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Resupinatus Gray, Nat. Arr. Brit. Pl. (London) 1: 617. 1821

= Asterotus Singer, Mycologia 35: 161. 1943 (according to Singer, 1975)

= Calathinus Quél., Enchir. fung. (Paris): 46. 1886 (according to Thorn and Barron, 1986)

= Phyllotus Karst., Bidr. Finl. Nat. Folk 32: 14. 1879 (according to Singer, 1951) = Rhodocyphella W.B. Cooke, Beih. Sydowia 4: 105. 1961 (according to Donk, 1962) = Scytinotopsis Singer nom. nud., Ann. Mycol. 34: 335. 1936 (according to Singer, 1951) = Stigmatolemma Kalchbr., Grevillea 10: 104. 1882 (according to Thorn et al., 2005) = StromatocyphellaW.B. Cooke, Beih. Sydowia 4: 104. 1961 (this study)

Type species: Resupinatus applicatus (Batsch: Fr.) Gray, Nat. Arr. Brit. Pl. (London) 1: 617. 1821.

Characters: those of the tribus; hymenophore lamellate, poroid or cyphelloid,

carpophores attached to the substrate directly via the pileus or by a lateral or eccentric pseudostipe; surface hairs repeatedly dichotomously branched or diverticulate;

cheilocystidia diverticulate.

Distribution: cosmopolitan; some species show geographic patterns of distribution.

Observations: Historically, Resupinatus was restricted to species with lamellate or poroid fruit bodies, and those that were cyphelloid were placed in the genus

Stigmatolemma. The only molecular phylogenetic analysis to date prior to this study showed that Stigmatolemma was paraphyletic within Resupinatus, and so all members of the former genus were transferred into the latter (Thorn et al., 2005). Data from this study support this synonymy, and add the cyphelloid Stromatocyphella as another synonym of Resupinatus. The history of these cyphelloid genera is discussed below. Species in Hohenbuehelia were initially separated from Resupinatus on the basis of morphological characters: the presence of metuloid cystidia (thick-walled, lanceolate, and often crystal-incrusted sterile cells in the hymenium) indicates a species belongs in

Hohenbuehelia where an absence indicates the species belongs in Resupinatus (Singer, 1951; 1975a). Possession of a nematophagous anamorph (previously referred to as

Nematoctonus) was added as a defining character of Hohenbuehelia by Thorn and Barron (1986), who treated in Hohenbuehlia some species that were previously thought to

belong in Resupinatus due to their lack of conspicuous metuloids,such as H. unguicularis. Separation of Hohenbuehelia and Resupinatus has been supported by molecular phylogenetic analyses (e.g., Koziak et al. 2007). There is, however, at least one species of Resupinatus with metuloids (because of which the specimen was originally identified as Hohenbuehelia nigra), but no nematophagous anamorph. The sequence from this culture grouped with the Resupinateae and not in Hohenbuehelia

(Pleurotaceae) so was deposited in GenBank as “Resupinatus sp.” (Alberto et al., 1998; Thorn et al., 2000). Asterotus was once considered a separate genus from Resupinatus due to the unusual morphological characters: fruit bodies are stipitate and spoon-shaped, rather than astipitate, orbicular, and dorsally or laterally attached, and surface hairs are digitate to dichotomously branched rather than diverticulate (Singer, 1943). Singer (1975) later changed his mind and included Asterotus within Resupinatus, and this synonymy is supported by the placement of a sequence of the type species, Asterotus dealbatus, within Resupinatus (Moncalvo et al., 2002; Thorn et al., 2005; this study).

Stigmatolemma Kalchbr., Grevillea 10: 104. 1882

= Resupinatus Gray, Nat. Arr. Brit. Pl. (London) 1: 617. 1821. (Thorn et al., 2005; this study)

Type species: Stigmatolemma incanum Kalchbr., Grevillea 10: 104. 1882 Resupinatus incanus (Kalchbr.) Thorn, Moncalvo & Redhead, in Thorn et al.,

Mycologia 97(5): 1148. 2006

Observations: until Talbot’s re-description of the type species (Talbot, 1956) the status

of the genus was considered doubtful (Donk, 1962). The original description by

Kalchbrenner was very brief (12 words), but did designate a type species (Kalchbrenner, 1882). Both Singer (1962) and Donk (1959) were satisfied with Talbot’s re-description after having also seen the small fragment of the holotype of Stigmatolemma incanum in BPI.

