheleaceae encompasses both Fayodiaceae and the /hydropoid clade of Matheny et al. (2007a); and Resupinatus is placed in the Resupinataceae (separate from the Pleurotaceae). Other differences are: Arrhenia (Typhulaceae in Funga Nordica not Hygrophoraceae), Cystoderma (Squamanitaceae rather than unplaced), Loreleia and Rickenella (unplaced rather than Repetobasidiaceae), Melanoleuca (unplaced rather than Pluteaceae), Mycenella (Physalacriaceae rather than unplaced), Panaeolus and Panaeolina (both Bolbitiaceae not Panaeolaceae), Psilocybe (Strophariaceae not Hymenogastraceae) and Xeromphalina (Typhulaceae rather than unplaced).
In the following: orders, families and other suprageneric taxa are as in the arrangement of genera and each taxon (entity) as used in FunKey (whether a genus, a genus represented by a single species or species group or an identification unit within a genus) is shown in bold once under its correct family.
In the discussion below, we usually use the generic placement and circumscription of Singer (1986) as a baseline against which to compare the ramifications of analyses of molecular data, which have mostly occurred since 2000. Molecular data are lacking for most agarics described from Australia, and Singer (1986) is the work that has most often been utilised in placing species into genera and infrageneric taxa on the basis of morphology. In particular, Grgurinovic (1997a) largely followed Singer (1986) in classifying the numerous Australian species described by Cleland.
AGARICALES
Agaricaceae
Vellinga (2004) used nLSU and ITS data to confirm that the Agaricaceae included the following Australian genera: Agaricus (including Gyrophragmium), Chlorophyllum, Coprinus sens. strict., Lepiota, Leucocoprinus (including Leucoagaricus), Macrolepiota, Melanophyllum and Montagnea. This circumscription was confirmed from multilocus molecular data by Matheny et al. (2007a), although fewer genera were sampled. The Agaricaceae also includes a number of gasteroid (puffball) genera such as Langermannia and Tulostoma. Apart from those, the delimitation of the Agaricaceae on molecular grounds agrees with its morphological scope as set out by Singer (1986) apart from the inclusion of Coprinus sens. strict., Gyrophragmium and Montagnea, and with the exclusion of some extra-Australian genera such as Cystoagaricus.
Agaricus has long been recognised as distinct (Singer 1986), at least its temperate species. Singer (1986) included it in the tribe Agariceae, and various species of Agaricus form a well-supported clade in molecular analyses (Moncalvo et al., 2002; Vellinga et al., 2003a; Vellinga, 2004; Walther et al., 2005; Matheny et al., 2007a). Singer (1986) tentatively placed the tropical Hymenagaricus (distinguished by the cellular pileipellis) under Agaricus. Molecular data from single species of each of Hymenagaricus and Micropsalliota (another genus in the Agariceae, with cheilocystidia) have been included in a number of studies. Isolates of the two genera are sister to Agaricus (Vellinga, 2004; Matheny et al., 2007a) but further taxon sampling from all three genera is required before the separation of Hymenagaricus and Micropsalliota can be confirmed. No species resembling Hymenagaricus or Micropsalliota have been reported from Australia.
Molecular data are available for one species of Gyrophragmium: G. dunalii (which is an earlier name for the type of the genus, G. delilei). This species falls within Agaricus in phylogenetic analyses (Moncalvo et al., 2002; Vellinga, 2004) and therefore has recently been formally transferred to that genus as has Longula texensis [which has at times been placed in Gyrophragmium] (Geml et al., 2004). For the moment Gyrophragmium inquinans is maintained in Gyrophragmium, but it would not be surprising if this species is eventually transferred to Agaricus sens. strict.
Coprinus sens. strict., consisting of C. comatus and a few other species, has been transferred to the Agaricaceae (Redhead et al. 2001) with the numerous other taxa formerly in Coprinus placed in several genera of the Psathyrellaceae. We refer to Coprinus sens. strict. as the Coprinus comatus group to emphasise that most Coprinus species now belong elsewhere. Fruit-bodies of Montagnea resemble a dried Coprinus, and while micromorphology is also similar, spores are not forcibly discharged. In the molecular analyses of Moncalvo et al. (2002) and Vellinga (2004), three species of Montagnea successively clustered at the base of the clade for Coprinus sens. strict.; hence the former is paraphyletic. One solution would be to synonymise Montagnea with Coprinus sens. strict. However, Redhead et al. (2001), on the basis of the earlier molecular analysis of Johnson & Vilgalys (1998) where a single species of Montagnea also clustered at the base of the Coprinus sens. strict. clade, argued that the two genera should be maintained pending further molecular analysis, because the evolution of forcible discharge (as in Coprinus) from ancestral species lacking that attribute (as in Montagnea) was unknown elsewhere in the euagarics.
Singer (1986) placed three genera lacking clamp connections (Leucoagaricus, Leucocoprinus and Seriomyces) in the tribe Leucocoprineae of the Agaricaceae. The latter genus (not known from Australia) contained only a few species, that were pure white and had spores that lacked a germ pore. He distinguished Leucocoprinus from Leucoagaricus by the plicate pileus margin of the former and the frequent occurrence of pseudoparaphyses (pavement cells) in the hymenium.
Vellinga (2004) gathered molecular data from numerous species of Leucocoprinus, Leucoagaricus and Sericeomyces, and found a large clade (albeit without strong statistical support) containing a mixture of members of all three genera, and several isolates of non-sporing fungi cultivated by ants. Vellinga (2004) noted that an alternative arrangement (where Leucocoprinus was monophyletic) could not be rejected, but Leucoagaricus and Sericeomyces were certainly polyphyletic. She advocated accepting the large assemblage of Leucoagaricus, Leucocoprinus and Sericeomyces as a single genus, pending further analysis that could result in this group being split into smaller taxa. The correct name for the combined genus is Leucocoprinus, but Vellinga (2004) did not formally transfer species of the other two genera to it. She also noted that some species that belong in Leucocoprinus only have valid names in Lepiota at present. Consequently, the genera Leucoagaricus and Leucocoprinus are retained in the sense of Singer (1986). The Leucoagaricus leucothites group is keyed out separately for species with a relatively smooth, white to brown or grey pileus. Fruit-bodies are rather similar to Agaricus in the field, except for the white lamellae. In Leucoagaricus (other), the pileus usually has darker fibrils and the fruit-body frequently stains red on cutting or bruising.
Chlorophyllum (green spore print) and Macrolepiota (white spore print) were also placed in the tribe Leucoprineae of the Agaricaceae by Singer (1986). Analyses of molecular data (Vellinga, 2003a; 2004; Vellinga et al., 2003) led to the surprising conclusion that some species included by Singer (1986) in Macrolepiota now belong in Chlorophyllum (which also includes the sequestrate Endoptychum agaricoides, although not other species of that genus). The genus Macrolepiota was restricted to species where the pileipellis is a trichoderm compared to the hymenidermal (or epithelial) pileipellis of Chlorophyllum. In addition, species of Macrolepiota have a hyaline cap over the germ pore which is absent in Chlorophyllum. Within Chlorophyllum, the green-spored Chlorophyllum molybdites is keyed out separately to the remaining species in Chlorophyllum (other), those having white spore prints. Chlorophyllum hortense is also keyed out separately due to its similarity to Leucocoprinus (by the sulcate pileus margin, reddening flesh and spores lacking a germ pore) rather than other members of Chlorophyllum, although it is separated from Leucocoprinus by the presence of clamp connections.
Lepiota, as characterised by Singer (1986) in the tribe Lepioteae of the Agaricaceae, is a large genus, most species of which have dextrinoid spores and clamp connections. He recognised three sections that did not have this combination of characters: section Amyloideae for a few species, including Lepiota lignicola (which has been transferred to Leucopholiota), with amyloid spores and clamp connections; section Anomalae for a few species that lacked clamp connections; and section Amylosporae for a couple of species with amyloid spores and no clamp connections. The remaining sections were characterised by the nature of the pileipellis and the shape of the spores. The pileipellis consisted of spines made up of sphaerocysts in section Echinatae (recognised at generic rank by some authors as Echinoderma), and it was a hymeniderm in section Cristatae. The other sections had a trichodermal pileipellis, with the spores spurred in section. Stenosporae, fusoid in section Lepiota and not fusoid in section Ovisporae. Singer (1986) also accepted in the tribe Lepioteae the genus Cystolepiota for species usually with clamp connections and with an epithelial pileipellis (of rounded elements) and inamyloid or dextrinoid spores. Pulverolepiota was subsequently described by Bon (1993) for species with an epithelioid pileipellis of elongate elements, rough spores and lacking clamp connections. Melanophyllum, another genus with an epithelioid pileipellis has finely warted spores and clamp connections. It was placed in the tribe Agariceae by Singer (1986) on account of the green spore print.
Vellinga (2003c) analysed numerous species from most of the sections of Lepiota outlined by Singer (1986) and potentially related genera such as Cystolepiota, Melanophyllum and Pulverolepiota. Interpretation of the analyses was complicated by some discordance between phylogenies from different DNA regions, and by low statistical support for some clades. Nevertheless, Vellinga (2003c) identified four main clades within Lepiota and allied species, although these clades ‘do not correspond to currently recognised genera, subgenera and sections’. Three clades consisted of species of Lepiota, one with an exclusively hymeniform pileipellis, and two other clades with a trichodermal pileipellis (in one clade made up of both long and short elements, and in the other made up solely of long elements). While the nature of the pileipellis was consistent within each of the three clades, spore shape varied widely. A few species of Lepiota were placed in between the three main clades, including Lepiota fuscovinacea (without clamps) from section Anomalae. The fourth clade identified by Vellinga (2003c) contained species of Cystolepiota, Melanophyllum, Pulverolepiota and Lepiota section Echinatae. She suggested that this group could be accommodated in a single genus. The generic name with priority appears to be Melanophyllum, but no formal transfers were made from other genera. Vellinga (2003c) did not comment on how the three clades of Lepiota should be treated, but Vellinga (2004) accepted a single genus for Lepiota, once section Echinatae had been removed. Further sampling, particularly of tropical species, is required before a new classification of the tribe Lepioteae is settled. For the moment, we recognise Lepiota as containing only species with dextrinoid spores and clamp connections. Consequently a number of species included by Grgurinovic (1997a) in Lepiota are not included in FunKey, because they either lack clamp connections (and hence probably belong in Leucocoprinus inclusive of Leucoagaricus) or they have amyloid spores (and hence are of uncertain generic disposition).
In FunKey, the sole Australian representative of Lepiota section Echinatae, Lepiota aspera, is keyed out separately, with the remaining species in Lepiota (other). Melanophyllum is also keyed out separately, in the restricted sense of Melanophyllum haematospermum. It is quite likely that species of Cystolepiota and Pulverolepiota occur in Australia but these await formal documentation.
Amanitaceae
Amanita (including Amanitopsis) and Limacella, as accepted on morphological grounds by Singer (1986), are also supported by molecular evidence as independent genera (Moncalvo et al., 2002). However, on the basis of evidence from a number of DNA regions, Matheny et al. (2007a) consider Pluteus and Volvariella, which Singer (1986) also included in the Amanitaceae, to belong in the Pluteaceae. Matheny et al. (2007a) did not sample Limacella, and the placement of this genus in Amanitaceae therefore follows the finding of Moncalvo et al. (2002), on the basis of nLSU sequences, that a clade containing several species of Limacella is sister to Amanita, although with weak statistical support.
While most species of Amanita have a well-developed volva, some do not, particularly those in section Lepidella subsection Solitariae and sections Amanita (where the volva is friable) and Validae. Therefore, Amanita (other) is keyed out separately from Amanita (no volva).
Bolbitiaceae
The Bolbitiaceae was characterised by Singer (1986) by the hymeniform pileipellis and brown spore print, and included Agrocybe, Bolbitius and Conocybe. Multilocus molecular data confirm the placement of Bolbitius and Conocybe in the Bolbitiaceae (Matheny et al., 2007a). However, Agrocybe is polyphyletic, and none of the species sampled belong in the Bolbitiaceae, most being close to the Strophariaceae (Moncalvo et al., 2002; Walther et al., 2005; Matheny et al., 2007a).
Singer (1986) and Arnolds (2005) treated Conocybe and Pholiotina as separate genera, distinguished by the presence of annulus (or veil remnants on the pileus margin) in the latter, and the abruptly capitate lamellar cystidia of the former. Watling (1982) had included Pholiotina under Conocybe (as subgenera Piliferae and Pholiotina). Walther et al. (2005) included ten species of Conocybe and Pholiotina (all as Conocybe) in a phylogenetic analysis of nLSU data. They found that the two species of Pholiotina fell in a well-supported clade that was sister to one containing species of Conocybe sens. strict. and a single collection of Bolbitius. The main Conocybe clade included representatives of three of the six sections in Conocybe and one of the three sections in Pholiotina recognised by Singer (1986). Species with rough spores (rarely found in both genera, and not known from Australia) were not sampled. Nevertheless, the available molecular data support the distinctiveness of Conocybe and Pholiotina.
Bolbitius was recognised on morphological grounds by Watling (1982), Singer (1986) and Arnolds (2005), being distinguished from Conocybe and Pholiotina by the fruit-body that rapidly collapses, the viscid, sulcate pileus, free lamellae and the lack of lecythiform cystidia. Molecular data on Bolbitius place it in the Conocybe plus Pholiotina clade, but either sister to Pholiotina (Moncalvo et al., 2002) or to Conocybe (Walther et al., 2005; Matheny et al., 2007a). All cladograms are consistent with an independent Bolbitius, and because the study of Matheny et al. (2007a) is based on more DNA regions, a relationship with Conocybe seems more probable, although the species of the three genera sampled in each of the three studies were different.
The genus Descolea is distinct morphologically (Horak, 1971; Singer, 1986), and all species for which DNA sequence data are available fall within a single clade, albeit intermixed with various sequestrate species (Peintner et al. 2001). There has been debate as to whether Descolea belongs in the Cortinariaceae (Horak, 1971) or Bolbitiaceae (Singer, 1986). The latter placement is more consistent with the presence of the hymeniform pileipellis, but the verrucose spores that lack a germ pore and the ectomycorrhizal habit are unusual in the Bolbitiaceae. A single nLSU sequence (Moncalvo et al., 2002) placed Descolea in an unsupported clade among members of the Bolbitiaceae and Panaeoloideae, and not with the various genera traditionally placed in the Cortinariaceae (that occurred in at least five separate clades). In contrast, Peintner et al. (2001), in a study utilising ITS sequences, treated Descolea a member of the Cortinariaceae. However, their analysis only included species traditionally placed in the Cortinariaceae, and no members of the Bolbitiaceae or Panaeoloideae. The placement of Descolea in the Bolbitiaceae was confirmed by the multi-locus study of Matheny et al. (2007a), where the Bolbitiaceae, including Descolea, is the sister group to the Cortinariaceae sens. strict., and genera of the Panaeoloideae fall in a separate part of the cladogram.
Clavariaceae
Camarophyllopsis is distinct morphologically, and was placed by Singer (1986) and Young (2005a) in the Hygrophoraceae (as Hygrotrama). Where molecular data are available, all other genera treated by Singer (1986) in this family fall in a single clade, with the exception of Camarophyllopsis, which rather surprisingly falls in the Clavariaceae alongside coral fungi such as Clavaria (Matheny et al., 2007a).
Cortinariaceae
Recent molecular studies of species from across the world (Peintner et al., 2002; 2004; Garnica et al., 2005) indicate that a number of genera formerly treated as independent entities within the Cortinariaceae on the basis of their morphology (Singer, 1986) belong in fact in Cortinarius. In addition, a morphology-based infrageneric classification of Cortinarius (Singer, 1986) is at odds with molecular data (Peintner et al., 2004; Garnica et al., 2005). However, a new classification that integrates all data sources is yet to be finalised, particularly with respect to relationships among the various clades revealed in the molecular analyses. Furthermore, many species do not fall within well-supported clades on the basis of the DNA regions so far sequenced, and taxon sampling of this very diverse genus is far from complete.
From analysis of multilocus molecular data (Matheny et al., 2007a), the Cortinariaceae is now restricted to Cortinarius. The family is sister to the Bolbitiaceae.
Due to the diversity and the range of morphology represented, ten identification units are used in FunKey for Cortinarius: Cortinarius australiensis, Cortinarius canarius, Cortinarius fibrillosus, Cortinarius mariae, Cortinarius subgenus Cortinarius, Cortinarius subgenus Dermocybe, Cortinarius subgenus Myxacium, Cortinarius morphogroup Rozites, Cortinarius morphogroup Cuphocybe, and Cortinarius (other) for remaining species.
In Cortinarius, four species are keyed out individually. Cortinarius australiensis has a massive fruit-body with a membranous annulus. It was originally placed in Rozites, but Horak (1981) recognised that it belonged in Cortinarius. Peintner et al. (2004) confirmed from DNA sequence data that C. australiensis is correctly placed in Cortinarius. Cortinarius canarius (a member of Cortinarius subgenus Dermocybe) is keyed out separately due to the unusual combination of very finely ornamented spores (that can appear smooth under the light microscope) and a rather membranous partial veil (that may leave an annulus, rather than the usual ring zone). Cortinarius fibrillosus has very finely ornamented spores, which can appear smooth under the light microscope. The pileus is radially fibrillose, and thus the species has often been placed in Inocybe, having also been described separately as Inocybe cystidiocatenata. Matheny & Ammirati (2003) confirmed the generic placement of Cortinarius fibrillosus. Cortinarius mariae was originally placed in the monotypic Rapacea (Horak 1999) due to the greenish spore print and very finely warty spores that can appear smooth under the light microscope, but molecular data show that it too is a Cortinarius (Peintner et al., 2004).
