History
From 23–79 AD, Pliny found a specimen belonging to a mushroom kind, known to the Greeks by the name ’Pezica’, which grows without root or stalk. Subsequently, Dillenius (1719) provided a Peziza description and categorized it as discomycetes or the cup-shaped fungal group (Kimbrough 1970). Linnaeus (1753) used the term Peziza in a similar sense. Dillenius (1719) and Linnaeus (1753) classified all the fungi into ten genera, of which two (Elvella and Peziza) accommodated discomycetous specimens at that time.
Persoon et al. (1801) established the first systematic classification of discomycetes, further developed by Fries. In 1821, a systematic classification of discomycetes (via the Friesian classification system) was established with four classes (Coniomycetes, Hyphomycetes, Gasteromycetes and Hymenomycetes) in which discomycetes specimens were included in Hymenomycetes except for Thelebolus which accommodated in Gasteromycetes (Fries 1821). Fries (1821) divided Peziza into 12 tribes, which later raised to generic rank. The Friesian classification was used for 60 years until 1849 when Fries (1821) modified the classification of discomycetes by establishing six families: Helvellaceae, Bulgariaceae, Dermateae, Patellariaceae, Phacidiaceae and Sticteae based on apothecial macromorphology (Kimbrough 1970). De Notaris (1864) listed 26 genera for discomycetes based on the ascal shape, paraphyses morphology, color of ascospores and the nature of excipulum cells. Nylander (1869) employed the iodine reaction for the first time to classify discomycete specimens besides the morphological examination, and his work was published in Monographia Pezizarum fennicarum. In his work, Nylander (1869) segregated Peziza into 25 subgenera (Karsten 1869). Additionally, some genera such as Aleuria, Phialea, Mollisia, Trochila, Patellaria Orbilia, Lachnella and Lachnea were noted.
Fuckel (1869) classified 80 genera of discomycetes into six families following the Friesian familial system based on apothecial micromorphologies and microchemical data as proposed by Nylander (1896). Based on sterile tissues in apothecia and ascospore shapes, Karsten (1871) divided discomycetes into three families, Helvellaceae, Pezizaceae, Phacidiaceae, and seven subfamilies in a more conservative way than Fuckel’s previous work. Crouan & Crouan (1857) did significant work for classifying discomycetes through their first illustration and description of the dehiscence of operculate asci. Subsequently, Cooke (1879) illustrated and described more than 400 discomycetous species based on authentic herbaria. Saccardo (1884) made a breakthrough in the classification of discomycetes based on the size, shape, color and septation of ascospores. In late 1800–1885, Boudier (1907) published the most significant work in the discomycete classification based on the presence or absence of an operculum, the difference of inoperculate and operculate discomycetes, the amyloid test and oil drops of ascospores. In 1889, Saccardo published his volume of Sylloge Fungorum using his previous discomycetes classification system and abandoning the Friesian system. In this work, Saccardo raised many subgenera into generic rank and published more than 200 genera with approximately 3500 species. In 1887–1896 influenced by Saccardo’s classification, Rehm divided discomycetes into two divisions (Pezizaceae and Helvellaceae), in which Pezizaceae segregate into five suborders (Phacidiaceae, Stictideae, Tryblidieae, Dermatiaceae and Pezizeae) (Rehm 1887, Saccardo 1887, Rehm 1896, Rehm 1898). In Rehm’s work, the suborder Pezizeae included Mollisiaceae, Helotiaceae, Eupezizeae and Ascoboloceae. This Rehm’s work was considered the most practicable book for discomycetes identification, however, this work is discredited due to the failure to recognize the operculate and inoperculate terms (Kimbrough 1970).
Durand (1900) introduced the classification of Pezizineae with mostly cup-shaped discomycetes and regarded four families: Pezizaceae, Ascobolaceae, Helotiaceae and Mollisiaceae. Durand (1900) mainly included mostly fleshy and leathery discomycetes in Pezizaceae and Ascobolaceae, respectively, while waxy and membranous discomycetous specimens were accommodated in Helotiaceae and Mollisiaceae. Höhnel (1903, 1909) described numerous authentic discomycetous specimens valuable for natural classification establishment.
Boudier (1885) proposed new discomycetes rank by grouping discomycetes bearing operculum into three tribes (Mitres, Cupules and Lenticules) while the inoperculate discomycetes into three tribes (Clavules, Carnoses and Cyathules). Boudier (1907) broadened his previous classification to include other subdivisions of seven operculate and 12 inoperculate families and raised the subfamily rank from tribes. In his work, Boudier (1907) used macromorphology, micromorphology and cytochemical characters to designate his specimens into tribe ranks, however, his descriptions were not based on authentic materials.
