Pseudoplectania Melaena (Pezizales): Taxonomical and Nomenclatural Note

Pseudoplectania melaena (Pezizales): taxonomical and nomenclatural note

Nicolas VAN VOOREN Summary: Pseudoplectania melaena (Fr.) Sacc. (= P. vogesiaca) is a widespread species, growing on dead Gilbert MOYNE wood of conifers (Abies sp. in Europe), during spring, in mountainous areas. In some regions, it is considered Matteo CARBONE as rare or uncommon, that’s why we propose a full description of this taxon based both on recent collections Jean-Marc MOINGEON and ancient specimens housed in Persoon’s herbarium (L). Taxonomical and nomenclatural aspects are pro- vided, and a lectotype is designated. Although limited, the risk of confusion with other species of the genus is discussed. Ascomycete.org, 5 (1) : 47-52. Keywords: Ascomycota, Sarcosomataceae, Pseudoplectania vogesiaca, Plectania, taxonomy. Janvier 2013 Mise en ligne le 03/01/2013 Résumé : Pseudoplectania melaena (Fr.) Sacc. (= P. vogesiaca) est une espèce largement répandue, poussant sur le bois mort de conifères (Abies sp. en Europe), durant le printemps, dans les zones montagneuses. Dans certaines régions, elle est considérée comme rare ou peu fréquente, c’est pourquoi nous proposons une description complète de ce taxon basée sur de récentes récoltes et également sur la révision de spécimens an- ciens conservés dans l’herbier Persoon (L). Des aspects taxinomiques et nomenclaturaux sont également fournis, et un lectotype est désigné. Bien que limité, le risque de confusion avec les autres espèces du genre est évoqué. Mots-clés : Ascomycota, Sarcosomataceae, Pseudoplectania vogesiaca, Plectania, taxinomie.

Riassunto: Pseudoplectania melaena (Fr.) Sacc. (= P. vogesiaca) è una specie ben diffusa, fruttificante in pri- mavera su legno morto di conifera (Abies sp. in Europa) in ambiente montano. Considerato che in alcune re- gioni viene considerata rara o poco comune, abbiamo deciso di proporre una dettagliata descrizione di questo taxon basata su raccolte recenti e quelle antiche presenti nella collezione di Persoon (L). Vengono for- niti inoltre gli aspetti tassonomici e nomenclaturali, e viene designato un lectotypus. Sebbene limitato, il ri- schio di confusione con le altre specie del genere viene discusso. Parole chiave: Ascomycota, Sarcosomataceae, Pseudoplectania vogesiaca, Plectania, tassonomia.

Introduction ≡ Peziza vogesiaca Moug. & Nestl., Stirp. Crypt. Vog.-Rhen., 6, no. 584 (1818); Peziza melanea var. vogesiaca (Moug. & Nestl.) Pers., Mycol. Europ., 1: 239 (1822) [“melania”3]. Pseudoplectania melaena (Fr. : Fr.) Sacc. is rather frequently cited in = Peziza spongiosa Peck, Bot. Gaz., 5: 35 (1880), after SEAVER (1913) literature on discomycetes either with this epithet or with the sy- and PADEN (1967). nonym combination Pseudoplectania vogesiaca (Moug. & Nestl.) Sea- ≡ Bulgaria spongiosa (Peck) Peck, Bot. Gaz., 6 (7): 240 (1881); ver. It is mainly characterized by a medium size blackish cup with a Pulparia spongiosa (Peck) Sacc., Syll. fung., 8: 612 (1889). well-defined stipe and microscopically by the rounded spores and the bent to coiled tips of the paraphyses. Description Apothecia at first deeply cupulate then spread, 20–60 mm in dia- Material and methods meter, shortly stipitate or, in some cases, more elongated, totally black and shiny on young specimens, then hymenium dark grey to The description is based on fresh material and synthesized from grey with olivaceous tinges, outer surface black, wrinkled. Margin several collections. Microscopic characters were observed with light eroded, darker than hymenium, hairy. Stipe more or less cylindri- microscopes, in water, especially pigmentation and spore size. 20 or cal, 5–10 (20) × 5 mm, flexuous, smooth, black. Flesh quite elastic, 30 spores were measured on each studied collection. greyish or olivaceous grey. Asci cylindrical, 320–450 × 11–13 μm, with a long attenuate base, aporynchous, octospored, inamyloid. Paraphyses filiform, 1.5–2 μm Taxonomy wide, septate, branched in lowest part, more or less diverticulated or curved to hooked at the apex or with nodules, greyish, up to 3– Pseudoplectania melaena (Fr. : Fr.) Sacc., Syll. fung., 8 : 165 (1889). 6 (–8) μm wide at the top. Hymenial hairs cylindrical, 3 (–4) μm ≡ Peziza melaena Fr., Syst. mycol., II: 60 (1822); Melascypha me- wide, with a single septum at the bottom, brownish, straight to cur- laena (Fr. : Fr.) Boud.1, Icones mycol., liste prélim.: 3 (1904); Plectania ved at the apex. Spores spherical, (11.5–) 12–13 μm in diameter, melaena (Fr. : Fr.) Paden, in Korf, Mycotaxon, 14 (1): 1 (1982). hyaline, with a quite thick wall, smooth, containing small guttules. = Peziza melanea Pers., Mycol. Europ., 1: 239 (1822)2, pro parte. Medullary excipulum of textura intricata, with hyaline hyphae with ≡ Aleuria melanea (Pers.) Gillet, Champ. Fr. Discom.: 37 (1879) an incrusted greyish pigment. Ectal excipulum of textura angularis [“melania”]. with thick-walled cells, brownish to black-brown especially in the = Pseudoplectania vogesiaca (Moug. & Nestl.) Seaver, Mycologia, most external layer. Hairs inconspicuous to abundant in places, 80– 5 (6): 300 (1913). 280 × 5–8 μm, mostly shorter than 100 μm long, straight to very