Cortinarius violaceus is the type of Cortinarius but occupies a rather isolated position within the genus as regards its morphology, along with the few other species that belong in subgenus Cortinarius. These species have cheilocystidia and pleurocystidia, and sometimes also caulocystidia and pileocystidia, and their spores can have a plage (Singer, 1986; Moser, 1987). Evidence from DNA sequences shows Cortinarius subgenus Cortinarius to be monophyletic, but it does not fully resolve relationships with other major clades within Cortinarius (Peintner et al., 2004; Garnica et al., 2005). Because the species of this group are distinctive macroscopically, by virtue of the violet pileus and stipe in combination with very dark or blackish lamellae, Cortinarius subgenus Cortinarius is keyed out separately.
The genus Rozites, long-established on morphological grounds, was thought to be distinct from Cortinarius because of its membranous annulus (Horak, 1981a; Singer, 1986). However, on molecular evidence, Peintner et al. (2002) showed that the type species of Rozites is referable to Cortinarius and, therefore, Rozites should be sunk into Cortinarius. Peintner et al. (2004) found that DNA sequences of other Rozites, including the Australian C. elacatipus (formerly R. foetens), also indicate synonymy with Cortinarius. Agerer (2006) questions the conclusions from molecular data and considers that on the basis of the structure of ectomycorrhizas formed by Rozites (including the lack of rhizomorphs and the amyloid reaction of hyphae), there ‘is not the slightest indication for a closer relationship of Rozites to the genus Cortinarius’ and suggests rather a relationship with Descolea or Inocybe. However, morphology of the fruit-body, apart from the annulus, is quite consistent with Cortinarius, and we follow Peintner et al. (2002; 2004) in synonymising Rozites. Because species formerly placed in Rozites fall in several clades within Cortinarius in the various molecular analyses, the group is not referred to a formal taxonomic unit, but rather species are assigned to Cortinarius morphogroup Rozites. It is possible that the various Australian species could be closely related, but sequence data are currently incomplete. The genus Cuphocybe, initially characterised by the squamulose pileus surface, was likewise shown by Peintner et al. (2002; 2004) to be a synonym of Cortinarius. However, because species referred to Cuphocybe fall in several clades (Garnica et al., 2005), they are referred to Cortinarius morphogroup Cuphocybe.
Cortinarius subgenus Myxacium, a very distinctive morphological group, is characterised by the pileus and stipe being viscid to glutinous. While it is employed for identification purposes, molecular data (Garnica et al., 2005) show that while its European species fall within a well-supported clade, the three Australian species analysed do not. Cortinarius archeri and C. sinapicolor cluster together, but C. erythraeus falls elsewhere within the cladogram.
Dermocybe was characterised by the presence of anthraquinone pigments (and thus frequently bright yellow, orange or red colours to the fruit-body), a usually non-hygrophanous pileus and the absence of a bulbous stipe base (Singer 1986), However, even on morphological and chemical grounds alone, Høiland (1983) preferred to recognise Dermocybe as a subgenus of Cortinarius. Molecular data clearly place Dermocybe within Cortinarius (Liu et al., 1997; Høiland & Holst-Jensen, 2000; Peintner et al., 2002; Garnica et al., 2005), and it is here treated as Cortinarius subgenus Dermocybe. One species of this subgenus, the bright yellow C. canarius, is keyed out separately (see above).
The various other species of Cortinarius which do not belong to any of the groups outlined above are aggregated as Cortinarius (other), which covers a wide range of macro-morphological forms, all of which have verrucose spores.
Crepidotaceae
The Crepidotaceae (including Crepidotus, Simocybe and Pleuroflammula) is well supported on multilocus molecular data and is sister to the Inocybaceae (Matheny et al., 2007a).
Singer (1986) accepted species with smooth or ornamented spores, as sections Crepidotus and Echinospori respectively. He differentiated the macroscopically identical Pleurotellus, with type P. hypnophilus, by the very pale spore print (‘cream buff’ rather than clay brown or cinnamon). The genus Melanomphalia (not yet reported from Australia) shared the verrucose spores of section Echinospori, but had a central stipe and often decurrent lamellae (Singer, 1986). Senn-Irlet (1995) treated Pleurotellus as a synonym of Crepidotus, including P. hypnophilusand Crepidotus herbarum under C. epibryus. Crepidotus epibryus has not been reported from Australia. The other species included in Pleurotellus by Singer (1986), P. chioneus, which does occur in Australia, is morphologically close to C. epibryus, the most significant difference being the broader spores (Moser, 1983). Therefore, it is likely that Pleurotellus chioneus also belongs in Crepidotus, and it is included in that genus in the key, although no formal transfer has yet been made.
Molecular data (Moncalvo et al., 2002; Aime et al., 2005) confirm that Pleurotellus hypnophilus belongs in Crepidotus. Aime et al. (2005) also show that the type of Melanomphalia (M. nigresecens from Europe) is closely related to several species of Omphalina sens. lat. [which are more correctly placed in Lichenomphalia and Arrhenia, following Redhead et al. (2002a)]. However, at least one other species of Melanomphalia (M. thermophilus) belongs in Crepidotus (Aime et al., 2005). Moncalvo et al. (2002) and Aime et al. (2005) found Crepidotus to be monophyletic, but there was not a separate clade for each of the smooth- and rough-spored species. Therefore, the two sections recognised by Singer (1986) are not monophyletic. For the purposes of identification, we recognise Crepidotus (smooth spores) and Crepidotus (rough spores). At present, centrally stipitate species of Crepidotus (such as would formerly have been placed in Melanomphalia) are not known from Australia.
The six species of Simocybe included in the molecular analysis of Moncalvo et al. (2002), presumably all from the Northern Hemisphere, form a separate clade (albeit with relatively low statistical support) that is sister to Crepidotus. Aime et al. (2005) also found that a subset of the same species of Simocybe fell in a weakly supported clade sister to Crepidotus.
On morphology, Singer (1986) placed Pleuroflammula in the Strophariaceae. Moncalvo et al. (2002) recovered two species of Pleuroflammula in a separate clade /pleuroflammula which did not fall within the Strophariaceae but was sister to a clade containing Crepidotaceae but also Inocybaceae. The multilocus study of Matheny et al. (2007a) placed one species of Pleuroflammula within the Crepidotaceae.
Cyphellaceae
The agaricoid genus Cheimonophyllum (so far represented in Australia by Cheimonophyllum candidissimum) is accepted as circumscribed by Singer (1986), who placed it in the tribe Collybieae of the Tricholomataceae. However, on molecular evidence it falls in a clade with cyphelloid genera, such as Cyphella, which lack lamellae (Matheny et al., 2007a).
Entolomataceae
Singer (1986) considered Entoloma (synonym Rhodophyllus) in a broad sense for species with spores that are angular in all views, irrespective of fruit-body morphology, thus including various segregates adopted by Pegler (1983c) and other authors, such as Alboleptonia, Eccilia, Claudopus, Inocephalus, Inopilus, Leptonia , Nolanea and Pouzarella. On molecular evidence (Moncalvo et al., 2002; Co-David et al., 2009) many of the segregates are polyphyletic or their recognition would render other genera paraphyletic. We follow Co-David et al. (2009) in recognising Entoloma in a broad sense. Due to lacking or having a lateral stipe, Entoloma section Claudopus is keyed out separately in FunKey, with remaining members of the genus included under Entoloma (other).
The other two genera of this family, Clitopilus and Rhodocybe are well established on the basis of spore morphology (Singer, 1986). However, multi-locus molecular analyses (Co-David et al., 2009) show that there is a clade for Clitopilus nesting within one containing species of Rhodocybe. Consequently the two genera must be united under Clitopilus, the older name. Due to the distinctive spore morphology of the two taxa, we key out Clitopilus morphogroup Rhodocybe (spores verrucose) and Clitopilus other (spores longitudinally ridged). Based on multi-locus data (Matheny et al., 2007a; Co-David et al., 2009) Entoloma (and segregates) and Clitopilus (including Rhodocybe) both fall within a well-supported Entolomataceae clade.
Fayodiaceae
Conchomyces is accepted as an independent genus for the echinulate-spored Conchomyces bursiformis, following Horak (1981b), although it has been placed in Hohenbuehelia by Reid (1963) and Singer (1986: in subgenus Reidia). On the basis of nLSU data (Moncalvo et al., 2002), Conchomcyes falls within a clade (designated as /fayodioid) containing mostly rough-spored agarics, such as the genera Fayodia, Gamundia and Myxomphalia. Singer (1986) treated all three under Fayodia, which he placed in the tribe Myceneae of the Tricholomataceae, whereas he had placed Conchomyces (under Hohenbuehelia) in the tribe Resupinateae. No members of the /fayodioid clade were sampled by Matheny et al. (2007a) and so the position of this clade in relation to other families of Agaricales remains uncertain, but it is reasonable to suggest it is likely to warrant family rank, in which case the name Fayodiaceae is available.
Hydnangiaceae
Laccaria is quite distinct morphologically and was considered by Singer (1986) to be the sole member of the tribe Laccariinae of the Tricholomataceae. On multilocus molecular data the genus falls within a lineage recognised as the family Hydnangiaceae, which also including the truffle-like Hydnangium (Matheny et al., 2007a).
Hygrophoraceae
The Hygrophoraceae was for a long time considered to be distinct from the Tricholomataceae and limited to Hygrophorus and Hygrocybe and a few close relatives, all characterised by the relatively thick lamellae and elongate basidia (Singer 1986; Young 2005a), although Young (1997d) emphasised the waxy appearance of the lamellae as being diagnostic and pointed out that the basidia were no longer than those in other families. Multilocus molecular data (Matheny et al., 2007a) demonstrate that several omphalinoid genera treated by Singer (1986) in the Tricholomataceae are referable to the Hygrophoraceae, in particular Lichenomphalia. Matheny et al. (2007a) did not include species of Arrhenia in their study, but because that genus is the sister taxon to Lichenomphalia on molecular data (Moncalvo et al., 2002; Redhead et al., 2002a), and the combined clade of species of these two taxa is sister to other genera placed by Matheny et al. (2007a) in the Hygrophoraceae, it too must be considered a member of this family. Omphalina sens. strict. (with O. pyxidata as type) and Gerronema sens. strict. (at least in the sense of the two species for which sequence data are available, G. strombodes and G. subclavatum) could also belong in the Hygrophoraceae, but these taxa were not sampled by Matheny et al. (2007a), and in the cladogram of Redhead et al. (2002a) they are basal to the other Hygrophoraceae, although the taxon sampling did not include other closely related families. Neither Omphalina sens. strict. nor Gerronema have been confirmed from Australia (see Excluded taxa).
Generic limits for the taxa traditionally assigned to the Hygrophoraceae follow Young (2005a) who accepts Humidicutis, Hygrocybe (including Bertrandia) and Hygrophorus (with one Australian species, Hygrophorus involutus). Young (2005a) also includes Camarophyllopsis in the Hygrophoraceae, but that genus belongs in the Clavariaceae. Other arrangements generally accept Humidicutis, Hygrocybe and Hygrophorus, but sometimes at different rank. Singer (1986) treated Camarophyllus as a genus; Young (2005a) included it under Hygrocybe, as subgenus Cuphophyllus. The genus Gliophorus was recognised as a segregate from Hygrocybe by Horak (1990) due to the glutinous pileus and stipe. Species placed in Gliophorus by Horak (1990) were treated in Hygrocybe subgenus Pseudohygrocybe by Young (2005a).
Multilocus molecular data support the inclusion of Humidicutis, Hygrocybe and Hygrophorus in the Hygrophoraceae (Matheny et al., 2007a). These data also support the delimitation of these genera, except that subgenus Cuphophyllus should be segregated from Hygrocybe and accepted as a distinct genus (Camarophyllus). It was also suggested that there might be other segregates from Hygrocybe, such as Gliophorus (although only one species of this genus was sampled) (Matheny et al., 2007a). For the moment (due to the lack of necessary combinations in some genera), we retain the generic delimitation used by Young (2005a) in his comprehensive treatment of the Australian Hygrophoraceae. Within Hygrocybe, due to the large number of species (more than 70), the three subgenera accepted by Young (2005a) are keyed out separately: Hygrocybe subgenus Cuphophyllus, Hygrocybe subgenus Hygrocybe and Hygrocybe subgenus Pseudohygrocybe.
Hygrocybe subgenus Pseudohygrocybe is characterised by the regular to subregular lamellar trama, composed of relatively short elements, in contrast to the very long (more than 1000 µm) elements of subgenus Hygrocybe, and the irregular trama of subgenus Cuphophyllus. Hygrocybe kandora has the long elements in the lamellar trama typical of subgenus Hygrocybe, but in FunKey it is keyed out in subgenus Pseudohygrocybe because it shows certain character states in common with other species of the latter (especially the subsquamulose pileus surface, the lamellae that can have a decurrent tooth and the pileipellis being a trichoderm); these character states are otherwise lacking in subgenus Hygrocybe (at least the Australian species). For Hygrocybe lilaceolamellata Young (2005a) observed that 'future studies might result in its transfer to Hygrophorus, due to its weakly divergent lamellar trama'.
The nomenclature and generic delimitation of Omphalina sens. lat. and its segregate genera are complex (see also Gerronema and Omphalina under Excluded taxa). Delimitation of Arrhenia and Lichenomphalia, the two omphalinoid genera of the Hygrophoraceae in Australia, has been clarified by Redhead et al. (2002a), and the latter is accepted for lichenised omphalinoid agarics. The names Botrydina and Phytoconis have also been utilised for Lichenomphalia, but both are typified by anamorphs and cannot be used for the teleomorph (basidia-producing) stage (Redhead et al., 2002a). Arrhenia is non-lichenised and ranges from agaricoid (centrally stipitate and lamellate) to having cyphelloid or nutant fruit-bodies with a smooth or vein-like hymenium (the morphology of the type of the genus). Singer (1986) placed species of Lichenomphalia (such as the Northern Hemisphere L. alpinum and L. hudsonii) mostly in Gerronema section Romagnesia subsection Venustissima, but he also included in this subsection species now accepted in Loreleia (Hymenochaetales). Singer (1986) placed agaricoid members of Arrhenia (such as the Northern Hemisphere A. epichysium and A. griseopallida) in Omphalina section Omphalina (which also contained the quite distinct O. pyxidata, which is the type of Omphalina) and in Leptoglossum, while restricting Arrhenia to A. auriscalpium, a species with a merulioid (vein-like) hymenium.
Hymenogastraceae
Phaeocollybia and Hebeloma are well circumscribed morphologically (Singer, 1986; Rees & Wood, 1996). Hebeloma victoriense is keyed out separately here because of its unusual deep reddish brown spore print and the well-defined annulus. Holland & Pegler (1983) suggested that while the spore print colour is consistent with Hebeloma subgenus Porphyrospora, the other characters, especially the annulus, are more typical of other subgenera or sections. Other Australian species of Hebeloma are included under Hebeloma (other).
Phaeocollybia and Hebeloma were formerly placed in the Cortinariaceae by Singer (1986), while Kühner (1980) included the former in the Strophariaceae and the latter in the Cortinariaceae. The Northern Hemisphere species of the two genera included in the molecular analysis of Moncalvo et al. (2002) fell in a separate monophyletic clade for each genus, but the position of the two genera in relation to others was not resolved. Two Australian collections of Hebeloma and one of Phaeocollybia clustered with Northern Hemisphere representatives of these respective genera on the basis of the nLSU sequences analysed by Rees et al. (2003). The multilocus study of Matheny et al. (2007a) placed Phaeocollybia and Hebeloma in the Hymenogastraceae, along with several other genera, including Galerina. The /psychedelia clade of Psilocybe (which is now known as Psilocybe) also belongs in the Hymenogastraceae while the remaining species of Psilocybe belong in Deconica which is in the Strophariaceae (see this family for further discussion). Due the recent split of Psilocybe into two genera in separate families, Australian species are all keyed out under Psilocybe. The Hymenogastraceae is the sister taxon to the Strophariaceae.
Singer (1986) circumscribed Galerina to include species with a rather diverse range of spore morphology and placed the genus in the Cortinariaceae, although Kühner (1980) maintained that it belonged in the Strophariaceae. Singer (1986) reserved subgenus Tubariopsis for species with verrucose spores lacking a plage, and generally lacking clamp connections and pleurocystidia. Within the other subgenus, Galerina, Singer (1986) distinguished nine sections, including Inocyboides (cystidia thick-walled), Porospora (spores smooth with a narrow germ pore) and Pseuodotubaria (spores smooth, lacking a germ pore). Species fo the remaining sections (Calyptrospora, Galerina, Naucoriopsis, Mycenopsis and Physocystis) usually had verrucose spores with a plage. Section Porospora was recognised by some authors as the genus Phaeogalera (with the type P. stagnina).
Moncalvo et al. (2002), Rees et al. (2002; 2003) and Walther et al. (2005) found on molecular data that the few Galerina species sampled fell in different places in their phylogenetic trees and Gulden et al. (2005), using molecular data from 36 Northern Hemisphere species of Galerina, confirmed that the genus is polyphyletic. Most species occur in four clades, designated as /mycenopsis, /naucoriopsis, /galerina and /tubariopsis consisting typically of species from sections Mycenopsis, Naucoriopsis, Galerina and Tubariopsis respectively. However, while there is a typical macro- and micro-morphology for each clade, there are exceptions, and some species from other sections are also present in each of the four clades. In addition, several species of Galerina fall outside the four main clades; these include Galerina nana (from section Inocyboides) and G. pumila (from section Mycenopsis).
Gulden et al. (2005) found that the three Galerina clades /naucoriopsis, /galerina and /tubariopsis were interspersed by species from other genera, such as Agrocybe, Hypholoma, Phaeocollybia and Pholiota, although many of the relationships between those other genera and the Galerina clades lacked statistical support. In their multilocus study Matheny et al. (2007a) included one species from each of /galerina, /naucoriopsis and /tubariopsis. These fell together in a clade with one species of Phaeocollybia, but there was no statistical support for a single clade for the three Galerina species. Interestingly, members of the genera that were interspersed among the three Galerina clades in the nLSU tree of Gulden et al. (2005) all fell outside of the Galerina plus Phaeocollybia clade in the multilocus tree of Matheny et al. (2007a). The fourth clade identified by Gulden et al. (2005), /mycenopsis, was quite separate from the large (unsupported) clade containing the other three Galerina clades. The /mycenopsis clade fell within a strongly supported clade within which it was sister to a clade consisting of a monophyletic Gymnopilus together with Galerina paludosa (from section Mycenopsis).