Nannfeldt (1932) critically re-evaluated many discomycetes genera by proposing higher ascomycetes groups, namely Plectoascales, Ascoloculares and Ascohymeniales. Nannfeldt (1932) separated the discomycetes in Ascohymeniales into four orders, Pezizales, Ostropales, Helotiales and Lecanorales. In Nannfeldt (1932), some Helotiales species were transferred to Ostroporales based on long cylindric asci and filiform ascospores, which raised controversy. Also, Nannfeldt (1932) divided Hyaloscyphaceae into three tribes, Lachneae, Hyaloscypheae and Arachnopezizeae, based on the size of the apothecia and the presence of subiculum.
In 1940, the operculate and inoperculate asci were first described by Chadefaud (1940), a significant finding, especially for Sarcoscyphaceae, which have an intermediate apical apparatus between operculate and inoperculate for which Le Gal (1946) suggested the name ‘suboperculate’. The suboperculate discomycetes were later included in the family of Sarcoscyphaceae and Pezizaceae. Subsequently, Le Gal (1947) provided important knowledge on the formation of ornaments and spore walls of operculate discomycetes and proposed families, Aleuriacae, Ascobolaceae, Hellvellaceae, Humariaceae, Morchellaceae and Sarcoscyphaceae. In 1953, after a new regulation about typification was released affecting Sarcoscypha and Velutaria, Korf (1963) proposed new families: Cyttariaceae, Pezizaceae and Sarcoscyphaceae. Berthet (1961) studied discomycetes and examined the nuclei numbers in apothecial structures, croziers formation and/or the presence of imperfect fungi. Dennis (1968) published a book entitled “British cup fungi and their allies”, which relied on Nannfeldt’s work for grouping inoperculate fungi (Nannfeldt 1932) and several other compilations on operculate discomycetes (Boudiers 1907, Le Gal 1946, 1947). Subsequent works on regional discomycetes were written by mycologists, such as Gamundi (1956, 1957, 1959, 1964) on discomycetes of Argentina; Eckbald (1957) wrote Sarcoscyphaceae of Norway; Thind & Batra (1957), Thind et al. (1958), Thind & Waraitch (1966) on operculates and inoperculates in India; Batra (1961a, b), Batra & Batra (1963) on Indian discomycete flora; Korf (1963) on status and scope of discomycetes in India, and Rifai (1968) on the Australian Pezizales.
Gäumann (1964), followed by Rifai (1968), confirmed the monotypic status of Pyrenomataceae, while Eckbald (1968) reevaluated the taxonomy and phylogeny of operculate discomycetes. However, the most significant study on operculate discomycetes was conducted by Eckblad (1968) by dividing Pezizales into Ascobolaceae, Helvellaceae, Morchellaceae, Otideaceae, Pezizaceae, Pyronemataceae, Rhizinaceae, Sarcoscyphaceae and Thelebolaceae. The most accepted study on Ascomycota was Dennis (1968) entitled ‘British Ascomycetes’, which includes five orders of discomycetes: Helotiales, Lecanorales, Pezizales, Phacidiales and Ostropales. Kimbrough (1970) published his work on the most current discomycetes classification at that time in which he divided inoperculate divisions into orders Helotiales, Ostropales and Phacidiales.
In the 19th century, advances in molecular studies with the DNA barcoding approach revolutionized studies on taxonomy and phylogeny of discomycetes. Landvik & Erikson (1994) documented the parsimonious tree of discomycetes, followed by Gargas & Taylor (1995), Landvik (1996) and O’Donnell (1997). These studies exhibited a paraphyletic assemblage for apothecial specimens, of which Helotiales and Pezizales were placed in distinct clades. Phylogenetic studies focused on discomycetes and Pyrenomycetes were then conducted by Lutzoni et al. (2004), James et al. (2006), Spatafora et al. (2006), Wang et al. (2006a,b) and Schoch (2009a,b). Jacklitsch (2016) incorporated discomycetes taxa in his Ascomycota classification. Afterwards, Ekanayaka et al. (2018) summarized all classes comprising apothecial bearing species and further updated discomycetous taxa based on multi-gene phylogeny and morphology by Ekanayaka et al. (2019a), Johnson et al. (2019), Pfister & Healy (2021) and Quandt & Haelewaters (2021), leading to the current classification of Ascomycota (Wijayawardene et al., 2022)
Recent Genus
RutstroemiaXeropilidium
Ionomidotis
Recent Species
Encoelia furfuraceaRutstroemia tiliacea
Xeropilidium dennisii