1 BOUDIER (1885) designated Peziza melaena as the type species of his genus Melascypha. 2 Although PERSOON (loc. cit.) did not provide any description of the taxon Peziza melanea, it is understood as an encompassing definition of the published varieties — P. melanea var. vogesiaca, var. fuscocana and var. sphagnophila — and the valid publication of the first variety generates automatically the auto- nym Peziza melanea var. melanea (art. 26.3 of ICBN). 3 After examining the label of Persoon’s collection no. 910.256-832 on which the name Peziza melanea is distinctly written, we are convinced that the epithet melania inscribed on page 239 in his Mycologia Europaea, sectio prima (PERSOON, 1822), is a typographical error that must be corrected (art. 60.1 of ICBN). 47 slightly curved, septate, brown or bistre-brown but with paler tips at giosa (Mexico), A. balsamea and A. grandis (USA, Canada). It is also re- the longer ones, with a thick wall, smooth, with dark setulae at their ported on Thuja plicata in the USA by PADEN (1983: 180-181). It is im- base. portant to note that, in many references found in literature, the simple information “on decaying conifer wood’’ is also rather com- Studied material: FRANCE. Doubs, Levier, route des Sapins, on mon. dead, decaying branch of Abies sp., 2.04.1978, leg. et det. G. Moyne. Doubs, Levier, route forestière du Rondé, 700 m a.s.l., on dead branches of Abies sp., 4.04.2008, leg. et det. J.-M. Moingeon, G. Moyne Distribution: In Europe, the species is recorded in the following & N. Van Vooren (pers. herb. NV 2008.04.02). Ibidem, 25.03.2010 and countries: Austria, Bosnia and Herzegovina, Bulgaria, Czech Repu- 10.03.2010, leg. et det. J.-M. Moingeon. Doubs, Esserval-Tartre, 820 m blic, Estonia, France, Germany, Italy, Poland, Russia, Slovakia, Spain, a.s.l., on dead trunk of Abies sp., 26.01.2012, leg. et det. J.-M. Moin- Sweden and Switzerland. Outside Europe, P. melaena is known in geon. ITALY. Veneto, Belluno, Pian Cansiglio, 1030 m a.s.l., on mossy the Americas (Canada, USA and Mexico) and in Asia (Japan, Taiwan dead wood of Abies alba, 11.05.2002, leg. et det. E. Campo (TUR-A and China). 145735). Veneto, Belluno, Pian Cansiglio,Vallorch, on mossy dead wood of Abies alba, 24.04.2006, leg. et det. E. Campo (TUR-A 176859). Systematics: Although the wide concept of Plectania Fuckel pro- Veneto, Belluno, Bosco del Cansiglio, on mossy dead wood of Abies posed by PADEN (1983) — where all species of the genus Pseudo- alba, 25.03.2012, leg. et det. E. Campo & M. Carbone (MCVE 27433). plectania were included in a section Sphaerosporae —, was followed Ecology: P. melaena seems to be always associated with dead by many authors, HARRINGTON et al. (1999: 46) confirmed that the two wood of Abies, mainly A. alba in Europe, although records have been genera should be separated. These first phylogenetic results were made on A. nordmanniana and A. holophylla. Outside Europe, col- finally validated by CARBONE et al. (2013) with several sequenced taxa, lections have been registered on Abies kawakamii (Taiwan), A. reli- including Pseudoplectania melaena.