Sequence data are available for only three Southern Hemisphere species of Galerina. The Australian G. subcerina [placed by Wood (2001) in section Calyptrospora on morphological grounds] was found by Rees et al. (2003) to form a well-supported clade with the Northern Hemisphere G. paludosa, a sister to Gymnopilus in the cladogram of Gulden et al. (2005). Rees et al. (2002) found that G. subcerina and another Australian species, G. tibiformis [placed by Wood (2001) on morphology in section Mycenopsis], clustered together in some analyses, but among some Northern Hemisphere species in others. Galerina tibiformis was found by Rees et al. (2003) to cluster with a species of Inocybe. This is surprising, since all species of Galerina sampled by Gulden et al. (2005) were distant from the two species of Inocybe included in their phylogenetic tree.
While further sampling of Galerina with multilocus data are required before precise delimitation of any segregate genera and their relationships can be determined, the possibility remains that the bulk of Galerina could be monophyletic. In particular, an integrated analysis of a broad sample of Southern Hemisphere Galerina and Gymnopilus would be informative in confirming the boundaries between these two genera, and indicating the placement of austral Galerina species in the four main clades identified among Northern Hemisphere Galerina.
For the purposes of identification, we separate species of Galerina into four groups. Galerina nana, with its distinctive metuloid cystidia, is keyed out separately. The other three groups are differentiated on spore morphology. Galerina (rough spores, no plage) contains two species placed by Wood (2001) in section Mycenopsis, that differ from other species in the section by the lack of a plage. Galerina (smooth spores) includes four species placed by Wood (2001) in section Porospora along with the smooth-spored G. subpumila from section Mycenopsis. The remaining species (all with verrucose spores and a plage) are keyed out as Galerina (other). The latter consists of species placed by Wood (2001) in sections Calyptrospora, Galerina, Mycenopsis, Naucoriopsis and Physocystis.
Inocybaceae
The family Inocybaceae contains only Inocybe (see below for Auritella), and is sister to the Crepidotaceae (Matheny et al., 2007a).
According to Singer (1986), Inocybe contained species with smooth, rounded spores (subgenera Inosperma and Inocibium, without and with metuloids, respectively) and those with nodulose spores (subgenus Inocybe). Horak (1977b; 1979a; 1980b) treated subgenus Inocybe of Singer (1986) at generic rank, as Astrosporina, and restricted Inocybe to species with rounded spores. Singer (1986) and Horak (1979a; 1980b) had different interpretations of the type of Inocybe, which is why Singer (1986) applied that name (as subgenus) to the nodulose-spored group, and Horak (1980b) applied it (as genus) to the group with rounded spores. A cladistic analysis of morphological characters by Kuyper (1986) suggested that nodulose spores evolved more than once in Inocybe. Kuyper (1986) redefined infrageneric taxa, introducing the subgenus Mallocybe (pileus woolly to squamulose, spores rounded, hymenial cystidia arising as terminal elements of tramal hyphae, and with brown necropigment in the basidia), and recognising subgenera Inocybe (almost always with metuloid pleurocystidia) and Inosperma (lacking pleurocystidia). Species with nodulose spores, placed by Singer (1986) in subgenus Inocibium, were dispersed by Kuyper (1986) in two supersections of subgenus Inocybe.
Analysis of DNA sequences from three loci by Matheny (2005) revealed that all species traditionally treated in Inocybe in the broad sense fall within a single, well-supported clade consisting of five main clades, corresponding to Inocybe sens. strict., Mallocybe, Inosperma, and two newly introduced taxa, /auritella (sister to Mallocybe) and /pseudosperma. The /auritella clade contained Australian species, which shared with those of Mallocybe the basidial necropigment. A single taxon with affinities to Inocybe maculata was placed in a fifth lineage, /pseudosperma (which was also stated to contain some members of section Rimosae). Species with nodulose spores did not form a monophyletic group, but all fell within the Inocybe sens. strict. clade, mixed with many species with rounded spores. Species from the other four clades all had rounded spores. Matheny & Bougher (2006) formally introduced the genus Auritella for the /auritella clade, which contains Southern Hemisphere species, morphologically close to Mallocybe, but with a relatively short stipe and long cheilocystidia (except in the single sequestrate species). Pending formal adoption of names at generic rank for all five lineages within Inocybe, and the placement of local species, we recognise two groups based solely on spore outline: Inocybe (nodulose spores) and Inocybe (rounded spores), the latter including Auritella.
Lyophyllaceae
This family, as defined in the multilocus study of Matheny et al. (2007a) accords to the tribes Lyophylleae and Termitomyceteae of Singer (1986), which he placed in the Tricholomataceae.
Singer (1986) had a broad circumscription of Lyophyllum, including species placed in Tephrocybe (= Lyophyllum section Tephrophana) by authors such as Moser (1983). In an analysis of data from several DNA loci, Hofstetter et al. (2002) showed that Lyophyllum and Calocybe, as circumscribed by Singer (1986), are not monophyletic, and nor is section Tephrophana. They identified four groups within the Lyophylleae, consisting of: (1) predominantly species of Calocybe (and species of Lyophyllum considered by some authors to belong in Calocybe), but also containing the type of Lyophyllum; (2) members of the Termitomyceteae, along with some species of section Tephrophana; (3) species of Asterophora and Hypsizygus, along with some of section Tephrophana, and one species of Calocybe; and (4) a mixture of species of section Tephrophana and other sections of Lyophyllum.
The nomenclature of these clades, and their constituent groups is yet to be resolved. Australian species of Lyophyllum are very poorly known, and for the purposes of identification, three units are recognised: Lyophyllum tylicolor group, Lyophyllum anthracophilum group and Lyophyllum (other). The last identification unit contains only one named species, Lyophyllum decastes, but it is coded to allow for some of the known variation in the genus to cope with potentially undescribed Australian species of Lyophyllum sens. lat. With regard to the four clades distinguished by Hofstetter et al. (2002), Lyophyllum tylicolor falls within Clade 3B while L. anthracophilum and L. decastes fall within Clade 4A.
The other Australian genus in the Lyophylleae, Asterophora, is well characterised and represented by Asterophora mirabilis. The only debate has been about the choice between the names Asterophora and Nyctalis. Singer (1986) favoured the latter, but we follow Redhead & Seifert (2001) in using Asterophora.
Macrocystidiaceae
The generic circumscription of Macrocystidia is stable (Singer 1986). It was placed in the subtribe Omphalinae of the Tricholomataceae by Singer (1986), but multilocus molecular data indicate that it is the sole member of the Macrocystidiaceae, a family whose placement is unresolved within the five main clades of Agaricales recognised by Matheny et al. (2007a).
Marasmiaceae
Multilocus sequence data delimit the family Marasmiaceae as containing Campanella, Chaetocalathus, Crinipellis, Marasmius and Tetrapyrgos (Matheny et al., 2007a). Singer (1986) placed three of these genera in the tribe Marasmieae of the Tricholomataceae, in subtribes Marasminae (for Marasmius) and Crinipellinae (for Chaetocalathus and Crinipellis); while a third subtribe Oudemansiellinae contained Oudemansiella and several other genera now placed in other families. A number of cyphelloid genera with non-lamellate hymenium were also included in the first two subtribes. Singer (1986) placed Campanella in the tribe Collybieae along with genera now mainly belonging in the Omphalotaceae, such as Anthracophyllum and Marasmiellus. He did not recognise Tetrapyrgos at generic rank.
The only significant morphological difference between Chaetocalathus and Crinipellis is that the former has a reduced or non-existent stipe (Singer, 1986). Some tropical species of Crinipellis (C. perniciosa and allies, some anamorphic) that are plant pathogens have recently been segregated into Moniliophthora, which is not known from Australia. Molecular data are consistent with monophyly of the three genera (Aime & Phillips-Mora, 2005; Kerekes & Desjardin, 2009).
Singer (1986) distinguished the laterally-stiped Campanella from Marasmiellus (with central or lateral stipe) by the gelatinised trama of the former in association with widely-spaced and often intervenose lamellae; although one section of Marasmiellus contained pleurotoid species with gelatinised trama. Both genera were separated from Marasmius by the non-hymeniform pileipellis. Horak (1983b) emended the genus Pterospora (later called Tetrapyrgos for nomenclatural reasons) to accommodate some species placed by Singer (1986) in Campanella and Marasmiellus which had ‘tetrahedral’ spores (with a stellate or triangular profile). Species of Tetrapyrgos, with both central and lateral stipe attachment, cluster together in molecular analyses (Moncalvo et al., 2002; Aime & Phillips-Mora, 2005; Wilson & Desjardin, 2005; Thorn et al., 2006; Nakasone et al., 2009) although sometimes mixed with Campanella (see below). Marasmiellus candidus, with a rounded spore outline, clusters in the Tetrapyrgos clade in the analysis of Wilson & Desjardin (2005), but on a long branch. With greater taxon sampling of Marasmiellus and Campanella, this species falls outside the Tetrapyrgos clade (Nakasone et al., 2009).
On molecular data Campanella is polyphyletic, with most species falling in the Marasmiaceae but with C. eberhardtii in the Omphalotaceae (Wilson & Desjardin, 2005; Nakasone et al., 2009). Various relationships of Campanella to Tetrapyrgos have been recovered. A clade consisting of several collections of Campanella can be: sister to Tetrapyrgos (Moncalvo et al., 2002); separate from Tetrapyrgos but with relationship unresolved (Aime & Phillips-Mora, 2005); entirely within the Tetrapyrgos clade (Thorn et al., 2006); or forming a separate clade but with C. junghuhnii within the Tetrapyrgos clade (Nakasone et al., 2009). Spores of C. junghuhnii are not obviously triangular, but they were described by Parmasto (1981) as having an ‘indistinct bulge’ on one side. The separation of Campanella from Tetrapyrgos requires further analysis.
In FunKey we adopt Tetrapyrgos, keying out the centrally stipitate Tetrapyrgos nigripes separately to the laterally stipitate Tetrapyrgos olivaceonigra, and retain Campanella in the sense of Singer (1986) exclusive of species now in Tetrapyrgos.
In Marasmius, Singer (1986) included 12 sections, all with hymeniform pileipellis with the exception of two sections Androsacei and Fusicystides.
On the basis of molecular data, species treated by Singer (1986) in Marasmius sections Alliacei and Androsacei now belong in the Omphalotaceae, in Mycetinus and Setulipes respectively. Tan et al. (2009) [quoting 'Desjardin unpbl.'] also place section Fusicystides, for which molecular data are lacking, under Setulipes. A few other species from section Alliacei have been transferred to Rhizomarasmius which belongs in the Physalacriaceae (Moncalvo et al., 2002; Ronikier & Ronikier, 2011). At least some members of Marasmius section Epiphylli also belong in the Physalacriaceae (Walther et al., 2005; Tan et al., 2009). Note that there are many tropical species that have not yet been transferred from sections Alliacei and Androsacei to Mycetinus and Setulipes, respectively. See further discussion under Omphalotaceae.
Members of the remaining six sections of Marasmius (sections Marasmius, Globulares, Hygrometrici, Leveilleani, Neosessiles and Sicci) for which there are molecular data form a monophyletic group in the Marasmiaceae (Wilson & Desjardin, 2005; Thorn et al., 2006; Douanla-Meli & Langer, 2008; Tan et al., 2009; Wannathes et al., 2009), although some of these analyses do not include representatives of all six sections. Wilson & Desjardin (2005) also indicate that section Scotophysini (monotypic, for M. scotophysinus), for which molecular data are lacking, belongs in Marasmius sens. strict. Molecular data are also lacking for section Inaequales which contains a few species with inamyloid trama. Within the core Marasmius clade, the few monophyletic sections have relatively low taxon sampling, and the large sections Globulares and Sicci are completely intermixed (Tan et al., 2009; Wannathes et al., 2009). The subsections and series recognised by Singer (1986) are also not monophyletic (Wannathes et al., 2009).
With one exception, species of Marasmius described from Australia belong in sections with a hymeniform pileipellis, including Alliacei, Epiphylli, Marasmius, Globulares and Sicci (Pegler, 1965; Grgurinovic, 1997a). Other species reported from Australia are all from sections Globulares, Marasmius or Sicci. When describing Marasmius tasmaniensis, which lacks a hymeniform pileipellis, Singer (1989) compared it to species of section Androsacei. It is omitted from FunKey pending clarification of its generic position. It is also possible that some of the other species from Australia, from sections Alliacei and Epiphylli belong outside of Marasmius sens. strict. However, until placement in genera such as Mycetinus is supported by molecular evidence, we treat all the other Australian species under Marasmius. We key out Marasmius oreades separately from Marasmius (other), due to the relatively large size of the fruit-body and the stipe that is not black (as in many other species of the genus).
Mycenaceae
For most Mycena species, we follow the generic circumscription of Grgurinovic (2003) in her comprehensive monograph of the Australian species, which essentially follows Singer (1986), but with the addition of several new sections for unique Australian species. Exceptions to Grgurinovic's (2003) scheme are the genera Cruentomycena and Roridomyces. The former was erected by Petersen et al. (2008) for the distinctive Cruentomycena viscidocruenta, which had been recognised by Grgurinovic (2003) as the only species of Mycena section Viscidocruentae; this differs from other Mycena by the viscid pileus and stipe and smooth pileipellis hyphae. Roridomyces was introduced by Rexer (1994) for Mycena rorida and other species with a hymeniderm pileipellis; these were included in Mycena section Roridae by Grgurinovic (2003).
Most Mycena have amyloid spores (see below for exceptions among Australian species), but some extralimital sections such as Mycena section Hiemales, have non-amyloid spores, thus complicating the generic delimitation against Hemimycena, which has non-amyloid spores and hyphae, as do many species of Hydropus (Singer, 1986). The presence in Australia of mycenoid genera other than Mycena, such as Delicatula (with a veil), Hemimycena and Hydropus, has not been confirmed. However, even though it would not be surprising if species of these genera were encountered, the three genera are not included in this edition of FunKey.
Mycena (other) is the identification unit for most of the Australian agarics currently placed in Mycena, and it includes representatives of 13 sections. Six other sections are separated for identification purposes because they differ, respectively, by: non-amyloid spores (Mycena section Adonideae, which belongs in the /hydropoid clade), more robust habit, purple to lilac fruit-bodies, smooth pileipellis hyphae and sometimes non-amyloid spores (Mycena section Calodontes [this section is equivalent to section Purae of Singer (1986)]), bright orange lamellae and overall similarity to Hygrocybe (Mycena leaiana in section Caespitosae), white squamules on the pileus and smooth pileipellis hyphae (Mycena nargan in section Nargan), acanthocysts on the pileal surface (Mycena section Sacchariferae), or large, thick-walled pleurocystidia in combination with criniform stipes (Mycena cystidiosa in section Metuloidiferae). Removal of these segregates means that all species in Mycena (other) have amyloid spores and repent pileipellis hyphae that are nodulose. The amyloid reaction can be weak in some species (such as noted by Grgurinovic (2003) for M. subalbida), but this is sometimes an effect of the age of material, which in some cases can be overcome by longer soaking in Melzer's reagent.
Moncalvo et al. (2002) investigated nLSU sequences and distinguished a clade /mycenaceae containing ten species of Mycena sensu Singer (1986), mixed with species of Favolaschia, Panellus, Mycenoporella, Poromycena and Resinomycena. Walther et al. (2005), also using the nLSU, recovered a clade of 20 species of Mycena that included Panellus stypticus. In both analyses, the Mycena species do not all cluster together and there are few statistically supported subclades within the /mycenaceae clade (such as could be linked to subgenera of Mycena). An exception is that in the Moncalvo et al. (2002) analysis three members of Mycena section Calodontes cluster together. This section was consequently recognised by Redhead et al. (2001) at generic rank as Prunulus, but it seems premature to separate out only this group at generic rank, while the relationships of the other members of the /mycenaceae clade remain unresolved. Both analyses found that Mycena rorida, the non-Australian type of Roridomyces, also fell within the /mycenaceae clade, but notably Walther et al. (2005) found that this species stood out as being on a very long branch in relation to all other Mycena species.
Cruentomycena viscidocruenta (as Mycena) and M. leaiana were the only Australian representatives in the /mycenaceae clade of Moncalvo et al. (2002), and there were no representatives from a number of sections uniquely Australian or austral, such as Cyanocephalae, Maldae, Metuloidiferae and Nargan. Concordance between the sections of Mycena, such as those of Singer (1986), and phylogeny has not been fully explored. Further gene and taxon sampling is necessary, especially from the Southern Hemisphere, to clarify monophyly and relationships among the various sections of Mycena and related genera.
In terms of family placement, Singer (1986) treated Mycena and Panellus in the Tricholomataceae, the former in the tribe Myceneae, the latter in the Panelleae. In their multilocus study, Matheny et al. (2007a) recognised the family Mycenaceae, represented by five species of Mycena, in a well-supported clade in which Panellus stypticus was nested (see below). Taxon sampling was not sufficient to resolve the rank of possible segregates from Mycena, such as Prunulus.
Both Moncalvo et al. (2002) and Matheny et al. (2007a) found that representatives of section Adonideae fell well outside of the Mycenaceae [see discussion of the /hydropoid clade (below)].
Singer (1986) accepted Dictyopanus, Panellus and Tectella in the tribe Panelleae. Burdsall & Miller (1975) recognised the close relationship between the lamellate Panellus stypticus and the type of the poroid genus Dictyopanus (D. pusillus), and transferred the latter species to Panellus. Jin et al. (2001) and Moncalvo et al. (2002) confirmed, using molecular data, the close relationship of Dictyopanus and Panellus. However, Moncalvo et al. (2002) found that the /panelloid clade containing Dictyopanus and Panellus also contained two species of Resinomycena and Mycena viscidocruenta (type of Cruentomycena). The molecular phylogeny of Petersen et al. (2008) placed Cruentomycena closer to Resinomycena and Panellus than to Mycena (although few species of the latter genus were included).