Donk (1959) also remarked that the type species of this genus was likely to be closely related to Peziza conspersa (now Resupinatus conspersus) and that this new cyphelloid genus showed remarkable micromorphological similarity to the agaric genus

Resupinatus, a suggestion that was also made by Pilát (1935), Romagnesi (1950), Kühner and Romagnesi (1953), and supported by Singer (1962). This suggestion that species with a reduced morphology were closely related to species with what was regarded to be a more “advanced” fruit body morphology (such as having gills, teeth, or pores) was not widely adopted in fungal classifications until the first molecular studies demonstrated this to be true (e.g., Hibbett and Binder, 2002; Bodensteiner et al., 2004). This study further demonstrates that species with cyphelloid fruit body morphologies are multiply derived within groups of species with lamellate or poroid fruit bodies (see Figure 2.1). This study therefore supports that Stigmatolemma is a synonym of Resupinatus.

Stromatocyphella W.B. Cooke, Beih. Sydowia 4: 104. 1961. = Resupinatus Gray, Nat. Arr. Brit. Pl. (London) 1: 617. 1821.

Type species: Stromatocyphella conglobata (Burt) W.B. Cooke, Beih. Sydowia 4: 104. 1961

Cyphella conglobata Burt, Ann. Mo. bot. Gdn 1: 375. 1915.

Observations: at the time of its description, this genus originally contained three species. Reid (1964) reported that two of these (Stromatocyphella lataensis and Stromatocyphella aceris) do not belong in the genus, and are synonyms of Calythella erucaeformis and Cyphellopsis anomala, respectively. One other species, Cyphella subgelatinosa, may be a close relative. This species shares the common stroma-like hyphal mass, but has cups in smaller groups. A full discussion of Cyphella subgelatinosa can be found in Chapter 5.

There has also been disagreement in the literature about the correct classification of Stromatocyphella conglobata. In the type description, Burt (1914) commented that this species superficially resembles Cyphella fasciculata (now Merismodes fasciculatus, a member of the Cyphellaceae), but suggested that it might deserve a new genus. Singer (1945) supported this idea, and Cooke (1961) created the genus Stromatocyphella, which was supported by Donk (Donk, 1964). Unfortunately, Singer (1973) also stated that this species displayed strong affinities to Campanella and Flavolaschia, and so

Stromatocypella has been classified with these genera in the Marasmiaceae ever since (as per Kirk et al., 2008), despite its morphology which places it as a close relative of

Resupinatus (Singer, 1945, 1986). Molecular data (see Figure 2.1) now support its inclusion within Resupinatus.

Rhodocyphella W.B. Cooke, Beih. Sydowia 4: 105. 1961. = Resupinatus Gray, Nat. Arr. Brit. Pl. (London) 1: 617. 1821.

Type species: Rhodocyphella cupuliformis (Berk. & Ravenel) W.B. Cooke, Beih. Sydowia 4: 105. 1961.

Cyphella cupuliformis Berk. & Ravenel, Grevillea 2: 5. 1873 (as “cupulaeformis”).

Chaetocypha cupuliformis (Berk. & Ravenel) Kuntze, Rev. Gen. pl. (Leipzig) 2: 847. 1891.

Observations: this genus is a synonym of Resupinatus. The name Rhodocyphella cupuliformis was originally used for all collections of cyphelloid fruit bodies lacking a subiculum found on Juniperus, regardless of geographic origin (Cooke, 1961), and was then treated as a synonym of Resupinatus taxi (Gilbertson & Blackwell, 1984), which is a later name for Resupinatus griseopallidus (Elborne, 2008). This is incorrect, as North American collections on Juniperus are Rhodocyphella cupuliformis, while European collections on a variety of substrates are Resupinatus griseopallidus (an unrelated species). Rhodocyphella cupuliformis has an identical sister species (Australian

cyphelloid Resupinatus 1) found only in New Zealand on Dacrycarpus which requires a new name. The type species of this genus is discussed in further detail in section 2.4 “Taxonomy” (below), the Australian taxon (Australian cyphelloid Resupinatus 1) in Chapter 4, and Resupinatus griseopallidus in Chapter 5.

One other species, Rhodocyphella grisea, may be a close relative or even a synonym of Resupinatus griseopallidus as it is morphologically nearly identical, but was found in Sri Lanka. Unfortunately, it is only known from the type collection, precluding a molecular analysis. A more thorough description of this species may be found in Chapter 5. Rhodocyphella cupuliformis is an example of what would have traditionally been

considered an “intermediate” form between the lamellate species of Resupinatus and the cyphelloid gregarious species of Stigmatolemma: fruit bodies resemble Resupinatus in occuring singly or in small groups and lacking a subiculum, but are cyphelloid as in Stigmatolemma. The molecular data show that this species falls within Resupinatus (see Figure 2.1), making Rhodocyphella a synonym of Resupinatus and a member of the Resupinateae.