Fig. 1 – Pseudoplectania melaena. Microscopic characters of the collection NV 2008.04.02. A. Ascus, paraphyses and hymenial hair (on left). B. Spores. C. External hair. Scale bars = 10 μm. Drawing: N. Van Vooren. 48 Pseudoplectania melaena Young and immature specimens Picture: J.-M. Moingeon

Pseudoplectania melaena Mature specimens Picture: J.-M. Moingeon

Pseudoplectania melaena Picture: M. Carbone

49 Nomenclature and typiﬁcation considered as the one housed in Persoon’s herbarium at the Uni- versity of Leiden (under no. 910.256-833) as it can be seen from the communication in Mougeot’s handwriting. Although we cannot Although two epithets are currently used to designate this fun- prove if these samples constitute part of the original material, this gus, the correct name is incontestably Pseudoplectania melaena. collection is unquestionably a sort of “pattern” that Mougeot used to Peziza melaena was published by FRIES (1822: 60) who cited Peziza publish the new species Peziza vogesiaca. This is why we think it is vogesiaca as a synonym and, after the article 15.1 of ICBN, names important to present its revision. sanctioned by Fries cannot be replaced by “earlier homonyms and competing synonyms”. Concerning the typification of this name, according to the article Collection no. 910.256-833 (Barcode L0116888) 7.8 of ICBN “anything associated with the name” published in a sanc- As written above, this collection was sent to Persoon by Mougeot. tioning work can be used to typify this name. In the case of Peziza It consists of three very well preserved black, discoid, stipitate apo- melaena, FRIES (loc. cit.) cited three references: i) a pre-Linnean name, thecia, up to 3 × 4 cm in diameter. The microscopic study has re- ii) Peziza vogesiaca Moug. & Nestl.4 and iii) Peziza melanea Pers. (ut vealed the following characters: Spores globose 11.5–13 μm, “melainia”). As no type can be attached to Peziza melaena (no plate smooth, with granular content, walls ca. 1–1.5 μm thick. Asci cylin- is referenced and no material is located in the Fries herbarium at drical, ca. 330 × 10–12 μm, inamyloid, operculate, 8-spored, with UPS), and because P. melanea var. melanea in the sense of Persoon walls slightly thickened and with a flexuous, tapered, aporynchous is not clearly defined, we think that its type must necessarily be cho- base. Paraphyses very difficult to see, but those seen were cylin- sen through the typification ofPeziza vogesiaca. The latter has been drical, on av. 2–3 μm in diam., septate, with curved to slightly coiled published, under the no. 584, in fasc. 6 of the Stirpes Cryptogamae tips; brownish in the upper half due to an extracellular amorphous Vogeso-Rhenanae, a series of exsiccata distributed in several herba- pigment. Hymenial hairs very difficult to see but the very few seen ria (PFISTER, 1985: 110). Each distribution of this collection no. 584 is were cylindrical, on average 3 μm diam., with one septum present considered as part of “original material” of P. vogesiaca, but it seems only in the basal part; their tips were straight to slightly curved and impossible to know if each part belongs to the same collection — the upper part was immersed in the same extracellular amorphous in the sense of art. 8.2 of ICBN — and, even in this case, which would pigment of the paraphyses. Subhymenium of thick textura intricata be the holotype. These exsiccata can be found probably in 16 her- with hyphae cylindrical, septate; uniformly brown to brownish at baria in the world (G, S, etc.) or more and the label associated with low magnification.Medullary excipulum of textura intricata with the material does not contain any information of help (see fig. 2). hyphae cylindrical, hyaline, septate, av. 4 μm diam. but up to 7–8 μm Because of this situation, and in accordance with article 9.2 of ICBN, in some parts, very slightly thick-walled (up to 0.5 μm). Ectal exci- we have to designate the lectotype among the existing material at- pulum of textura globulosa~angularis to angularis with cells up to tached to the protologue: Peziza vogesiaca Moug. & Nestl., lectotypus 25 μm long or wide, with thickened walls; dark brown mainly for an hic designatus: exs. no. 584 in Stirpes Cryptogamae Vogeso-Rhenanae, epimembranaceous and extracellular pigment. External hairs very in herbario Genavense conservatur (G). inconspicuous and difficult to find, but those seen were cylindrical to slightly tapered upwards, 5–8 μm diam., on average less than 100 μm long, smooth, walls up to 1 μm thick, septate, mostly straight or also very slightly curved; light brown for an epimembranaceous pigment.