Pending further taxon sampling, we recognise Dictyopanus at generic rank because its monophyly remains a possibility. However, it is clear that other species of Panellus, such as P. mitis and P. serotinus (Sarcomyxa), do not belong in the genus, because on molecular evidence they fall outside the /panelloid clade, as does Tectella (Jin et al. 2001; Moncalvo et al., 2002; Matheny et al., 2007a). Nineteenth century Australian records of Panellus mitis and Tectella patellaris are doubtful, and the species are not included in FunKey.
Panellus stypticus, the only species of Panellus sens. strict. is keyed out separately, as is Panellus ligulatus because it differs from Panellus sens. strict. by its non-amyloid spores and the presence of gloeosphex cystidia, a combination of characters that suggests a relationship with Hohenbuehelia. Molecular data are not available for this species. For the other Australian species, P. longinquus, see under Unplaced within Agaricales. 3.
Omphalotaceae
Lentinula was treated as an independent genus in tribe Collybieae of the Tricholomataceae by Pegler (1983a), although Singer (1986) recognised the same taxon as Lentinus subgenus Edodes. Molecular data (Moncalvo et al., 2002; Wilson & Desjardin, 2005; Mata et al., 2006; Matheny et al., 2007a) place Lentinula as a monophyletic group within the Omphalotaceae, distant from the Polyporaceae (where Lentinus now belongs). The genus is represented in Australia by the Lentinula edodes group. Anthracophyllum is a well-circumscribed genus (Pegler & Young, 1989). It was included by Singer (1986) in tribe Collybieae, but molecular data also place it within the Omphalotaceae (Moncalvo et al., 2002; Matheny et al., 2007a). The single species in Australia is Anthracophyllum archeri. The well-defined genus Omphalotus was placed in the Paxillaceae (Boletales) by Singer (1986). Molecular data initially placed it instead in the Agaricales (Binder et al., 1997) and later specifically in the Omphalotaceae (Moncalvo et al., 2002; Mata et al., 2006; Matheny et al., 2007a). The single species Australian species is Omphalotus nidiformis.
Collybia was placed in the tribe Collybieae of the Tricholomataceae by Singer (1986), with nine sections, including Collybia, Iocephalae, Levipedes, Striipedes, Subfumosae and Vestipedes. The genus in the sense of Singer (1986) is now known to be a highly artificial assemblage, and Collybia is now restricted to section Collybia, including the type, C. tuberosa, and a few other species. These are characterised by the white spore print, the pileipellis a cutis, and often grow on other fungi and produce sclerotia (Antonín & Noordeloos, 1997). Microcollybia is a later synonym of Collybia in this narrow sense. Collybia racemosa, which was placed alongside C. tuberosa and allied species by Antonín & Noordeloos (1997), has recently been segregated in the genus Dendrocollybia, which has the stipe covered in short outgrowths terminating in tiny sterile pilei (Hughes et al., 2001). Collybia sens. strict. belongs in the Tricholomataceae (actually within the main Clitocybe clade) and Dendrocollybia is also excluded from the Omphalotaceae, although its family placement is not yet resolved (Moncalvo et al., 2002; Matheny et al., 2007a). Collybia in the narrow sense does not occur in Australia, nor does Dendrocollybia.
Rhodocollybia has been taken up for species placed by Singer (1986) in two groups within Collybia section Striipedes: stirpes Maculata and Butyracea (Antonín & Noordeloos, 1997; Antonín et al. 1997). The genus is characterised by a pinkish spore print and spores that are cyanophilous and dextrinoid. Molecular data supported the monophyly of Rhodocollybia within the Omphalotaceae (Mata et al., 2004a, 2004b, 2006; Wilson & Desjardin, 2005). However, the recently erected genus Connopus (for Collybia acervatus, which does not occur in Australia) renders Rhodocollybia paraphyletic in Bayesian analysis of nLSU sequences; although alternative trees with Connopus sister to Rhodocollybia could not be rejected statistically (Hughes et al., 2010). Connopus, which has non-dextrinoid spores, is sister to a clade comprising members of the R. butyracea group and this clade is sister to the R. maculata group (Hughes et al., 2010). Rhodocollybia has been confirmed from Australia (with e.g. dextrinoid spores), and the genus is included in FunKey inclusive of the R. butyracea and R. maculata groups.
Gymnopus is an old name (introduced by Persoon at infrageneric level), but one that was not widely recognised until Antonín & Noordeloos (1997) and Antonín et al. (1997) restricted Collybia to the species around C. tuberosa, excluded Rhodocollybia (as above) and took up Gymnopus for species that had been placed by Singer (1986) in Collybia sections Iocephalae, Levipedes, Subfumosae and Vestipedes, and in stirps Fusipes of section Striipedes. Antonín & Noordeloos (1997) and Antonín et al. (1997) characterised Gymnopus against other collybioid agarics by the non-insititious stipe base, with basal mycelium, combined with a white to cream spore print and non-dextrinoid spores (in contrast to the pinkish spore print of Rhodocollybia) and frequently with encrusting pigment (in contrast to Collybia sens. strict.). Gymnopus as thus circumscribed includes species with a pileipellis which is a cutis or a trichoderm, made up of hyphae that have either: (1) sparse or abundant nodules, (2) short and broad branches or lobes (a 'Dryophila-structure' where the elements can resemble pieces of a jig-saw puzzle), or sometimes (3) 'coralloid' diverticulae (approaching a 'Rameales-structure', where the diverticulae are numerous and narrow and can branch several times). Species with the latter type of pileipellis seem to be predominantly tropical, and were placed by Singer (1986) in Collybia section Subfumosae.
Antonín & Noordeloos (1997) pointed out the close similarity of Gymnopus to Marasmiellus, with the 'only real difference' being that the stipe is insititious in the latter, but with a basal mycelium in the former. Singer (1973; 1986) characterised Marasmiellus against Marasmius as lacking the hymeniform pileipellis of the latter, with the pileipellis 'typically' having a 'Rameales-structure', although across the genus the structure was often 'poor or weak', and in some species the pileipellis was composed of non-diverticulate hyphae. He differentiated Marasmiellus from other members of the tribe Collybieae (especially Micromphale) by a complicated combination of characters, exemplified by his contrast of Marasmiellus against Micromphale as the former possessing 'gelatinized hyphae ... in the trama of the pileus ... only if the stipe is insititious ... and there is a Rameales-structure'. The circumscription of Marasmiellus by Singer (1986) included ten sections with a mix of centrally stipitate and pleurotoid species.
Micromphale was recognised as a separate genus by Singer (1986) due to the combination of insititious stipe and gelatinous trama (or gelatinised pileipellis). Micromphale was treated as a synonym of Marasmiellus by Antonín & Noordeloos (1997) and Antonín et al. (1997) because of the similarity of the pileipellis structure in the two genera, although they also transferred to Gymnopus some species that had been placed by Singer (1986) in Micromphale. Setulipes was created by Antonín (1987) for M. androsaceus (type of Marasmius section Androsacei), which differs from other Marasmius in the pileipellis which is not hymeniform at maturity.
Molecular data place the types and nearly all other species sampled of the four genera Gymnopus, Marasmiellus, Micromphale and Setulipes within one clade in the Omphalotaceae which also contains Lentinula and Rhodocollybia (Moncalvo et al., 2002; Mata et al., 2004b; Wilson & Desjardin, 2005 [see this work especially for placement of type species]; Walther et al., 2005). Initially, Mata et al. (2004b) transferred to Gymnopus the types of Marasmiellus (M. juniperinus), Micromphale (M. foetidum) and Steulipes (S. androsaceus, formerly Marasmius, and type of Marasmius section Androsacei), as well as a couple of other species of Marasmiellus for which molecular data were available, while indicating that their analysis showed Rhodocollybia to nest within this broadly circumscribed concept of Gymnopus. Wilson et al. (2004) found Gymnopus section Levipedes to be monophyletic but section Vestipedes was not.
With greater taxon sampling of nLSU data, Wilson & Desjardin (2005) recognised a strongly supported clade ‘A’ for nearly all sampled members of Gymnopus, Lentinula, Marasmiellus, Micromphale, Rhodocollybia and Setulipes. Within this clade, Gymnopus was polyphyletic, broadly dividing into two clades for each of (a) sections Gymnopus (i.e. Collybia section Striipedes) and Levipedes and (b) sections Vestipedes and Iocephalae; along with strongly supported clades for Lentinula and Rhodocollybia. Recognition of Gymnopus in a broad sense would have required sinking of these other morphologically distinct genera. Therefore, Wilson & Desjardin (2005) recommended acceptance of the two clades for Gymnopus at generic rank. Gymnopus in the strict sense included the type of this genus (G. fusipes) along with types of Setulipes and Micromphale; an arrangement accepted by Noordeloos & Antonín (2008). The second Gymnopus clade (‘b’ above) included the type of Marasmiellus and therefore technically it should be called Marasmiellus. However, Wilson & Desjardin (2005) refrained from making the transfers to Marasmiellus because they considered that greater taxon sampling was required before redefining Marasmiellus. In addition, the Gymnopus ‘b’ clade recovered by Wilson & Desjardin (2005) that contained the type of Marasmiellus did not have strong statistical support, and indeed, in the strict consensus tree of Hughes et al. (2010), members of this second clade fell in four clades in a polytomy that also included Rhodocollybia and a single branch for Marasmiellus ramealis. Furthermore, Wilson & Desjardin (2005) found that a few species of Gymnopus and Marasmiellus fall outside of the two main Gymnopus clades; notably Marasmiellus ramealis is sister to a species of Campanella and M. palmivorus clusters in the Maramsiaceae. In addition, Matheny et al. (2007a) found Gymnopus contrarius to cluster at the base of the Omphalotus+Anthracophyllum clade, rather than with other Gymnopus. Using ITS data, Mata et al. (2006) recovered a wide range of Gymnopus species not interspersed by Lentinula or Rhodocollybia in three main clades, corresponding to the two groups of Wilson & Desjardin (2005) except that Gymnopus sens. strict. occupied two clades, a small clade containing the types of Gymnopus and Setulipes and a larger one for the other species. Further sampling of this group with multilocus data will assist in clarifying generic boundaries.
Compared to Gymnopus, Marasmiellus is poorly sampled in molecular analyses, as is Setulipes (i.e. Marasmius section Androsaceus). Therefore, it cannot be assumed that all species of these two groups will belong in the same genus as their type specimens. This is especially true for Marasmiellus, which according to Singer (1986) contained ten sections, representatives of only four of which were included by Wilson & Desjardin (2005), and then by one or only a few species that fell in several different places in their tree, including outside the Gymnopus clade that contained the type of Marasmiellus.
Within their clade ‘A’, Wilson & Desjardin (2005) also identified a distinct clade containing members of Marasmius section Alliacei (with a hymeniform pileipellis) and Marasmiellus opacus for which they resurrected the genus Mycetinus. Desjardin et al. (1993) had already showed that the pileipellis of Marasmiellus opacus was unusual because it was subhymeniform when young, and remained so at the disc in mature fruit-bodies. Mycetinus species often have a garlic-like odour, although strong odours are also present in other lineages of clade ‘A’. Species placed in Mycetinus were also recovered as monophyletic by the analyses of Mata et al. (2004b; 2006), Hughes et al. (2010) and Ronikier & Ronikier (2011) and the genus was accepted by Noordeloos & Antonín (2008).
As regards Australian species of this group, some have been reported under Gymnopus, Micromphale and Marasmiellus. In addition, many remain in Collybia (e.g. Grgurinovic, 1997) but they are most likely referable to Gymnopus, being excluded from Rhodocollybia by their non-dextrinoid spores. Molecular data are lacking for all Australian taxa, as is detailed information on pileipellis structure for some species. In FunKey, we key out Gymnopus in a broad sense [equivalent to both major clades of Wilson & Desjardin (2005)] inclusive of Australian species placed in Micromphale and Marasmiellus as well as species of Collybia not evidently belonging in Rhodocollybia. The exception is the very distinctive Marasmiellus affixus which is keyed out separately due to its symbiotic growth with an algal mat and the strong odour.
Panaeolaceae
Singer (1986) recognised the genera Panaeolus, Anellaria, Copelandia and Panaeolina in the subfamily Panaeoloideae of the Coprinaceae. Panaeolina was distinguished from Panaeolus by the ornamented spores. Copelandia differed by having blue-staining fruit-bodies and the presence of metuloids, while Anellaria was characterised by the fleshy, non-hygrophanous, viscid pileus in combination with chrysocystidia-like pleurocystidia (differing from true chrysocystidia by not yellowing in ammonia solution). Singer (1986) pointed out that the Anellaria type of pleurocystidium [termed sulphidia by Gerhardt (1996) from the positive reaction with sulphovanillin] were also present in some species of Panaeolus. Gerhardt (1996) accepted Panaeolus and Panaeolina on morphological grounds, the former including three subgenera (Panaeolus, Anellaria and Copelandia). Within subgenus Panaeolus, one section (Verrucispora) contained species with very fine ornamentation.
Molecular data consistently recover a single clade encompassing Panaeolus (including Anellaria and Copelandia) and Panaeolina (so far represented only by P. foenisecii), but relationships among taxa vary. Using nLSU, Moncalvo et al. (2002) recovered Anellaria semiovata (as Panaeolus) embedded within the Panaeolus clade, while in analyses of the same region, Panaeolina is sister to the main Panaeolus clade (Moncalvo et al., 2002; Walther et al. 2005). In contrast, Garnica et al. (2007), with the addition of data on RPB1, recovered a clade with Anellaria semiovata basal to a supported clade of Panaeolina foenisecii and Panaeolus acuminatus. Copelandia cyanescens is at the base of the Panaeolus clade (Moncalvo et al., 2002) or not (Maruyama et al., 2003). Panaeolus olivaceus, the type of section Verrucispora, falls among other species of Panaeolus (Walther et al., 2005).
Further analyses of molecular data, especially with increased taxon sampling, are required to confirm the circumscription of the genera of the Panaeoleae. For the moment, the morphological classification of Gerhardt (1996) is accepted because it remains consistent with most molecular phylogenies. We key out Panaeolina foenisecii, Panaeolus (other) (including Anellaria) and also Panaeolus cyanescens, the last having distinctly ornamented spores.
The position of Panaeolus + Panaeolina [called tribe Panaeoleae by Matheny et al. (2007a)] varies with respect to other higher taxa. In analyses of the nLSU alone, Panaeoleae has support as sister to the Bolbitiaceae (Walther et al., 2005), or is placed next to Bolbitiaceae without support (Moncalvo et al., 2002). However, in analyses of other DNA regions, the Panaeoleae is not sister to Bolbitiaceae. Firstly, in the nLSU + RPB1 analysis of Garnica et al. (2007) the three species of Panaeoleae sampled form a supported clade that is sister to the Psathyrellaceae. Secondly, in the six-gene tree of Matheny et al. (2007a), the Panaeoleae falls within a supported clade also containing Tubariaceae and Crepidotaceae (the exact relationship not being resolved), with the Gymnopileae as an intervening clade between these and other families, and the Bolbitiaceae (and distant to the Psathyrellaceae). Therefore, we do not accept Panaeoleae as belonging to the Bolbitiaceae, but rather as an independent taxon that should be recognised at family rank as Panaeolaceae [an invalidly published name that requires validation].
Panaeolopsis (molecular data lacking) is included in the Agaricaceae in the Dictionary of the Fungi, but its morphological similarity to Panaeolus (lenticular-mitriform spores with distinct germ pore and cheilocystidia present) means that it is more likely to belong in the Panaeolaceae. The only species known from Australia is Panaeolopsis nirimbii.
Physalacriaceae
Four Australian agaric genera, Armillaria, Cyptotrama, Flammulina and Oudemansiella, belong in the family Physalacriaceae, along with some genera that do not form agaricoid fruit-bodies, such as Physalacria. The family is well-supported in the multilocus study of Matheny et al. (2007a), and the same clade was recovered in the Moncalvo et al. (2002) phylogeny based on nLSU sequences. The four genera had been placed by Singer (1986) in various subunits of the Tricholomataceae: Armillaria in the subtribe Omphalinae, Cyptotrama and Flammulina in the tribe Pseudohiatuleae, and Oudemansiella in the subtribe Oudemansiellinae. Other genera of the Physalacriaceae were formerly placed elsewhere in the Tricholomataceae. Indeed, the placement of these genera in various parts of the Tricholomataceae (Singer, 1986) exemplifies the common disparity between suprageneric classifications based solely on morphology with phylogenetic reconstructions based on molecular data. However, in this instance, the delimitation of the genera themselves has changed little or not at all with the addition of molecular data.
Armillaria has been rather stable in its circumscription, although Singer (1986) used the name Armillariella for it, and applied the name Armillaria to Floccularia. Cyptotrama aspratum, the only Australian species, has been placed in 14 different genera, although the genus Cyptotrama is now well differentiated (Redhead & Ginns, 1980; Singer, 1986). Flammulina is also a well-defined genus (Singer, 1986) and, pending revision, all Australian records are treated in FunKey under Flammulina velutipes.
