TAXON 70 (2) • April 2021: 429–430 Vogan & al. • (2803) Change conserved type of Podospora

PROPOSALS TO CONSERVE OR REJECT NAMES Edited by John McNeill, Scott A. Redhead & John H. Wiersema (2803) Proposal to change the conserved type of Podospora, nom. cons. () Aaron A. Vogan,1 Andrew N. Miller2 & Philippe Silar3 1 Systematic Biology, Department of Organismal Biology, Uppsala University, Norbyvägen 18D, 752 36, Uppsala, Sweden 2 Illinois Natural History Survey, University of Illinois Urbana-Champaign, Champaign, Illinois 61820, U.S.A. 3 Université de Paris, Laboratoire Interdisciplinaire des Energies de Demain (LIED), 75006, Paris, France Address for correspondence: Aaron A. Vogan, [email protected] DOI https://doi.org/10.1002/tax.12478 First published as part of this issue. See online for details.

(2803) Podospora Ces. in Rabenhorst, Klotzschii Herb. Mycol., 1849) and Malinvernia Rabenh. (in Hedwigia 1: 116. 1857), and their ed. 2: No. 259 (vel 258). Jan–Jun 1856, nom. cons. types and nomenclatural statuses. The history of the typifications of Typus: P. anserina (Rabenh.) Niessl in Hedwigia 22: 156. Podospora and Schizothecium themselves are relevant and will be 1883 (Malinvernia anserina Rabenh. in Hedwigia 1: 116. discussed in more detail below. 1857), typ. cons. prop. The recent placement of one species in particular, , in a separate , Triangularia Boedijn (in Ann. Mycol. The generic name Podospora is currently conserved against 32: 302. 1934), by Wang & al. (in Stud. Mycol. 93: 155–252. 2019) Schizothecium Corda (Icon. Fung. 2: 29. Jul 1838), and the history precipitated our proposal. Podospora anserina is a model filamentous of the conservation of the name Podospora is long and convoluted. that has served as a workhorse for molecular biology and genet- Attempts were made in 1910 to conserve Podospora (Saccardo in ics for over a century. It has been central to the fields of senescence, Briquet, Rec. Doc. Sect. Nomencl. Congr. Bruxelles: 46. 1910) and heterokaryon incompatibility, sexual reproduction, and prion biology, in 1930 to reject Podospora (Maire in Briquet, Rec. Syn. Cambridge: among others, providing one of the most tractable models to study 118. 1930). Podospora also has a conserved non-original type, these complex phenomena. As a result, it is the focal organism of P. fimiseda (Ces. & De Not.) Niessl (in Hedwigia 22: 156. Oct numerous laboratories and has provided the basis for thousands of 1883), based on Sordaria fimiseda Ces. & De Not. (in Comment. research articles (e.g., Stahl & al. in Molec. Gen. Genet. 162: 341–343. Soc. Crittog. Ital. 1: 226. Jan 1863), which has its own conserved type 1978; Cummings & al. in Molec. Gen. Genet. 171: 239–250. 1979; specimen: Rabenhorst, Klotzschii Herb. Mycol., ed. 2: No. 259 (S). Coustou & al. in Proc. Natl. Acad. Sci. U.S.A. 94: 9773–9778. An image of the exsiccata label with a full description of the genus 1997; Dufour & al. in Proc. Natl. Acad. Sci. U.S.A. 97: 4138–4143. and species was reproduced by Braun (in Schlechtendalia 35: 8, fig. 2000; Krause & al. in J. Biol. Chem. 279: 26453–26461. 2004; Espagne 22. 2018). The type of Schizothecium is S. fimicola Corda (l.c.). & al. in Genome Biol. 9: 1–22. 2008; Grognet & al. in PLOS Genet. Schizothecium is no longer considered to be taxonomically syn- 10: e1004387. 2014). The fungal genus Podospora encompasses a onymous with Podospora as typified. Lundqvist (in Symb. Bot. Upsal. large number (127 currently accepted names in Species Fungorum, 20: 1–374. 1972), in pre-molecular taxonomy, recognized both genera http://www.speciesfungorum.org/names/Names.asp) of diverse, often separately. In more recent literature, based on modern DNA sequence- coprophilous, species united taxonomically by the morphology of based analyses, Schizothecium and Podospora are well separated, their ascospores. Their mature ascospores possess a dark, pigmented and in fact, Schizothecium is the type of the newly named family, head with a hyaline pedicel and often gelatinous caudae on both ends Schizotheciaceae Y. Marin & Stchigel (in Microorganisms 8(9, no. (Lundqvist, l.c.). It has long been recognized that these characters do 1430): 24. 2020). Nearly contemporaneously with the recognition not represent synapomorphies within the group and that Podospora, of the Schizotheciaceae was the recognition of the Podosporaceae as currently circumscribed, encompasses many disparate taxa (Miller X. Wei Wang & Houbraken (in Stud. Mycol. 93: 223. 2019), further & Huhndorf in Molec. Phylogen. Evol. 35: 60–75. 2005). Although separating the two genera taxonomically. the genus contains a large number of described species, P. anserina is The history of the existing conservation and rejection entries of the only one that has received any amount of attention in the scien- Podospora and Schizothecium in App. III and Sordaria fimiseda in tific literature beyond original descriptions, taxonomic treatments, App. IV of the ICN (Wiersema & al., Int. Code Nomencl. App. I–VII. and ecological surveys. Podospora has been placed in the Lasiospha- 2018–; https://naturalhistory2.si.edu/botany/codes-proposals) dates eriaceae Nannf. (in Nova Acta Regiae Soc. Sci. Upsal., ser. 4, 8: 50. back to a proposal by Donk (in Regnum Veg. 34: 25–31. 1964) to 1932), which has been found to be paraphyletic within the order Sor- conserve Podospora against Schizothecium following a lengthy ana- dariales Chadef. ex D. Hawksw. & O.E. Erikss. (in Syst. Ascomyce- lysis of several other generic names, including Sordaria Ces. & De tum 5: 182. 1986). Molecular phylogenies have shown that there Not. (l.c.: 225) nom. cons., Pleurage Fr. (Summa Veg. Scand.: 418. exist three or four main lineages that contain species assigned to