Collection no. 910.256-832 (Barcode L0116889) This collection is made up of five very well preserved black, discoid and stipitate apothecia. According to the annotation slips present in the herbarium voucher, it seems that they were collected in the Vosges mountains. The microscopic observation revealed they are identical to the previous collection. Discussion

Until the beginning of the XXth century, all authors used the Frie- sian name: COOKE (1876: 112), BOUDIER (1885: 103, 1905-1910: 191), SACCARDO (1889: 165), REHM (1896) and MIGULA (1913: 971). Only GIL- Fig. 2 — Peziza vogesiaca. Lectotype. Collection no. 584 (Stirpes Crypt.) LET (1879: 37) applied the epithet “melania” proposed by PERSOON from herbarium G. (1822: 239). However, the American mycologist Seaver proposed to restore the name Peziza vogesiaca (due to its anteriority) combining it in the genus Pseudoplectania (SEAVER, 1913: 300). He was not fol- Revision of Peziza vogesiaca in Persoon’s herba- lowed by several European authors such as GRELET (1938: 80), FAVRE (1948: 30), IMBACH (1949: 1), during the first half of the XXth century. rium In the second part of this century, P. vogesiaca was more often cited or used partially in literature: MOSER (1963: 121), KRIEGLSTEINER (1975: As mentioned above and illustrated on fig. 2, the name Peziza vo- 181), DENNIS (1978: 78), BREITENBACH & KRÄNZLIN (1981, no. 123) and gesiaca was suggested by Persoon in a letter (in litt.) sent to Mou- CETTO (1987, no. 2112), whilst KREISEL (1962: 75), RAITVIIR (1965: 532), geot after the examination of a collection made by the latter in the PADEN (1967: 179), OTANI (1973: 414), RYMAN (1979: 328) and SVRČEK Vosges mountains (north-eastern France). This collection could be (1981: 74) kept the correct name. It seems that since the revision of

4 Regarding the authority of the epithet, in the Stirpes we have seen the indication “Pers. in litt.” but, according to art. 46.7 of ICBN (ex. 36) there is no inter- nal evidence that Persoon had supplied the validating description. Authorship is therefore to be cited as “Mougeot & Nestler” or, at least, “Persoon ex Mou- geot & Nestler” because the 6th fascicle was published only by Mougeot & Nestler (PFISTER, 1985: 110). 50 Fig. 3 — Collections of Peziza melaena var. vogesiaca from Persoon’s herbarium in L.