Dörfelt (e.g. 1983, 1984) and Petersen & Methven (1994) accepted the genus Xerula for species with no partial veil and a pseudorrhiza, and reserved Oudemansiella for species with a partial veil and annulus and growing directly on wood. However, Singer (1986) and Pegler & Young (1987) treated Xerula as a subgenus of Oudemansiella. In a comprehensive revision of the ‘Xerula/Oudemansiella complex’, Petersen & Hughes (2010) accepted eight genera based on morphological characters. In addition to Xerula and Oudemansiella, they recognise Dactylosporina, Hymenopellis (for Oudemansiella radicata and allies), Mucidula (for O. mucida and allies), Ponticulomyces, Paraxerula and Protoxerula. In addition, Petersen & Hughes (2010) carried out phylogenetic analyses of an extensive set of molecular data and found that some of the accepted genera lacked statistical support or were polyphyletic (in particular, Hymenopellis), as pointed out by Vellinga (2010). A classification consistent with molecular phylogeny, as adopted by Yang et al. (2009) and discussed by Vellinga (2010), is to accept Xerula sens. strict. (for X. pudens and allies), Paraxerula and a broad Oudemansiella (including all the other genera). This arrangement is also consistent with the nLSU analysis of Walther et al. (2005) where Xerula pudens is in a clade with several other genera of Physalacriaceae, rather than clustering directly with the clade consisting of Xerula radicata and Oudemansiella mucida. In FunKey we use Oudemansiella in this broad sense including Hymenopellis and Protoxerula (sensu Petersen & Hughes, 2010), both of which are represented in Australia. Xerula in the restricted sense of Yang et al. (2009) does not occur in Australia, although many Australian species have been placed in the genus. In FunKey, most species are keyed out under Oudemansiella other, for species with fruit-bodies growing on the ground (probably attached to buried wood) which usually have a distinct pseudorhiza. Oudemansiella exannulata is keyed out separately, because it produces fruit-bodies directly on exposed wood and has a partial veil when young.
Pleurotaceae
In the 19th century, Pleurotus was a repository for all pale-spored pleurotoid agarics (with the stipe lateral or absent). The circumscription was narrowed in the middle of the 20th century with the removal of many species to genera such as Conchomyces, Hohenbuehelia, Omphalotus and Resupinatus. However, Singer (1986) retained a somewhat broad concept of Pleurotus, and included in section Lentodiellum species that Corner (1981) and Pegler (1983b) had placed in Lentinus or Panus. Molecular data confirm that these latter species belong in Panus (Hibbett & Vilgalys, 1993). Species from the remaining sections of Pleurotus accepted by Singer (1986), including section Tuber-regium, form a monophyletic group in molecular analyses (Moncalvo et al., 2002). In addition to the main entry for Pleurotus (other), three species are keyed out separately. Pleurotus tuber-regium from section Tuber-regium of Singer (1986) had been placed in Lentinus or Panus by some authors, but has been confirmed as a Pleurotus based on both the molecular data and also the production of nematode toxin on secretory processes (Hibbett & Thorn 1994; Thorn et al. 2000; Moncalvo et al., 2002). Pleurotus giganteus is unique in having a radiate lamellar trama and a partial veil. It was previously placed in Panus (Corner, 1981) or Lentinus (Pegler, 1983b), where it was aberrant due to the veil, the rather short, ellipsoid spores and the presence of capitate cheilocystidia, but molecular data (Karunarathna et al., 2011) confirm its placement in Pleurotus. Pleurotus eugrammus is distinctive because of its luminosity. It is the type of Nothopanus, and this genus, in the strict sense of the type collection of the type species (Lentinus eugrammus), and as interpreted by Corner (1981), is a synonym of Pleurotus (Petersen & Krisai-Greilhuber, 1999). This placement was adopted by Singer (1986). Note, however, that Singer (1944) misapplied Nothopanus eugrammus for a distinct taxon, for which the genus Neonothopanus has been introduced (Petersen & Krisai-Greilhuber, 1999). Note also that the material of Nothopanus (RV PR-27), which Thorn et al. (2000) found to be distinct from Pleurotus on molecular evidence and the lack of nematode toxic secretions, was re-identified as Neonothopanus nambi by Moncalvo et al. (2002).
Resupinatus and Hohenbuehelia are distinct morphologically, gloeosphex cystidia being found only in the latter (Thorn & Barron, 1986; Singer, 1986). Molecular data (Moncalvo et al., 2002) support their separation, but also indicate that various cyphelloid and poroid species should be added to Resupinatus (Moncalvo et al., 2002; Thorn et al., 2006).
Singer (1986) placed Pleurotus in the tribe Lentineae of the Polyporaceae, alongside Lentinus and Panus, but included Hohenbuehelia and Resupinatus in the tribe Resupinatae. Earlier molecular studies recovered a relationship between Hohenbuehelia and Pleurotus, but the /resupinatus clade did not fall within the Pleurotaceae (Moncalvo et al., 2002). Multilocus molecular data group Hohenbuehelia, Pleurotus and Resupinatus together, in the Pleurotaceae (Matheny et al. (2007a).
Pluteaceae
Melanoleuca, Pluteus and Volvariella are well-delimited morphologically (Singer, 1986), and there is also support for these genera on molecular evidence (Moncalvo et al., 2002). Singer (1986) included Pluteus and Volvariella in the Amanitaceae, but placed Melanoleuca in the Tricholomataceae alongside Leucopaxillus, which also has spores with amyloid ornamentation. However, on the basis of multilocus molecular evidence, Melanoleuca, Pluteus and Volvariella (represented by V. gloiocephala) all belong in the Pluteaceae, which is a separate clade to the Amanitaceae, and also distant to the Tricholomataceae (Matheny et al., 2007a). A similar relationship was found by Moncalvo et al. (2002) on nLSU data, except that Volvariella (represented by two other species) did not cluster with Melanoleuca and Pluteus, but was the sister taxon to the Schizophyllaceae.
NOTE ADDED IN PROOF: Volvariella has been found to be polyphyletic, on the basis of multi-locus data (Justo et al., 2011), and the type and most other species do not belong in the Pluteaceae. Their family placement requires further taxon sampling, but on this latest data, Volvariella sens. strict. is most closely related to Cantharocybe and a species of Camarophyllus. A few species of Volvariella, including V. gloiocephala, remain in the Pluteaceae. The new genus Volvopluteus has been erected for these species.
Psathyrellaceae
The genus Coprinus was formerly regarded as a distinct genus of the Psathyrellaceae on the basis of the deliquescence of the fruit-body, although this character was not well-developed in a few species such as Coprinus disseminatus (Singer, 1986). The classification of Coprinus was re-organised by Redhead et al. (2001) and the genus restricted to the type (Coprinus comatus) and a few other species. Coprinus in the strict sense has been transferred to the Agaricaceae. Most of the numerous other species formerly placed in Coprinus were dispersed in three genera of the Psathyrellaceae: Coprinellus, Coprinopsisand Parasola. Each of the three genera has some distinguishing morphological characteristics, and, at the time of their creation by Redhead et al. (2001), all appeared to be supported by molecular data, although Psathyrella was under-sampled in comparison to the coprinoid genera.
Subsequently, a number of studies using molecular data found the segregate genera from Coprinus (Coprinellus, Coprinopsis and Parasola) to be monophyletic, albeit in some cases with weak statistical support and low taxon sampling, but suggested that Psathyrella was not monophyletic (Moncalvo et al., 2002; Walther et al., 2005; Matheny et al., 2007a). Recent studies including a broader range of Psathyrella species, as well as additional coprinoid species, have all confirmed that (1) Psathyrella is polyphyletic, (2) internal phylogenetic structure within Psathyrella sens. lat. does not correspond well to previous morphology-based infrageneric classifications, and (3) deliquescence has been gained and/or lost several times in the Psathyrellaceae (Padamsee et al., 2008; Vasutova et al., 2008; Larsson & Orstadius, 2008; Nagy et al., 2009, 2010, 2011). The three coprinoid genera Coprinellus, Coprinopsis and Parasola are intermixed with several different clades comprised of species of Psathyrella. The genus Cystoagaricus also falls within the broad Psathyrellaceae clade. The relative positions of units within Psathyrellaceae varies according to different studies, utilising different gene regions and different taxon sampling. For example, Padamsee et al. (2008) and Vasutova et al. (2008) recovered Coprinellus within a large clade otherwise containing species of Psathyrella, including the type, P. gracilis; while Nagy et al. (2010) found Coprinellus as sister to a clade of several groups of Psathyrella, including the type section; but Larsson & Orstadius (2008) [at least in some analyses] and Nagy et al. (2011) recovered two separate clades for Coprinellus, both of which were sister to species of Psathyrella. Differences in the number of genes analysed and the methods of analysis and in taxon sampling no doubt contribute to differences in the recovered phylogenies.
Whatever the details, either (1) a very broad Psathyrella will have to be recognised, inclusive of all three coprinoid genera and also of Cystoagaricus and Lacrymaria or (2) if the three coprinoid genera, Coprinellus, Coprinopsis and Parasola, are to be maintained, then species of Psathyrella will need to be distributed among several novel genera, and Psathyrella sens. strict. restricted in scope. This would entail introduction of a number of new genera to deal with the phylogenetic structure within the Psathyrellaceae, because not all of the clades have existing names at generic rank.
On molecular data, all three coprinoid genera now contain a few non-deliquescent species, some long-accepted as coprinoid (such as Coprinellus impatiens), others recently transferred from Psathyrella, such as Parasola conopilus (Larsson & Orstadius, 2008). Once such additions are taken into account, Parasola is monophyletic (Nagy et al., 2009). The broader Coprinopsis clade contains several species of Psathyrella, such as P. subatrata, that remain in that genus (Padamsee et al., 2008). However, monophyly of Coprinopsis exclusive of these species of Psathyrella cannot be rejected (Padamsee et al., 2008) and whether to transfer such species to Coprinopsis has not been resolved. Coprinellus is monophyletic in some studies (Padamsee et al., 2008; Vasutova et al., 2008; Nagy et al., 2010) but not in others (Larsson & Orstadius, 2008; Nagy et al., 2011).
Lacrymaria, with ornamented spores, has been treated as a subgenus of Psathyrella (Singer, 1986) or at generic rank (Watling, 1979a). Molecular studies show that Lacrymaria falls well within Psathyrella sens. lat. (Moncalvo et al., 2002; Walther et al., 2005; Matheny et al., 2007a; Padamsee et al., 2008; Vasutova et al., 2008; Larsson & Orstadius, 2008; Nagy et al., 2010, 2011) as a distinct clade but clustering with some smooth-spored Psathyrella species such as P. spadicea. Whether to add such species to Lacrymaria has not been resolved.
Given the uncertainty in the classification of the Psathyrellaceae, we recognise the genera Psathyrella, Coprinellus, Coprinopsis and Parasola, as distinguished on morphological grounds by Redhead et al. (2001) and with addition of a few non-deliquescent species to the latter three genera following Padamsee et al. (2008) and Nagy et al. (2009, 2010, 2011). We also recognise Lacrymaria at generic level, following Redhead et al. (2001) and Padamsee et al. (2008).
The Coprinus cordisporus group (containing also C. patouillardii), although deliquescent, is kept separate from Coprinus sens. strict. and from the three coprinoid genera (Coprinellus, Coprinopsis and Parasola) for the moment because on molecular data it does not fall within any of the coprinoid genera, rather clustering with Psathyrella species (non deliquescent). A position within Coprinellus could not be ruled out statistically by Padamsee et al. (2008), but Nagy et al. (2010) found that ‘the hypothesis that C. patouillardii belongs to Coprinellus could be rejected’.
Schizophyllaceae
Schizophyllum (represented in Australia by Schizophyllum commune) belongs to an independent lineage that also includes poroid and cyphelloid genera such as Fistulina and Porodisculus (Matheny et al., 2007a).
Strophariaceae
Singer (1986) recognised two subfamilies, Stropharioideae and Pholiotoideae, differentiated primarily by spore print colour, i.e. purple-brown or dark brown in the former and rusty, cinnamon or ochre-brown in the latter. These two subfamilies correspond to the two genera Psilocybe and Pholiota, both in the broad sense, as accepted by Kühner (1980), within the tribe Pholioteae of the Strophariaceae. Kühner (1980) also recognised the tribes Bolbiteae, Crepidoteae, Gymnopileae, Panaeoleae and Tubarieae, each of which is now recognised at family level in molecular-based classifications, although no valid name at family rank is available for some of the tribes.
Singer (1986) accepted four genera in the Stropharioideae, namely Stropharia, Hypholoma (which he called Naematoloma), Psilocybe and Melanotus in the Strophariaceae, using characters such as the degree to which the pileus was hygrophanous, stipe position, and the presence or absence of a subcellular hypodermium and chrysocystidia. He noted that some authors combined all four genera, because of the difficulty of delimiting individual genera, but he considered that his arrangement presented ‘a solution to all difficulties encountered’. Kühner (1980) treated these same four genera under Psilocybe with subgenera Psilocybe (including Deconica and Melanotus), Hypholoma and Stropharia. Noordeloos (1999) also accepted Psilocybe as the only genus of the Stropharioideae, with six subgenera. Four subgenera, Psilocybe, Melanotus, Hypholoma and Stropharia, more-or-less corresponded to the four genera as recognised by Singer (1986). Two additional subgenera were accepted by Noordeloos (1999) for infrageneric units recognised by Singer (1986), viz. subgenus Stercophila for Stropharia section Stercophila of Singer (1986) and subgenus Stropholoma for Naematoloma section Stropholoma of Singer (1986).
Recent molecular analyses (Moncalvo et al. 2002; Walther et al. 2005; Matheney et al., 2007) do not unambiguously support the classifications of Kühner (1980), Singer (1986) or Noordeloos (1999). For Psilocybe, most species fell within a single clade /psilocybe in the molecular analysis of Moncalvo et al. (2002). However, a group of psilocybin-containing taxa from section Caerulescentes as circumscribed by Singer (1986), including species such as Psilocybe semilanceata, P. cyanescens and the Australian P. subaeruginosa, were quite separate from the core species of Psilocybe, and were designated as the /psychedelia clade. These two separate clades were also recovered by Maruyama et al. (2003) [albeit with no sampling of intervening clades], Walther et al. (2005) and Bridge et al. (2008). The separation of the /psilocybe clade (which includes the type of the genus, P. montana) from the group around P. cyanescens was confirmed from multilocus data by Matheny et al. (2007a), who placed the former in the Strophariaceae and the latter in the Hymenogastraceae. A proposal by Redhead et al. (2007) to change the type of Psilocybe (by conservation) to P. semilanceata has been accepted. Consequently, the correct name for the /psychedelia clade is Psilocybe, and the remaining species from Psilocybe sens. lat. are now placed in the genus Deconica.
All Australian species of Deconica and Psilocybe sens. strict. (equivalent to the /psychedelia clade) are keyed out in FunKey together under Psilocybe, pending formal placement of Australian species in the two genera. On molecular grounds Melanotus is clearly nested within Deconica (Moncalvo et al., 2002: under /psilocybe). The two genera differ only in the position of the stipe, and they should be combined (as Deconica), but Melanotus hepatochrous is retained for the key because that is the current name in Australian literature. Even after it is recombined as Deconica, it would be sensible to key out species with non-central stipe as a distinct unit.
The circumscription and relationships of Phaeogalera and Kuehneromyces (neither of which is confirmed as occurring in Australia) have been the subject of some debate (Kühner 1980; Singer, 1986). Phaeogalera was treated as section Porospora of Galerina in the Cortinariaceae by Singer (1986), but with some species in Pholiota, while Kühner (1980) recognised it as subgenus (Porospora) of Naucoria in the tribe Tubarieae of the Strophariaceae. Kuehneromyces was treated as an independent genus of the Pholiotoideae by Singer (1986), but as a subgenus of Pholiota by Kühner (1980). The position of Phaeogalera and Kuehnermyces is yet to be conclusively established. In the /psilocybe clade of Moncalvo et al. (2002) 17 species of Psilocybe clustered together, at the base of which clustered one Psilocybe (P. subcoprophila) together with one species each of Hypholoma (H. udum, from section Psilocyboides), Phaeogalera (P. stagnina) and Kuehneromyces (K. mutabilis). There was poor statistical support for the relationships among these species, but the overall /psilocybe clade did have some statistical support. Moncalvo et al. (2002) suggested that Psilocybe sens. strict. (i.e. the current Deconica) might be extended to include these three species from other genera. Moncalvo et al. (2002) found that Pholiota oedipus [placed in this genus by Singer (1986) but by others in Phaeogalera] clustered not with /psychedelia, but at the base of the /tubarioid clade. Intriguingly, Walther et al. (2005) found that Kuehnermyces mutabilis did not fall in the clade for Psilocybe sens. strict, but was basal to the psilocybin-containing clade, along with Phaeogalera oedipus. A different species of Kuehneromyces (K. rostratus) was included by Matheney et al. (2007a), and this was basal to the Psilocybe sens. strict. clade, while Hypholoma udum did not cluster with Psilocybe, rather with other species of Hypholoma (see below). Gulden et al. (2005), using molecular data from a wide sampling of Galerina species, found that several collections of Phaeogalera stagnina were in an isolated position sister to a single Galerina species (G. badia), but not near other Galerina species or species of Strophariaceae. Moreover, two species of Kuehneromyces clustered together in the vicinity of Psilocybe phillipsii [as Melanotus, which is a Deconica] within the cladogram, but without strong statistical support for specific relationships. The distinctiveness of Phaeogalera and Kuehneromyces and their relationship to Deconica and Psilocybe needs further research.
Singer (1986) differentiated four sections in Hypholoma (as Naematoloma). The single species of section Cyanoloma had blueing fruit-bodies, does not occur in Australia, and there are no molecular data available. For section Stropholoma see below under Leratiomyces. Most species fell in two sections: Psilocyboides for species growing singly or in small groups, on litter or mulch or often among mosses, and the type section [which Singer (1986) called Naematoloma], members of which grow in dense clusters on wood. This separation was followed by Noordeloos (1999), as sections Psilocyboides and Fasciculares, respectively, within Psilocybe subgenus Hypholoma. In the tree of Moncalvo et al. (2002), species of section Psilocyboides were in three different places: (1) Hypholoma udum was well away from the other Hypholoma, at the base of the core Psilocybe clade; (2) H. subericaeum (from section Psilocyboides) clustered with a single species of Pholiota; and (3) H. ericaeum was at the base of the clade containing four taxa of section Naematoloma (= section Fasciculares). In the molecular tree of Walther et al. (2005) three species from each of the two main sections of Hypholoma fell within adjacent, well-supported clades, with the Psilocyboides clade including Hypholoma udum, in accord with the placement of this species by Singer (1986) and Noordeloos (1999). However, Jacobsson & Larsson (2007) recovered a clade with a mixture of Hypholoma and Stopharia species, with Phaeonematoloma myosotis [the type of this genus but also placed in Pholiota, as by Singer (1986) or Hypholoma, as by Knudsen & Vesterholt (2008] at the base of this clade. Further analysis is required to confirm relationships within Hypholoma and to Stropharia. In the meantime we key out Hypholoma section Psilocyboides separately from Hypholoma (other) [the latter being Naematoloma section Naematoloma of Singer (1986) and Psilocybe subgenus Hypholoma section Fasciculares of Noordeloos (1999)].