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Version of Record 429 Vogan & al. • (2803) Change conserved type of Podospora TAXON 70 (2) • April 2021: 429–430

Podospora (Kruys & al. in Fungal Diversity 70: 101–113. 2015; increasingly blurred as of late, as the same authors who created the Marin-Felix & al. in Microorganisms 8(9, no. 1430): 1–39. 2020). new combination, T. anserina, also claimed that the type of Podos- Recently (Wang & al., l.c.), the clade containing the currently con- pora is P. fimicola, the originally included species, instead of served type of Podospora, P. fimiseda, was named as the new family P. fimiseda, the currently conserved type (Wang & al., l.c). The error Podosporaceae, which is composed of three subclades. Nonetheless, is understandable as the history of the species names is confusing. species named as Podospora remained in all three subclades, inter- The generic name Podospora was originally based on P. fimicola mingled with species of at least seven other recognized genera. The (basionym: Schizothecium fimicola). When the genus Sordaria Ces. three subclades within Podosporaceae were treated as three genera, & De Not. was erected, both Sphaeria fimicola and P. fimicola were to which the names Cladorrhinum Sacc. & Marchal (in Bull. Soc. to be included, and so P. fimicola was given a new epithet as Sordaria Roy. Bot. Belgique 24: 64. 1885), Podospora, and Triangularia were fimiseda. At the same time, the authors doubted whether P. fimicola applied. As P. fimiseda occurs in a different subclade from that of really was the same as Schizothecium fimicola. Currently, it is P. anserina, a new combination was made for the latter, Triangularia believed that Schizothecium fimicola is typical of species related to anserina (Rabenh.) X. Wei Wang & Houbraken (l.c.: 231). Wang P. curvula, and thus Schizothecium is considered the proper name & al. (l.c.) overlooked the generic name Malinvernia Rabenh. (l.c.), for those taxa in modern nomenclature (Cai & al. in Fungal Diversity which has priority over Triangularia and was monotypic when pub- 19: 1–21. 2005) and is now the type of the family name Schizothecia- lished, typified by M. anserina. Additionally, neither the generic name, ceae. When Podospora was resurrected by Niessl (l.c.), he created the Triangularia, nor any of the other generic names that have been new combination, P. fimiseda, which currently serves as type of that included in the Podosporaceae have any amount of noticeability that generic name. even comes close to rivalling P. anserina’s fame; Google Scholar pro- Donk (l.c. 1964: 31) originally proposed conservation of Podos- vides ~11,500 hits for Podospora anserina and only 2110 for Trian- pora typified by P. fimicola against Schizothecium typified by gularia. “Triangularia” is also an adjective in Latin and a verb in S. fimicola. Reference to Donk’s earlier proposal was made by the Portuguese, and there is a fossil Gasteropod, Triangularia F. Frech Committee for Fungi and Lichens (Donk in Taxon 17: 579. 1968) (in Z. Deutsch. Geol. Ges. 46: 446–479. 1894), and, therefore, many without mentioning the type of either generic name. The listing in of the hits do not reference the fungal genus. If the term “fungus” is the Code eventually became “P. fimicola sensu Cesati”. However, appended to the search query, the number of hits plummets to 307. following changes to the Code, Nicolson & al. (in Taxon 33: 314– Moreover since the publication of the paper of Wang & al. (l.c.), 315. 