Boudier’s name published in Icones Mycologicæ made by KORF (in Smardaea/Greletia, Sowerbyella. Zeitschrift für Mykologie, 71 (2): BRUMMELEN et al., 1985) and probably as a consequence of the ICBN 121-164. adopted during the Congress in Sydney in 19815, the epithet me- BOUDIER E. 1885. — Nouvelle classification naturelle des discomy- laena has been definitely applied: ONADINID (1987: 218), CHMIEL & cètes charnus connus généralement sous le nom de Pézizes. Bul- SAŁATA (1988), IMASEKI et al. (1988: 552), TYLUTKI (1993: 26), DISSING & letin de la Société mycologique de France, 1, p. 91-120. ECKBLAD (2000: 126), MEDEL & CHACÓN (2000: 12), BENKERT (2005: 124), BOUDIER E. 1905-1910. — Icones mycologicæ ou Iconographie des PÉREZ-DE-GREGORIO et al. (2009: 57), DIMITROVA & GYOSHEVA (2009: 24) and champignons de France. Paris, Paul Klincksieck, 4 vol. MOYNE & MOINGEON (2010: 42), with the exception of WANG (2011: 55) BOUDIER E. 1907. — Histoire et classification des discomycètes d’Europe. and TRUDELL & AMMIRATI (2009: 293) who used the epithet vogesiaca Paris, Paul Klincksieck, 222 p. (although the latter indicated “better known in North America as BRUMMELEN J. (van), KORF R.P., CLÉMENÇON H., JÜLICH W. & DEMOULIN V. Pseudoplectania melaena”). ITURRIAGA et al. (2012) strangely consider 1985. — Icones Mycologicæ par Émile Boudier. Tome V : liste préli- P. vogesiaca and P. melaena as two different species. minaire et explication des planches. Lausanne, Piantanida, 300 p. BREITENBACH J. & KRÄNZLIN F. 1981. — Champignons de Suisse. Tome 1. The characters of Pseudoplectania melaena are distinctive enough Les Ascomycètes. Lucerne, Mykologia, 310 p. (phenology, habitat, size, microscopic features) for not being confu- CARBONE M., AGNELLO C. & ALVARADO P. 2013. — Phylogenetic studies in sed with other members of the genus: P. nigrella (Pers.) Fuckel has the family Sarcosomataceae (Ascomycota, Pezizales). Ascomy- smaller, entirely black, stalkless apothecia and grows on spruce lit- cete.org, 5 (1): 1-12. ter; P. sphagnophila (Pers.) Kreisel seems to be even smaller, with a CETTO B. 1987. — I funghi dal vero. Vol. 5. Trento, Arti Grafiche Satur- nia, 722 p. very small pseudostipe and grows in bogs among Sphagnum; P.eri- CHMIEL M.A. & SAŁATA B. 1988 [1986]. — Grzyby z rodzaju Pseudoplec- cae Donadini is sessile and grows under Ericaceae species in the Me- tania Fuck. w Polsce. Folia Societatis Scientiarum Lublinensis, 28 (1): diterranean area. Further Pseudoplectania species are unknown in 3-10. Europe. Among these latter, the only ones reported to be stipitate COOKE M.C. 1876. — Mycographia, seu icones fungorum. Vol. I. Disco- are: P. carranzae (Calonge & M. Mata) M. Carbone, Agnello & P. Alva- mycetes. London, Williams and Norgates, part. 3: 87-136 + pl. 41- rado and P. ryvardenii Iturr., Mardones & H. Urbina. However, both 60. protologues describe two very different species if compared toP. DENNIS R.W.G. 1978. — British Ascomycetes. Vaduz, J. Cramer, 585 p. + melaena. plates. DIMITROVA E. & GYOSHEVA M. 2009. — Bulgarian Pezizales: diversity, dis- Acknowledgements tribution and ecology. Phytologia Balcanica, 15 (1): 13-28. DONADINI J.-C. 1987. — Étude des Sarcoscyphaceae ss. Le Gal (1). Sar- We thank Jacques Melot for his assistance on some nomenclatu- cosomataceae et Sarcoscyphaceae ss. Korf. Le genre Pseudoplec- ral points. We thank Jukka Vauras (TUR-A Herbarium) for the loan of tania emend. nov. P. ericae sp. nov. (Pezizales). Cytologie et some collections. M. Carbone also thanks Prof. G. Thijsse, chief col- scanning des asques. Mycologia Helvetica, 2: 217-246. lection manager of the Leiden Herbarium (L), for the kindness and FAVRE J. 1948. — Les associations fongiques des hauts-marais juras- hospitality showed during his trip. siens et de quelques régions voisines. Beiträge zur Kryptogamen- flora der Schweiz, 10 (3): 1-228. FRIES E.M. 1822. — Systema mycologicum, sistens fungorum ordines, References genera et species, huc usque cognitas. Vol. II, 1st part. Gryphiswal- diae, Ernesti Mauritii, 274 p. BENKERT D. 2005. — Beiträge zur Kenntnis einiger Gattungen der Pe- GILLET C.C. 1879. — Les Discomycètes. Champignons de France. Alen- zizales (Ascomycetes): Plectania/Pseudoplectania, Ramsbottomia, çon, chez l’auteur, 230 p. + planches

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Nicolas Van Vooren 36 rue de la Garde 69005 Lyon France [email protected] Gilbert Moyne 12 rue Radieuse 25000 Besançon France [email protected] Matteo Carbone Via Don Luigi Sturzo 173 16148 Genova Italy [email protected] Jean-Marc Moingeon 2 rue de la Corvée 25520 Goux-les-Usiers France [email protected]