In Stropharia, Singer (1986) recognised three sections, Mundae (with, e.g. S. coronilla and S. rugosoannulata), Stropharia and Stercophila, and the same three taxa were also recognised by Noordeloos (1999) under Psilocybe, with Stropharia and Stercophila as subgenera, and Mundae as a section within subgenus Stropharia. In the tree presented by Moncalvo et al. (2002), two members of section Stercophila were in a clade together with an unidentified species of Stropharia, while five other species, representing the two other sections, formed a separate clade. The relationships of the two Stropharia clades in relation to each other and to several other clades were not resolved, including the clade containing species of Hypholoma section Fasciculares. Bridge et al. (2008) recovered the separate clade for sections Stropharia and Mundae, except that S. hornemanii clustered at the base of a clade otherwise containing species of Hypholoma. The distinction between section Stercophila and species of the other two sections was also observed by Walther et al. (2005) and by Jacobsson & Larsson (2007). In the latter study, Stropharia semiglobata fell among Hypholoma species but with low statistical support for relationships. Species of Stropharia section Stercophila are characterised morphologically by the combination of glutinous pileus and stipe, growth on dung and chrysocystidia sometimes lacking; and in addition acanthocytes appear to be absent [not mentioned for any species of the section by Noordeloos (1999)]. Therefore, Stropharia section Stercophila is keyed out separately to Stropharia (other), which includes members of both sections Mundae and Stropharia.
The species that has been commonly known as Hypholoma aurantiacum or Stropharia aurantiaca does not fit well in either Hypholoma or Stropharia. It differs from typical Hypholoma by having a pileus that is viscid when fresh, but is aberrant in Stropharia due to the subcellular hypoderm. Singer (1986) placed it in Hypholoma (as Naematoloma), in section Stropholoma, alongside N. magnivelare and N. squamosum, noting that species of this section, some of which lacked chrysocystidia, were intermediate between Stropharia, Hypholoma (as Naematoloma) and Psilocybe. Noordeloos (1999) accepted section Stropholoma within Psilocybe, with the same circumscription as Singer (1986), except for the addition of Psilocybe percevalii. The name Hypholoma aurantiacum, as widely used in recent decades, is a misapplication of Psilocybe ceres. Using nLSU molecular data, Bridge et al. (2008) showed that Hypholoma aurantiacum sensu auct. belongs in a clade with some secotioid fungi from the genus Leratiomyces (including the type of that genus, L. similis), two species formerly placed in the secotioid genus Weraroa (but not the type, which falls in the /psychedelia clade of Psilocybe), and the other species of section Stropholoma accepted by Singer (1986) and Noordeloos (1999). The same clade had been recovered by Moncalvo et al. (2002), and informally named /magnivelaris. Bridge et al. (2008) recognised the clade at generic rank, for which the correct name is Leratiomyces. Thus, Hypholoma aurantiacum sensu auct. is now known as Leratiomyces ceres, and this is the sole agaricoid species of the genus occurring in Australia. The one species of Leratiomyces (as Stropharia squamosa) sampled by Walther et al. (2005) was separate to the clades of Stropharia, Hypholoma and Psilocybe, in accord with Bridge et al. (2008), but was sister to an isolate labelled Agrocybe pediades. This isolate was possibly misidentified, because Bridge et al. (2008) noted that its nLSU sequence differed significantly from others identified as the same species.
Singer (1986) recognised five genera in the Pholiotoideae: Kuehneromyces (see above), Pachylepyrium, Phaeomarasmius (inclusive of Flammulaster), Pholiota and Pleuroflammula, separated by characters including spore print shade, fruit-body habit (pleurotoid in Pleuroflammula), hygrophaneity of the pileus and pileipellis structure and gelatinisation. Although Singer (1986) considered that the distinction between Kuehneromyces and Pholiota was ‘well enough in evidence’, the separation of the two (and indeed other genera) was rather complicated, as evidenced by the paragraph-length couplets in his key to the genera of the Pholiotoideae. Molecular evidence (Matheny et al., 2007a) places Pleuroflammula in the Crepidotaceae and Phaeomarasmius in the Tubariaceae.
Within Pholiota, Singer (1986) recognised five subgenera: Pholiota (chrysocystidia and annulus usually present), Flammula (annulus usually lacking), Hemipholiota (chrysocystidia lacking, mostly with well-developed annulus), Phaeonaematoloma (glutinous stipe surface and spore print often dark or violet brown) and Ploccoloma (very small fruit-bodies). An indication of the difficulties faced by Singer (1986) in trying to describe poorly circumscribed subgenera can be seen in the following contorted and qualified characterisation of subgenus Flammula: ‘pleurocystidia always present unless the spores are … [dextrinoid], either as chrysocystidia or as … [thin-walled cystidia] or metuloids or both; pileus not scaly if only chrysocystidia are present, or if no pleurocystidia are present … sometimes [with] white or pallid velar [remnants] … which tend to disappear … but in this case conspicuous non-chrysocystidioid pleurocystidia present’.
Kühner (1980) had recognised five subgenera: Pholiota, Kuehneromyces, Hemipholiota, Flavidula [treated by Singer (1986) as a section of subgenus Pholiota] and Lubricula [treated by Singer as section Lubricae of subgenus Flammula]. Species placed by Singer (1986) in subgenus Flammula had been divided by Kühner (1980) between subgenera Flavidula and Lubricula. For European species of Pholiota, Noordeloos (1999) accepted all of the subgenera of Singer (1986), except for Ploccoloma which did not occur in Europe, but he also recognised subgenera Lubricula and Flavidula [following Kühner (1980)] as well as Sordidae [for Pholiota oedipus, placed by Singer (1986) in section Sordidae of subgenus Hemipholiota]. In addition, Noordeloos (1999) treated Kuehneromyces as a subgenus of Pholiota. If Pholiota alnicola and its close allies are treated at generic level (see below), the correct genus is Flammula (typified by this species). Other species of subgenus Flammula sensu Singer (1986) consequently must be treated under a different name, i.e. subgenus Lubricula, as used by Kühner (1980) and Noordeloos (1999).
By means of molecular data, Moncalvo et al. (2002) recovered a main clade of 13 species from four sections within subgenera Pholiota and Lubricula. They also found that a number of species of Pholiota did not cluster with the rest of the genus. For example: Pholiota myosotis (as Phaeonaematoloma) did not fall within the core Pholiota clade, but closer to Stropharia and Hypholoma; three species (P. lucifera, P. destruens and P. populnea, all as Hemipholiota) clustered together in a clade designated /hemipholiota; Pholiota alnicola (as Flammula) and Pholiota albocrenulata (as Stropharia) were quite separate from the core Pholiota clade; Pholiota subochracea clustered with Hypholoma subericaeum (this cluster outside of the main Hypholoma clade); and for Pholiota oedipus see above under Phaeogalera. Similar placements outside of core Pholiota were also recovered by Walther et al. (2005, for Pholiota alnicola, P. lucifera and P. tuberculosa), Garnica et al. (2007, for Pholiota alnicola) and Jacobsson & Larsson (2007, for Pholiota albocrenulata, P. alnicola, P. myosotis, P. populnea, P. subochracea and P. lignicola). In their multigene study, Matheny et al. (2007a) sampled a single species of Pholiota sens. strict. (P. squarrosa), but did confirm that P. alnicola (as Flammula) was separate (belonging at the base of the Hymenogastraceae).
For core Pholiota, Walther et al. (2005) and Jacobsson & Larsson (2007) both recovered six species (five common to the two studies) in two clades at the base of the Strophariaceae, one paraphyletic in relation to the other. Moncalvo et al. (2002) had recovered little supported structure within core Pholiota, but one subclade contained species predominantly from subgenus Pholiota, while the remaining species were all from subgenus Lubricula. The two clades recovered by Walther et al. (2005) and Jacobsson & Larsson (2007) also corresponded to the two subgenera, except that Jacobsson & Larsson (2007) found that P. squarrosa fell in the same clade as members of subgenus Lubricula, rather than subgenus Pholiota, as found in the other two molecular studies.
Molecular studies have confirmed that the following are distinct from Pholiota sens. strict.: Flammula (for P. alnicola and a few close allies), Hemipholiota (for P. destruens and a few close allies), Hemistropharia (for P. albocrenulata), Meottomyces (for P. oedipus, for which the correct name is Meottomyces dissimulans, and several other species) and Phaeonematoloma (for P. myosotis). None of these genera are confirmed from Australia, and the placement of several other species that fall outside of Pholiota sens. strict. is yet to be resolved.
Australian species of Pholiota have been rarely represented in molecular phylogenies. Rees et al. (2003) included a single sequence from an unnamed collection of Pholiota. In FunKey all Australian species are treated under Pholiota (other) with the exception of Pholiota malicola, which is keyed out separately because it is larger than most other Pholiota and grows in dense clusters.
Agrocybe has traditionally been classified in the Bolbitiaceae, where the brown spore print and hymeniform pileipellis were consistent with other genera of that family (Singer, 1986). However, all studies employing molecular data clearly place the genus elsewhere (Moncalvo et al., 2002; Walther et al., 2005; Garnica et al., 2007; Matheney et al., 2007). In the multilocus study of Matheney et al. (2007a) three species of Agrocybe (including the type, A. praecox), form a well-supported clade within the Strophariaceae. However, results from other studies are at variance with this placement, with A. praecox basal to the Strophariaceae + Hymenogastraceae clade (Garnica et al., 2007) or in a clade of three species of Agrocybe, and this outside the clade containing Strophariaceae and Hymenogastraceae, and basal to a large one containing Psathyrellaceae (Walther et al., 2005), or in a poorly supported clade /agrocybe of several species of Agrocybe which also contains a species of Leratiomyces, that is outside of the Strophariaceae (Monvalco et al., 2002). In addition, some studies have found species of Agrocybe clustering away from the clade with the type. Matheney et al. (2007a) observed that A. erebia is at the base of the Tubarieae clade, and Walther et al. (2005) noted that A. dura and A. vervacti did not cluster with A. praecox, but were sister to the Psathyrellaceae, but A. pediades fell within the Strophariaceae, sister to Stropahria squamosa (which belongs in Leratiomyces). While these differences could be due to sampling of different DNA regions, the data strongly suggest that Agrocybe is polyphyletic. There has been no attempt to relate the different placements of species of Agrocybe to morphological variation in the genus, such as presence or absence of an annulus or germ pore. Further sampling is clearly required of taxa and DNA regions to resolve the generic limits and relationships. The most comprehensive molecular data are that of Matheney et al. (2007a), and thus we follow their placement of the genus in the Strophariaceae, at least as far as the type and allied species.
Tricholomataceae
As circumscribed by Singer (1986), this family contained most of the white-spored agarics. However, many genera have been transferred to segregate families, leaving few in the Tricholomataceae in the strict sense. Some segregate families correspond well with the tribes and subtribes recognised by Singer (1986), but others do not (see discussion under Hydnangiaceae, Hygrophoraceae, Marasmiaceae, Mycenaceae, Omphalotaceae, Physalacriaceae and the /hydropoid and /xeromphalinoid clades). The genera remaining in Tricholomataceae sens. strict. which occur in Australia are Clitocybe, Lepista, Leucopaxillus, Porpoloma and Tricholoma. Placement of most of these genera was confirmed by multilocus molecular data (Matheny et al., 2007a), but that of Porpoloma is based on the occurrence of an un-named species of Porpoloma in the /tricholomatoid clade of Moncalvo et al. (2002) alongside species of Leucopaxillus and Tricholoma.
Singer (1986) included three genera with amyloid spores, Leucopaxillus, Porpoloma and Melanoleuca, in the tribe Leucopaxilleae; but the last genus now belongs in the Pluteaceae. According to Singer (1986), Porpoloma had tricholomatoid fruit-bodies with smooth spores, while Leucopaxillus had clitocyboid or tricholomatoid fruit-bodies in two sections: Leucopaxillus (ornamented spores) and Aspropaxillus (smooth spores). Within Porpoloma, Singer (1986) accepted three subgenera: Porpoloma (for ectomycorrhizal species, predominantly from the Southern Hemisphere), Pogonoloma for P. spinulosum (with intracellular pigment) and Pseudotricholoma for the remaining species. Kuhner (1980) accepted genera having either amyloid or non-amuloid spores and, while recognising Leucopaxillus, he placed Aspropaxillus under Clitocybe and Porpoloma under Tricholoma.
Vizzini et al. (2012), using nLSU and ITS data, found Leucopaxillus and Porpoloma to be polyphyletic. Several species of Leucopaxillus (including the type species) formed a clade sister to a species of Porpoloma from South America (un-named, but presumably in subgenus Porpoloma). Distant from this clade, several smooth-spored species of Leucopaxillus formed a clade sister to several Northern Hemisphere taxa of Porpoloma (two species from subgenus Pogonoloma). Consequently, Vizzini et al. (2012) resurrected Aspropaxillus for smooth-spored Leucopaxillus. However, they did not re-name the second Porpoloma clade, pending molecular data on the type of Porpoloma, nor did they change the placement of Porpoloma metapodium which did not cluster with other species of the genus, and was near to Tricholoma in their ITS tree. Vizzini et al. (2012) also found that three species of Leucopaxillus were not related to Leucopaxillus sens. strict. or Aspropaxillus, and introduced the new genera Giacomia (for L. mirabilis), Notholepista (for L. subzonalis) and Pseudoclitopilus (for L. rhodoleucus). Evidently, numerous lineages containing clitocyboid or tricholomatoid taxa with amyloid spores (ornamented or not) have evolved independently in the Agaricomycetidae. For the moment, FunKey includes Leucopaxillus and Porpoloma with the former having ornamented spores. There are few Australian species in each genus, and molecular data will be required to confirm generic placement.
Most species included by Singer (1986) in Tricholoma (the only genus of his subtribe Tricholomatinae) lacked clamp connections. Some of the clamped species from section Leucorigida have been removed, notably those now accommodated in the genus Macrocybe (see under Unplaced within Agaricales. 1). Tricholoma species form a well-supported clade in molecular studies (Moncalvo et al., 2002; Walther et al., 2005; Matheny et al., 2007a; Vizzini et al., 2012). Most sampled species lack clamp connections, but one from section Pardinicutis (T. pardinum, with clamp connections) falls within the clade. Molecular data are not available for the other section with clamp connections, section Rigida. Most Australian species of Tricholoma lack clamp connections.
Subtribe Clitocybinae, as circumscribed by Singer (1986), contained Clitocybe, Lepista and Tricholomopsis; however, the last genus does not belong in the Tricholomataceae (see under Unplaced within Agaricales. 4). There has been confusion about the typification and limits of Clitocybe, particularly in relation to Lepista. Harmaja (1976) separated Clitocybe from Lepista by the degree of the cyanophilic reaction of the spores, with Lepista having smooth or ornamented spores that were strongly cyanophilic and often adhered in tetrads and collapsed when examined in mounts from dried lamellae, while spores of Clitocybe (which he typified with C. gibba) were cyanophobic. However, Singer (1986) and Bas et al. (1995) distinguished the two genera by spore ornamentation: verrucose in Lepista and smooth in Clitocybe. Molecular data show Clitocybe as conceived by Singer (1986) [with the type C. gibba] to be highly polyphyletic (Moncalvo et al., 2000; Walther et al., 2005; Mathney et al., 2007). Consequently, the following segregate genera have been erected or recognised: Ampulloclitocybe, Cleistocybe (in the /catathelasma clade), Infundibulicybe, Singerocybe (see under Unplaced within Agaricales. 5) and Trichocybe (Harmaja, 2003; Redhead et al., 2002b; Ammirati et al., 2007; Vizzini et al., 2010). Harmaja (2003) accepted C. nebularis as the type of Clitocybe [following Bas et al. (1995) and Redhead et al. (2002b)] and moved the cyanophobic species to Infundibulicybe, which is also characterised by the non-hygrophanous pileus and tear-drop shaped rather than ellipsoid spores. Subtle distinguishing morphological characters can be recognised for the other segregate genea, such as the non-hygrophanous pileus and interwoven hymenophoral trama of Ampulloclitocybe (Harmaja, 2003) or the often ephemeral veil and interwoven hymenophoral trama in Cleistocybe (Ammirati et al., 2007). After the removal of segregate genera, Harmaja (2003) concluded that Lepista must be merged with Clitocybe, under the latter name. However, other authors continue to recognise the two genera (e.g. Knudsen & Vesterholt, 2008). Nevertheless, synonymy of the two genera is supported by molecular data which consistently recovers a clade (or 'grade' in some analyses) containing a mixture of Clitocybe and Lepista species, with Lepista species not forming a monophyletic subclade (Moncalvo et al., 2000; Walther et al., 2005; Mathney et al., 2007; Vizzini et al., 2010, 2012). The core Clitocybe clade also contains one or a few species of Collybia (in the strongly restricted sense of the type C. tuberosa and a few allied species that fruit on fungal fruit-bodies). Several other species of Clitocybe, including C. clavipes and C. lateritia, fall ouside of the core Clitocybe clade and the segregate genera in molecular analyses. The clitocyboid habit (lamellae decurrent, stipe central, lacking an annulus) is a rather simple fruit-body form, and it is clear that this habit (in combination with smooth, inamyloid spores) has evolved independently in many lineages.