1984) recommended changing the listing to “P. fimiseda (Cesati 10 new articles all using P. anserina have been published. To do as et De Notaris) Niessl” as a conserved type and that matched Cesati’s Wang & al. have proposed would result in a large number of unnec- concept of “P. fimicola”. Later, the Code was further changed to per- essary new combinations and would require either conservation of mit typification of the basionym, Sordaria fimiseda Ces. & De Not. Triangularia against Malinvernia or more combinations in Malin- by the exsiccata specimen Rabenhorst, Klotzschii Herb. Mycol., vernia. Currently, P. fimiseda is the only species of the clade (14 ed. 2: No. 259 (S), which matches the place of publication for the known species) that belongs to Podospora, whereas the clade with generic name, i.e., the exsiccata label for the specimen. P. anserina (27 known species) contains at least six other species of There have also been a number of historical debates surround- Podospora. It is clear that to maintain stability and consistency in ing whether or not Podospora anserina was the correct name of the the literature, the name Podospora anserina should be protected against taxon in question (Traverso, Fl. Ital. Crypt. 2: 431–432. 1907; Mirza all competing names including both Malinvernia anserina and also & Cain in Canad. J. Bot. 47: 1999–2048. 1969; Lundqvist, l.c.), or if Triangularia anserina if Wang or others were to propose conserva- instead Podospora pauciseta (Ces.) Traverso (l.c.: 431) should be tion of Triangularia. The solution of simply synonymizing Triangu- used. Thankfully, this discourse was satisfactorily settled when it was laria under Podospora has been recently proposed (Ament-Velásquez discovered that the known collection of P. anserina strains represents & al. in MycoKeys 75: 51. 2020), which would protect the status of a species complex of seven separate lineages (Boucher & al. in Cryp- P. anserina temporarily. However, synonymizing all of these generic tog. Mycol. 38: 485–506. 2017). The standard laboratory strain S names may require that all species within the Podosporaceae be con- that is at the centre of most published scientific literature was “can- sidered Podospora, and furthermore, as P. fimiseda and P. anserina lie onized” as the ex-epitype culture of P. anserina (Silar in Schlechten- in separate clades, the status of P. anserina will continue to be in jeop- dalia 34: 1–2. 2018), while a separate strain (CBS 237.71) became ardy of future taxonomic changes. the representative of P.pauciseta (Boucher & al., l.c.). Thus, the appli- Furthermore, a number of issues exist regarding the current type cation of P. anserina is no longer in question. Conserving Podos- of Podospora, P. fimiseda, due in large part to the past uncertainty pora with P. anserina as its type would not only preserve a widely regarding the identity of the organisms on which various taxa were used name, but would eliminate much of the confusion involved described. The confusion centers around the names Schizothecium with P. fimiseda. fimicola, (Roberge ex Desm.) Ces. & De Not. (l.c. 226), and Podospora fimiseda, all of which are currently names Author information of the type of their respective generic names, as well as the name AAV, https://orcid.org/0000-0003-2013-7445 P. fimicola (Corda) Ces. (in Hedwigia 1: 103. 1856) (often incor- ANM, https://orcid.org/0000-0001-7300-0069 rectly attributed as “P. fimicola Ces.”). This issue has become PS, https://orcid.org/0000-0003-0104-987X

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