Australian species of Clitocybe have been assigned to sections Candicantes, Disciformes and Vernae (Grgurinovic, 1997a), but sequence data are lacking. Most Northern Hemisphere taxa of Candicantes and Disciformes included in molecular analyses fall within the core Clitocybe clade (Vernae is unsampled as yet). Therefore, in FunKey we make a simple distinction between Clitocybe other (inclusive of Singerocybe) with smooth spores and a white or cream spore print and Lepista with ornamented spores and a cream to pinkish brown spore print. The common species Clitocybe semiocculta is keyed out separately because of the habit on wood in combination with usually excentric stipe. Its generic position needs further study.
Several poorly known Australian species currently placed in Clitocybe are omitted from FunKey pending confirmation of their generic placement. These include C. subfrumentacea, C. tortipes and C. peraggregata, all of which lack clamp connections (Grgurinovic, 1997a). However, the absence of clamp connections is rare in Clitocybe (a few species of section Disciformes; Singer, 1986). Clitocybe subfrumentacea was originally included by Cleland (1934) in Clitopilus, which has pink spore print, and Cleland also noted that the spores were ‘rather irregular’. These characters, along with the large, pinkish fruit-bodies are consistent with Rhodocybe, but a re-examination of type material is required. Clitocybe tortipes is also aberrant due to the combination of cylindrical spores, sinuate to adnate lamellae and ‘fibrously splitting’ pileus surface. Clitocybe peraggregata is unusual in having a fruit-body with fan-shaped pilei arising from common branching stipe, and lack of clamp connections. In addition, it has been noted as growing on the ground (certainly the prefered substrate for Clitocybe), but ‘probably from rotten wood’. Species occurring on wood need further examination, especially since the removal of C. lignatilis to Ossicaulis (Redhead & Ginns, 1985). Therefore, C. straminea Cleland, which occurs at the base of stumps or on logs (Cleland 1934; Willis 1963), is also omitted from FunKey. Clitocybe straminea could be referable to Trogia (and then, not to be confused with Trogia straminea Corner), due to the lignicolous habit and the pileus described as smoky yellow brown due to 'fine fibrils'. Trogia grows on wood or litter and, according to Singer (1986), differs from Clitocybe in having a trichodermal pileipellis (although pileipellis hyphae can become repent in age).
Tubariaceae
Kühner (1980) recognised the Tubarieae as a tribe within the Strophariaceae, consisting of Galerina, Phaeocollybia and Naucoria. Within Naucoria, he recognised subgenera Tubaria, Phaeomarasmius, Naucoria and Porospora. Singer (1986) placed Tubaria at generic rank in the Crepidotaceae, and Phaeomarasmius at generic rank in the Strophariaceae, and included within it Flammulaster (as subgenus Carpophilus). Both Horak (1980) and Vellinga (1986) accepted Flammulaster and Phaeomarasmius at generic rank. Naucoria subgenus Naucoria sensu Kühner (1980) corresponds to Flammulaster, and the same taxon has also been called Naucoria section or subgenus Floccularia. Singer’s (1986) concept of Naucoria was based on a different type to that of Kühner (1980), who used Alnicola for Naucoria sensu Singer (1986). The latter is not confirmed from Australia. Naucoria subgenus Porospora is called Phaeogalera at generic rank, and was placed under Galerina by Singer (1986); this too has not been confirmed as occurring in Australia.
In regard to this complicated variation in classification, molecular data indicate that Naucoria (= Alnicola), Galerina and Phaeocollybia belong in the Hymenogastraceae, but confirm the suggestion by Kühner (1980) of a relationship between Tubaria, Phaeomarasmius and Flammulaster. These three genera were placed together in a clade designated by Matheny et al. (2007a), in the absence of a suitable name at family rank, as Tubarieae, which is the tribe name used by Kühner (1980). It seems reasonable to accept the Tubarieae at family rank, and the name Tubariaceae has been formally published. It is sister to the clade comprising Crepidotaceae together with Inocybaceae (Matheny et al., 2007a).
In the molecular analyses of Moncalvo et al. (2002) and Aime et al. (2005), a small group of Tubaria species forms a clade that is distant from other members of the Crepidotaceae, which is where Singer (1986) placed the genus. The Tubaria species studied were different in the two analyses, but both included the Australian T. rufofulva. The multi-locus study of Matheny et al. (2007a) included four species of Tubaria (some still referred to Pholiota and Naucoria, but transferred to Tubaria by Matheny et al., 2007b), in the Tubarieae clade. A wider sampling of more than a dozen species of Tubaria by Matheny et al. (2007b) recovered a clade comprising all species with the exception of T. minima (which clustered with a collection of Pachylepyrium). Tubaria confragosa, a species treated by Singer (1986) as a Phaeomarasmius, despite its transfer to Tubaria by Harmaja (1978), fell within the main Tubaria clade (Matheny et al., 2007b). Singer (1986) accepted rough-spored species in Tubaria, and although no molecular data are available such species, none are known from Australia.
The molecular analyses of Moncalvo et al. (2002), Aime et al. (2005) and Matheny et al. (2007a) all had a few species of Phaeomarasmius and Flammulaster at the base of the clade of Tubaria species. In a wider sampling of the two genera by Matheny et al. (2007b) there was a grade at the base of the Tubaria clade comprising seven species of Phaeomarasmius and Flammulaster, neither genus being monophyletic. One collection from each genus fell within the Tubaria clade, but Matheny et al. (2007b) suggested that these collections were misidentified.
Singer (1986) subsumed Flammulaster under Phaeomarasmius (as subgenus Carpophilus), but Horak (1980e) and Vellinga (1986) accepted Flammulaster at generic rank. Horak (1980e), focusing on New Zealand species of the two genera, emphasised spore shape (limoniform in Flammulaster, but rounded, amygdaliform or phaseoliform in Phaeomarasmius) as a key distinguishing feature. Horak (1980e) accepted in both genera species with an epithelial pileipellis (made up of chains of globose elements) or with a trichoderm, but allowed for species with cystidioid (differentiated) terminal elements in the pileipellis only in Flammulaster. Vellinga (1986) commented that the European taxa of Flammulaster were a rather heterogeneous assemblage, and that the differences emphasised by Horak (1980e) between Flammulaster and Phaeomarasmius did not hold up in other geographic areas. Further sampling of the two genera for molecular studies, and an integration of molecular and morphological data from a much broader geographical range are required before generic limits can be clarified. Flammulaster and Phaeomarasmius are likely to occur in Australia, both being represented in New Zealand (Horak, 1980e), but they are omitted from FunKey, pending re-assessment of possible Australian records.
Un-named family (tribe: Cystodermateae)
Cystoderma as delimited by Singer (1986) contained two sections, Granulosa, with non-amyloid spores, and Cystoderma, with amyloid spores, and was placed in the tribe Cystodermateae of the Agaricaceae. Only species of section Cystoderma have been confirmed as occurring in Australia. Moncalvo et al. (2002) found that representatives of the two sections fell in separate clades (one also containing Floccularia and the other Ripartitella), but there was poor or no statistical support for relationships within and between these clades. Harmaja (2002) erected Cystodermella for species of section Granulosa, which, as well as the non-amyloid spores, also differ in ploidy level, cystidia characters and in lacking a strong association with bryophytes. The multilocus study of Matheny et al. (2007a) only included Cystoderma amianthinum, a member of section Cystoderma. This was the sister taxon to the Nidulariaceae, and was designated as belonging to the tribe Cystodermateae. Species of Cystodermella were not sampled, and nor were members of Floccularia and Ripartitella.
Un-named family (tribe: Gymnopileae)
Singer (1986) distinguished Gymnopilus from Galerina, both of which he included in the Cortinariaceae, on the basis of the former genus generally having more robust fruit-bodies, styrylpyrone pigments, a black KOH reaction on the pileus and spores lacking a plage. Kühner (1980) placed Gymnopilus in the Strophariaceae as the sole member of the tribe Gymnopileae (and also considered Galerina to belong in the Strophariaceae). Rees et al. (1999) discussed a number of Southern Hemisphere species of Galerina (such as Galerina eucalyptorum), often with an excentric stipe and lacking styrylpyrone pigments, but with microcharacters similar to Gymnopilus, that blurred the distinction between the two genera. Nevertheless, they retained Galerina eucalyptorum. In a study utilising ITS sequences of a variety of Northern and Southern Hemisphere species, Rees et al. (2002) found that Gymnopilus species fell within a single clade (that had low statistical support) along with Galerina eucalyptorum, which was consequently transferred to Gymnopilus (where it is correctly known as G. perplexus). Sequence data for the two other Galerina species (G. bulliformis and G. incrustata) discussed by Rees et al. (1999) in relation to Gymnopilus are not yet available.
Various studies, including that of Rees et al. (2003) on 13 Australian collections, utilising data from the nLSU and sometimes also other regions, have found that species assigned to Gymnopilus on morphology form a distinct, usually well-supported clade (Moncalvo et al., 2002; Gulden et al., 2005; Matheny et al., 2007a). This clade does not fall within the Cortinariaceae or the Strophariaceae, but is treated by Matheny et al. (2007a) as distinct from other families, and is designated by the tribal name Gymnopileae. Only two species of Gymnopilus were sampled by Matheny et al. (2007a), nor did they sample members of the /mycenopsis clade of Galerina, which was found by Gulden et al. (2005) to be sister to a clade comprising Gymnopilus and Galerina paludosa. The distinction between Gymnopilus and Galerina needs further investigation with increased sampling from Southern Hemisphere species of the latter genus.
The genus Pyrrhoglossum was accepted by Singer (1986) as distinct from Gymnopilus on the basis of the excentric stipe. However, Rees et al. (2003) found that ITS sequences from the type, P. pyrrhum (known from Australia), placed the species within Gymnopilus.
Un-named family (/hydropoid clade)
Moncalvo et al. (2002), in their analysis of nLSU data, found that two members of Mycena section Adonideae (the non-Australian M. aurantiidisca and M. adonis) fell in the /adonis clade. While the position of this clade was not resolved in relation to others, it was quite separate from the /mycenaceae clade that contained various other species of Mycena. A position well outside the Mycenaceae for M. aurantiidisca and another member of section Adonideae (M. amabilissima) was confirmed by Matheny et al. (2007a), who found these species to belong in the /hydropoid clade, as did the Japanese Mycena auricoma (described in section Radiatae). We key out Mycena section Adonideae separately, but retain the species in Mycena pending formal recognition of a new genus for the section.
Un-named family (/xeromphalinoid clade)
Singer (1986) accepted Xeromphalina in a wide sense, inclusive of Heimiomyces, which he treated as one of two subgenera. We follow Horak (1979d) in recognising Heimiomyces at generic rank, for species with collybioid (rather than omphalinoid) fruit-bodies, usually adnate lamellae and a rounded to umbonate or papillate pileus, along with typically diverticulate cheilocystidia. According to the data provided by Horak (1979), some species of Heimiomyces have a well-marked, gelatinous zone in the upper pileus trama (which could be interpreted as a subcutis), subtended by thick-walled hyphae, whereas this type of gelatinous zone is lacking in Xeromphalina. In addition, the pileipellis in Heimiomyces varies from an epithelium (in non-Australian species) to a cutis or trichoderm, often with the terminal elements branched or diverticulate, whereas in Xeromphalina the pileipellis is often a cutis with un-modified terminal elements (Horak, 1979d). Recognition of Heimiomcyes is consistent with molecular data (Moncalvo et al., 2002) that show the two genera as monophyletic, forming adjacent clades, in a clade designated /xeromphalinoid. Placement of Heimiomyces under Xeromphalina would also be consistent with the molecular data. The close relationship of the two genera is clear, but their position within the Agaricales is now uncertain, although it is reasonable to suggest that this clade is likely to have family rank. Singer (1986) had placed Xeromphalina in the tribe Myceneae of the Tricholomataceae, but in the study of Moncalvo et al. (2002) the /xeromphalinoid clade did not cluster with other genera of this tribe, such as Mycena, which fell in a clade which they designated Mycenaceae. In the multilocus study of Matheny et al. (2007a), a single species of Xeromphalina was unplaced in any family, clustering adjacent to members of the Pterulaceae and Typhulaceae, but distant from the Mycenaceae.
Unplaced within Agaricales. 1
Pegler et al. (1998) erected Macrocybe for some agarics formerly placed in Tricholoma section Leucorigida (Singer 1986), but differing from other species of Tricholoma in the presence of clamp connections and in not being ectomycorrhizal. In addition to the morphological distinction from Tricholoma, Pegler et al. (1998) demonstrated on molecular analysis that the one species of Macrocybe sampled did not cluster with the several species of Tricholoma studied, but with a species of Calocybe. However, a limited selection of other genera were included, and siderophilous granulation, characteristic of Calocybe, is absent in Macrocybe. In the molecular analysis of Moncalvo et al. (2002), using nLSU data, two species of Macrocybe formed a clade sister to one clade containing some species of Callistosporium, and these two clades were sister (with low statistical support) to some taxa of Entoloma, but distant to any species of Calocybe. The multilocus study of Matheny et al. (2007a) did not sample Macrocybe, but Callistosporium fell within a clade designated as /catathelasma, which was separate from the Entolomataceae (where Entoloma belongs), Lyophyllaceae (where Calocybe belongs) and Tricholomataceae (where Tricholoma belongs). Data from other regions of DNA are required to confirm the family position of Macrocybe. Currently, Macrocybe crassa is the only known Australian member of this genus.
Unplaced within Agaricales. 2
Mycenella (so far represented in Australia only by Mycenella margaritispora) is accepted in the sense of Singer (1986), who distinguished it from Xerula (which he included in Oudemansiella) by the rough spores, or in the case of the single smooth-spored species (the European Mycenella salicina), by the combination of mycenoid habit and small spores. Singer (1986) placed Mycenella in the subtribe Oudemansiellinae of the Tricholomataceae, alongside Oudemansiella. Using two DNA regions, Garnica et al. (2007) found that Mycenella bryophila is a member of the Agaricales, but its familial position was not resolved, although it did not fall in the same clade as Xerula radicata.
Unplaced within Agaricales. 3
Panellus longinquus was included in Panellus by Singer (1986) and Corner (1987a), although Horak (1983a) placed it Pleurotopsis. In the molecular study of Jin et al. (2001), P. longinquus fell in a clade quite separate to the type of Panellus (P. stypticus), along with P. violaceofuscus and P. ringens. Taxon sampling in that study did not include many other genera. Nevertheless, in studies with wide taxon sampling, Moncalvo et al. (2002) and Matheny et al. (2007a) confirmed that Panellus longinquus (as Pleurotopsis) did not cluster with P. stypticus. It was an isolated lineage (Moncalvo et al. (2002) or clustered with a single species of Hemimycena (Matheny et al., 2007a).
Panellus longinquus clearly does not belong in Panellus, and Jin et al. (2001) recommend using the name Pleurotopsis for the clade containing P. longinquus. We retain the species in Panellus, pending the introduction of a suitable generic name, because Pleurotopsis is actually a synonym of Hohenbuehelia, as indicated by Singer (1986) and discussed by Thorn et al. (2006). This is because Pleurotopsis (Henn.) Earle is typified by Marasmius spodoleucus Berk. & Broome which is a synonym of Agaricus cyphelliformis Berk., and this species belongs in Hohenbuehelia (Thorn & Barron, 1986; Singer 1986).
Unplaced within Agaricales. 4
Tricholomopsis was placed by Singer (1986) in subtribe Clitocybinae of the Tricholomataceae, alongside Clitocybe and Lepista. Singer (1986) included two sections in Tricholomopsis: section Tricholomopsis for species such as T. decora and the only known Australian representative, Tricholomopsis rutilans, and section Platyphyllae for species such as T. platyphylla. The latter species has been transferred to Megacollybia. Molecular data (Matheny et al., 2007a) place Tricholomopsis as the sister taxon to the Amanitaceae, and not with genera of the Tricholomataceae, whilst Megacollybia is within the /hydropoid clade.
Unplaced within Agaricales. 5
Singerocybe contains species formerly in Clitocybe that have inflated elements in the trama of the pileipellis. Molecular data confirm that Singerocybe is separate from the core Clitocybe clade in the Tricholomataceae, but do not allow placement into a family (Vizzini et al., 2010). Clitocybe clitocyboides belongs in Singerocybe, but in FunKey it is keyed out under Clitocybe.
BOLETALES
Boletaceae, Hygrophoropsidaceae, Paxillaceae, Serpulaceae and Tapinellaceae
The order Boletales was introduced for fungi with fleshy, stipitate fruit-bodies and tubular-poroid hymenium ('boletes'). However, for a long time some lamellate genera have been admitted in Boletales. Thus, for example, Kühner (1980) included Gomphidius (not known from Australia) and Phylloporus in the Boletaceae, Omphalotus and Hygrophoropsis in the Hygrophoropsidaceae and Paxillus in the Paxillaceae. Singer (1986) treated ‘boletes’ at a different rank (suborder Boletineae of the Agaricales) but also accepted Phylloporus (Boletaceae) and Hygrophoropsis, Omphalotus and Paxillus (Paxillaceae). In addition, Meiorganum (which now contains some lamellate species) was included in the Boletaceae by Singer (1986), but at that time the genus contained only the type M. neocaledonicum, which is poroid at maturity.
The delimitation of families within the Boletales changed significantly from that of Kühner (1980) and Singer (1986) with integration of molecular data. We follow the classification outlined by Binder & Hibbett (2007) based on a multigene phylogeny, which is set out in the ‘Taxonomy supplement’ to their paper. In this arrangement, the Boletales encompasses a wide variety of morphology, including fleshy, stipitate fruit-bodies with the hymenium tubular-poroid (boletes) or lamellate (agarics); gasteroid fruit-bodies; or resupinate fruit-bodies with smooth, hydnoid or merulioid (wrinkled) hymenium (Binder & Hibbett, 2007).
On morphological and chemical evidence, Phylloporus was treated as an independent genus of Boletales (Kühner, 1980; Watling & Gregory, 1991) or Boletineae (Singer, 1986) with affinities with boletes such as Xerocomus (Boletaceae) rather than the lamellate Paxillus (Paxillaceae). This relationship has been confirmed by molecular data, which place Phylloporus in the Boletaceae (Binder & Hibbett 2007). As regards generic limits, Phylloporus nests within one of the several clades of Xerocomus (which is polyphyletic) (Binder & Hibbett, 2007; Neves et al., 2012). Indeed, Bresinsky & Besl (2003) [not seen, quoted by Neves et al. (2012)] went as far as transferring a few species of Phylloporus to Xerocomus, while also suggesting that a new genus was needed for other species of Phylloporus. Neves et al. (2012) argue for the retention of Phylloporus for the moment, especially due to the polyphyly of Xerocomus, from which several genera have been segregated by Šutara (2008). Nevertheless, all species of Xerocomus sampled by Neves et al. (2012) belong in Xerocomus in the strict sense, and not to segregate genera such as Xerocomellus, although there is low support in the backbone of their phylogeny. Therefore, we retain Phylloporus as an independent genus, pending studies that include wide taxon sampling from the Southern Hemisphere.
Hygrophoropsis was distinguished from Paxillus by the pale spore print and placed alongside that genus in the Paxillaceae by Singer (1986), but in the Hygrophoropsidaceae, with Omphalotus, by Kühner (1980). Molecular analyses confirm the distinctiveness of Hygrophoropsis and its placement in the Hygrophoropsidaceae, and show that the closest relative within that family is Leucogyrophana (with merulioid hymenium) rather than the more distantly related Paxillus (Paxillaceae; Binder & Hibbett, 2007). Omphalotus is now considered to be outside of the Boletales, in the Omphalotaceae (Binder et al., 1997).
Austropaxillus was segregated from Paxillus on the basis of molecular evidence, corroborated by chemical and morphological characters such as the absence of pleurocystidia, lack of clamp connections in some species and often elongate spores (Bresinsky et al., 1999). Singer (1986) had placed species now in Austropaxillus in sections Defibulati, Parapaxillus and Veluticipites of Paxillus.
Several wood-inhabiting taxa were included by Singer (1986) in Paxillus: Paxillus panuoides (type of Tapinella) and P. curtisii in section Panuoides and P. atrotomentosus in section Atrotomentosi. Kühner (1980) had also accepted these species in Paxillus, in subgenus Tapinella. Redhead & Ginns (1985), while recognising close morphological similarities between Paxillus and Tapinella, argued that trophic status, mycorrhizal in the former and wood-decaying in the latter, supported recognition at genus rank of Tapinella (in which they placed T. panuoides and T. atrotomentosa).
Molecular evidence excluded species of Tapinella from Paxillus (Bresinsky et al., 1999), and multigene data place Tapinella atrotomentosa with T. panuoides in the Tapinellaceae, alongside Pseudomerulius and the poroid Bondacevomyces (Binder & Hibbett, 2007). The Tapinella panuoides group is keyed out in FunKey.
With the exclusion of Austropaxillus, Tapinella and Paxillus curtisii (see below), Paxillus is restricted to the section Paxillus of Singer (1986), which is native to the Northern Hemisphere; Austropaxillus is a Southern Hemisphere genus.
The lignicolous Paxillus curtisii has lamellae with numerous interconnections, and the fruit-bodies lack a stipe. Redhead & Ginns (1985) transferred it to Pseudomerulius whose type (P. aureus) has a resupinate fruit-body with a merulioid (wrinkled) hymenium; this placement is followed by most authors (e.g. Baldoni et al., 2012). However, Singer et al. (1990) considered that Paxillus curtisii to be better placed in Meiorganum, alongside the type of that genus, the poroid, pseudostipitate M. neocaledonidum, presumably because both have pileate fruit-bodies. Australian material of a wood-inhabiting, pileate fungus with non-amyloid spores has been placed under Meiorganum olivaceoflavidum by Watling & Li (1999), who also included M. curtisii. For this reason, in FunKey we accept the Meiorganum olivaceoflavdium group (including M. curtisii) for Australian species with non-dextrinoid spores but whose fruit-bodies are otherwise similar in appearance to Tapinella panuoides. Spores of Tapinella are dextrinoid, whereas those of Pseudomerulius, as well as Meiorganum curtisii and M. neocaledonicum, are not (Singer et al., 1990; Baldoni et al., 2012). In terms of family placements, Singer et al. (1990) retained Tapinella panuoides in Paxillus, which they placed in the Paxillaceae, while Meiorganum was included in the Boletaceae. Binder & Hibbett (2007) placed Meiorganum in the Paxillaceae, with Tapinella and Pseudomerulius in the Tapinellaceae. Within the Tapinellaceae (designated by them as Tapinellineae), the molecular analysis by Binder et al. (2010) recovered separate clades for each of Tapinella (T. atrotrometosa and T. panuoides) and Pseudomerulius (P. aureus and P. curtisii), but no species of Meiorganum were sampled. The delimitation of Meiorganum and its relationship to Pseudomerulius requires molecular data for the former genus, including for the extra-limital Meiorganum agathidis (originally described in Merulius) which, according to Corner (1971), has dextrinoid spores.
CANTHARELLALES
Cantharellaceae
Corner (1966) characterised Cantharellus by the usually terrestrial fruit-bodies with the pileus margin initially incurved, a thickening hymenium, and with hyphae nearly always with clamp connections, and lacking secondary septation. He also recognised Craterellus, with the pileus margin straight from the first, and lacking clamp connections, as well as Pseudocraterellus, with hyphae lacking clamp connections but with secondary septation. Within Cantharellus Corner (1966) recognised subgenus Cantharellus for brightly coloured species such as C. cibarius and subgenus Phaeocantharellus for dark, often deeply infundibuliform taxa such as C. tubaeformis. According to Corner (1966), the latter subgenus included C. fuligineus, which lacks clamp connections, and C. cinereus, which he illustrated with clamped hyphae, but which was interpreted by Pegler et al. (1997) and others as lacking clamps, and placed in Pseudocraterellus (Pegler et al., 1997) or Craterellus (Breitenbach & Kranzlin, 1986).
Using molecular data, Dahlman et al. (2000) showed that at least some species of subgenus Phaeocantharellus (also known as subgenus Leptocantharellus) are more closely related to Craterellus than to species of subgenus Cantharellus. Furthermore, at least the type of Pseudocraterellus (P. sinuosus) also belongs in Craterellus, which is characterised morphologically by the hollow stipe. Using molecular data from several DNA regions, Moncalvo et al. (2007) confirmed the distinction between Cantharellus and Craterellus, which fell in an un-named clade also containing Hydnum (which has a toothed hymenium), and this clade was within the larger /cantharelloid clade. The same larger clade was designated as Cantharellales by Hibbett (2007), but we use the family name Cantharellaceae for the clade containing Cantharellus, Craterellus and Hydnum.
No Australian specimens have been included in molecular phylogenetic analyses, but we assume that Australian collections with a dark brown or grey pileus (with clamps or not) are referable to Craterellus, and we restrict Cantharellus for the bright yellow, orange or pink species. The poorly known species Cantharellus ochraceoravus, with a golden brown to cinnamon pileus, is retained in Cantharellus for the present.
GLOEOPHYLLALES
Gloeophyllaceae
Thorn et al. (2000) recovered a clade from analysis of nLSU data which included several species of Neolentinus along with one Gloeophyllum. They placed this clade within the Polyporales but with larger taxon sampling Binder et al. (2005) recovered a clade for Gloeophyllum, Neolentinus and several other genera that was outside of the Polyporales clade and distant from Lentinus. Subsequently, Kirk et al. (2008) recognised the order Gloeophyllales with the single family Gloeophyllaceae. Using multi-locus data and a wide taxon sampling, Garcia-Sandoval et al. (2011) confirmed the distinctiveness of the Gloeophyllales, which contains mostly brown-rot fungi with a range of fruit-body types, including pileate (poroid or lamellate) and resupinate (poroid or corticioid).
Neolentinus was erected for several species previously placed in Panus (Corner, 1981; Pegler, 1983b as a subgenus) which have brown rather than white rot (Redhead & Ginns, 1985). Note that the isolate labelled Neolentinus dactyloides (an Australian species) in the analysis of Hibbett & Vilgalys (1993), which did not cluster with other Neolentinus but rather with a species of Pleurotus, was misidentified and actually belongs in Pleurotus (Thorn et al., 2000). Four species of Neolentinus form a well-supported clade in the molecular analyses of Garcia-Sandoval et al. (2011), with a species of Veluticeps (with a resupinate, corticioid fruit-body) present on a long branch within this clade, but only in some analyses.
Gloeophyllum contains both lamellate and poroid species (Ryvarden & Gilbertson, 1993) and molecular data analysed by Garcia-Sandoval et al. (2011) shows that it is polyphyletic. One clade consists of predominantly lamellate species (such as G. sepiarium and G. striatum), while another has predominantly poroid species (such as G. odoratum). The second clade may contain subgroups, with the name Osmoporus available for its core members. The lamellate species included in FunKey fall within Gloeophyllum in a strict sense.
HYMENOCHAETALES
Repetobasidiaceae
The Hymenochaetales exhibit a diverse array of fruit-body types, from resupinate to pileate, with the hymenium varying from corticioid or tubular-poroid to lamellate. The few lamellate taxa all resemble Omphalina (with omphalinoid fruit-body), are associated with bryophytes, and fall within the /rickenella clade, one of five within the order identified by Larsson et al. (2007). Other fruit-body types in this clade include the stipitate-pileate Cotylidia, which has a smooth hymenium, and resupinate, corticioid genera such as Repetobasidium and Resinicium. Larsson et al. (2007) did not assign a rank to the five major clades, but family rank seems appropriate, and Larsson (2007) referred to the 'Rickenella family'. The family name Repetobasidiaceae is available.
The omphalinoid Hymenochaetales, represented by Loreleia postii group and Rickenella in Australia, have only recently been transferred from the Agaricales (Redhead et al., 2002b; Larsson et al., 2007). Most were included by Singer (1986) in Gerronema which, in retrospect, is highly heterogeneous, with section Romagnesia subsection Venustissima including species of Loreleia (lacking clamps and pleurocystidia) and Lichenomphalia (Hygrophoraceae), and with section Fibulae (typified by G. fibula) now recognised at generic rank as Rickenella, characterised by the presence of distinct pleurocystidia and pileocystidia, as well as clamp connections (Redhead et al., 2002a; 2002b). The generic distinctiveness of Loreleia and Rickenella is confirmed by molecular evidence (Moncalvo et al. 2002; Redhead et al. 2002b; Larsson et al., 2007), although sequence data are available for only one species of the first genus.
POLYPORALES
Polyporaceae
Most members of the Polyporaceae are poroid; exceptions are the lamellate Lenzites, Lentinus and Panus and one lamellate species of Trametes (T. elegans).
Lentinus and Panus are largely circumscribed according to the treatments of Corner (1981) and Pegler (1983b), with the latter author recognising Panus as a subgenus of Lentinus. In both treatments Lentinus and Panus are dimitic, with skeleto-ligative hyphae in the former and sleletal hyphae in the latter. Panus also differs in that some species have thick-walled pleurocystidia. The arrangement of Singer (1986) differed significantly from that of Corner (1981), in that most species of Lentinus were within Panus, and Lentinus was restricted to species now placed in either Neolentinus or Lentinula. Singer (1986) also treated some species of Lentinus and Panus in Pleurotus.
Current exceptions to the circumscription of Lentinus and Panus by Corner (1981) and Pegler (1983b) which have been confirmed by molecularanalyses are the placement of Panus giganteus and P. tuber-regium in Pleurotus (Thorn et al., 2000; Karunarathna et al., 2011). In addition, Panus tenebrosus has been transferred to the monotypic Austrolentinus, characterised by the lamellae that are very shallow ridges and the presence of coralloid hyphae in the pileipellis similar to pileipellis structures in the poroid Polyporus blanchettianus (Ryvarden, 1991). No DNA sequences are available for Austrolentinus. Furthermore, several species formerly placed in Panus sections Pulverulenti, Squamosi and Cirrhosi (Corner, 1981; Pegler, 1983b) are now included in Neolentinus. The latter is characterised by the brown rather than white rot (Redhead & Ginns, 1985), and on molecular data it falls in the gloeophyllum clade (i.e. Gloeophyllales) which is quite separate to the Polyporales (Binder et al., 2005).
The emended Panus is monophyletic and quite distinct from Lentinus based on molecular evidence (Hibbett & Vilgalys, 1993; Thorn et al., 2000; Douanla-Meli & Langer, 2010). The latter (with lamellate hymenium) is very close to certain species of Polyporus (with tubular-poroid hymenium) designated as the ‘polyporellus group’. Indeed, in some molecular analyses species of the former genus are adjacent to (Grand et al., 2011) or within a clade otherwise consisting of species of Polyporus, such as P. tricholoma (Krüger & Gargas, 2004; Krüger et al., 2008; Sotome et al., 2008, 2009). Krüger & Gargas (2004) went so far as to transfer the type and another species of Lentinus to Polyporus (although introducing illegitimate names for them). Synonymy of Lentinus and Polyporus has not been taken up by other authors. Sotome et al. (2008) argue that, because Polyporus is highly polyphyletic (with six major clades, some containing other genera) and typification of the genus remains controversial, the transfer of additional species into Polyporus should wait until the status and nomenclature of the various lineages of Polyporus is settled. Therefore, we retain lamellate species in Lentinus.
Ryvarden (1991) recognised Lenzites as being separate from Trametes, in which he accepted T. elegans because it lacked the emergent binding hyphae with acute, sword-like apices that are present in Lenzites. Ryvarden & Johansen (1980) and Corner (1987b), however, had retained T. elegans in Lenzites, reporting tapered endings to the binding hyphae in some collections, but obtuse tips in others. Cunningham (1965) had placed Lenzites under Daedalea, along with T. elegans (as Daedalea palisoti) and Daedaleopsis confragosa (for the latter, see Excluded taxa). Molecular data from two regions place the type of Lenzites (L. betulina) and T. elegans within a single clade along with other species of Trametes, in which is nested a monophyletic Pycnoporus (Tomsovsky et al. 2006). On the basis of a two locus molecular analysis, Welti et al. (2012) segregated Artolenzites (for T. elegans), Pycnoporus and Leiotrametes (for T. lactinea and allies) from a restricted Trametes (in which was placed L. betulina). Several other species of Trametes did not fall in the core Trametes clade or in the segregate genera. Justo & Hibbett (2011), using a five-locus data set, had previously concluded that rather than adopt segregate genera (up to 10 were considered), the best solution was to accept a broad Trametes inclusive of Artolenzites, Lenzites, Pycnoporus and several other genera. However, while we therefore adopt Trametes elegans, we continue to treat Australian species in Lenzites; in particular because, as pointed out by Justo & Hibbett (2011), names in Trametes are not yet available for L. acuta and L. vespacea, and molecular data are lacking for those species.
In terms of family placement, because there is no consensus on family circumscription within the Polyporales, Lentinus and Panus are included in the Polyporaceae. However, it should be noted that the two genera always fall in different major clades, as in the study of Binder et al. (2005) where the former was in the ‘core polyporoid clade’ and the latter in the ‘residual polyporoid clade’. Douanla-Meli & Langer (2010) suggest that Panus is closely related to the Meruliaceae. Both Lenzites and Trametes are in the ‘core polyporoid clade’ of Binder et al. (2005).
RUSSULALES
Auriscalpiaceae
The genus Lentinellus is well circumscribed morphologically (Petersen & Hughes, 2004). Six Northern Hemisphere species form a well-supported clade in molecular analyses (Larsson & Larsson, 2003), and molecular data also support placement in the Auriscalpiaceae (Miller et al., 2007).
Russulaceae
Russula and Lactarius belong in the Russulaceae based on their morphology and molecules (Singer, 1986; Miller et al., 2007), and the two genera have been recognised for more than two centuries as being independent. They are currently separated by the former having sphaerocytes in the lower portion of the lamellae (which are rare in Lactarius) and often lacking lamellulae, and the latter producing latex (Singer, 1986). The several species of the Lactarius clarkeae group (L. clarkeae, L. subclarkeae and Russula flocktoniae) are somewhat intermediate between the two and are keyed out as a separate identification unit to Lactarius (other). Lactarius clarkeae is sometimes found without conspicuous latex (and could then be confused with Russula). Russula flocktoniae lacks latex, but it closely resembles L. clarkeae, especially in the orange pileus and thick-walled terminal elements in the pileipellis. In addition, R. flocktoniae clusters with L. clarkeae and L. subclarkeae on the basis of DNA sequences (Jenny Tonkin, pers. comm.). It is notable that L. clarkeae has sphaerocytes in the lower parts of the lamellae (McNabb, 1971): an attribute that is rare in Lactarius but the usual state in Russula.
A recent multilocus phylogenetic analysis of Lactarius and Russula (Buyck et al., 2008) found four main clades. Most species of Russula fell within one clade, but a few from subsection Ochricompactae grouped with one species of Lactarius. The genus Multifurca was introduced for this latter clade, and Lebel et al. (2013) have recently transferred the Australian Lactarius stenophyllus to Multifurca. Buyck et al. (2008) found that the remaining species of Lactarius fell in two distinct clades: Lactarius sens. strict. (typified by L. torminosus) and another genus later designated as Lactifluus by Verbeken et al. (2011) and Stubbe et al. (2012a). No Australian collections of Lactarius or Russula were analysed by Buyck et al. (2008), although Stubbe et al. (2010) did sample Australian collections of Lactarius subgenus Gerardii (which belongs in Lactifluus). In the analysis of ITS sequences alone by Lebel et al. (2013) collections of Lactifluus fell in two clades, interspersed by Lactarius sens. strict.. One of these clades also contained some Australian collections from the Lactarius clarkeae group, including Russula flocktoniae. For the moment, we continue to use Lactarius in FunKey in a broad sense (inclusive of Lactifluus and Multifurca) pending multilocus analysis of a wider range of collections, especially those from Australia.
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