The generic name Abrothallus (Abrothallales

Suija & al. • Abrothallus species on parmelioid hosts

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The generic name Abrothallus (Abrothallales, Dothideomycetes), and names proposed in the genus by Giuseppe De Notaris, Søren Christian Sommerfelt, and Ignaz Kotte Ave Suija,1 David L. Hawksworth2 & Sergio Pérez-Ortega3 1 Institute of the Ecology and Earth Sciences, University of Tartu, 40 Lai street, 51005 Tartu, Estonia 2 Comparative Plant and Fungal Biology, Royal Botanic Gardens, Kew, Surrey TW9 3DS, U.K.; Department of Life Sciences, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K. 3 Real Jardín Botánico, CSIC, Plaza de Murillo 2, 28014, Madrid, Spain Author for correspondence: Ave Suija, [email protected] DOI https://doi.org/10.12705/676.13 Abstract The date of publication of the generic name Abrothallus is discussed, along with typification of the Abrothallus names proposed by Giuseppe De Notaris, Søren Christian Sommerfelt, and Ignaz Kotte. Pertinent historical collections have been examined, and each typication is supplied with a description in order to facilitate correct identifications. The new combination A. santessonii (≡ Vouauxiomyces santessonii) is proposed, and the species description updated to include the sexual morph. Keywords dual nomenclature; lichenicolous fungi; Parmeliaceae

INTRODUCTION The purpose of this contribution is to clarify the date and place of publication of the generic name Abrothallus De Not. (Abrothallales, Dothideomycetes), the issue of the type, as well as the nomenclatural status and typification of several species described in the nineteenth and early twentieth centuries. The genus was originally described as lichenized (De Notaris, 1846, 1849), although De Notaris expressed some doubts and hinted that Abrothallus might be a parasite comparable to two species now placed in Scutula Tul. Montagne (1851) was also sceptical that Abrothallus was a lichenized fungus, and the lichenicolous habit was unequivocably established by the critical microscopic studies of Tulasne (1852) and Lindsay (1857). This obligately lichenicolous genus is characterized by black roughly strongly convex to almost subglobose apothecioid ascomata, often covered with greenish pruina, asci which are bitunicate in structure and fissitunicate, 4–8-spored, and 2–4-celled, brown, asymmetric ornamented ascospores, in some cases separating into part-spores while still in the ascus, and branched and anastomosed interascal filaments. The asexual morph was described under the name Vouauxiomyces Dyko & D.Hawksw. (Hawksworth & Dyko, 1979; Pérez-Ortega & al., 2011), but since the termination of dual nomenclature for morphs of the same fungal species with different reproductive modes, that generic name is treated as a synonym. In fact, Abrothallus is an excellent example of the issues associated with dual nomenclature in fungi because asexual morphs may be found associated with the sexual ones in some species, while other species are found primarily or exclusively as the asexual morph (Tulasne, 1852; Suija & al., 2015).

Species delimitation in Abrothallus has been a subject of long-standing controversy, resulting in differently interpreted species names (e.g., Tulasne, 1852; Lindsay, 1857; Kotte, 1909; Vouaux, 1912–1914; Keissler, 1930; Hawksworth, 1983; Clauzade & al., 1989; Santesson, 1993; Diederich 2004; Santesson & al., 2004; Ihlen & Wedin, 2008). This discrepancy is evident when examining collections in which, under the same species name, one can find specimens growing on the same host genus but with clearly different morpho-anatomical features (Diederich, 2004; Ihlen & Wedin, 2008; Suija & Pérez-Ortega, pers. obs.). The problem is particulaly acute for species that grow on hosts of the lichen family Parmeliaceae (e.g., Diederich 2004; Ihlen & Wedin, 2008). The cosmopolitan family Parmeliaceae (Lecanoromycetes, Ascomycota) is one of the largest and evolutionary youngest families of mainly lichen-forming fungi with complex and variable morphologies (Divakar & al., 2015). During our ongoing studies on the genus Abrothallus (Suija, 2006; Pérez-Ortega & al., 2011, 2014; Suija & al., 2011, 2015), we realized the necessity to correctly interpret names published by earlier authors to clarify the taxonomy of Abrothallus species growing on hosts of the family Parmeliaceae. For this reason, we examined the original historical materials studied by G. De Notaris (1846, 1849), S.C. Sommerfelt (1826) and I. Kotte (1909). We propose a new combination A. santessonii and provide a new description including the sexual stage of the fungus.

MATERIALS AND METHODS This contribution is based on our examination of type and other material deposited in BM, BR, H, Institut für Vegetationskunde und Landschaftsökologie, Hemhofen,

Article history: Received: 5 Apr 2018 | returned for (first) revision: 3 Jun 2018 | (last) revision received: 11 Jul 2018 | accepted: 20 Sep 2018 | published: online fast track, N/A; in print and online issues, 13 Dec 2018 || Associate Editor: James C. Lendemer || © International Association for Plant Taxonomy (IAPT) 2018, all rights reserved

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Germany (not in Index Herbariorum, hereafter IVL), K (incl. IMI), M, NY, O, PC, RO, TU, and UPS. The specimens were examined with microscopes available in the herbaria cited above. Colour reactions were performed with 10% potassium hydroxide (K), 50% nitric acid (N), and Lugol’s (I Lugol) or Melzer’s solution (IMelzer) either without or with pretreatment with K (K/I). When 15 or more measurements are summarized in the description, the measurements are given as minimum value–(average ± standard deviation)–maximum value, followed by the number of measurements (n) given in brackets; minimum value–(average)–maximum value or minimum and maximum values are given when n < 15. The macrophotographs on Fig. 7 were taken with a Leica SP8APO compound stereomicroscope, and the free software CombineZM was used to combine images for increasing the depth of field. The scanning electron micrographs on the same figure were prepared from air-dried material mounted on stubs, coated with thin layer of gold, and observed using a FEI INSPECT Scanning Electron Microscope (MNCN, CSIC, Madrid).

THE GENERIC NAME ABROTHALLUS This generic name has been generally accepted as published by De Notaris in 1845 (Jatta, 1911; Farr & al., 1979; Greuter & al., 1993; Diederich, 2004; Kirk & al., 2011; Suija & al., 2011), although Tulasne (1852) and Lindsay (1857) both referred to two papers of De Notaris (1846, 1849). To further add to the confusion, Santesson (1960) cited De Notaris (1849) as the place of publication of the single originally included species, but did not comment on the place of publication of the generic name. The reason for these discrepancies is that the original account of the genus Abrothallus and the species name A. bertianus De Not. was presented to a meeting of the Real Academia of Torino on 15 June 1845 (Fig. 1), but not published in a printed format until 1849 (De Notaris, 1849). However, there are at least two copies of this publication (in BPI [herb. Stevenson] and G-BOIS) that have been interpreted as

undated preprints of the 1849 paper, set in an identical typography but with the pages numbered 1–4 and not 351–354 as in De Notaris (1849) and an identical plate, and the statement “Exhib. 15 iunii 1845”. There is no doubt that there was an exhibit introducing this new generic name communicated at the Real Academia of Torino on that date from the manuscript records of the Academy, where it is recorded that permission was given to print this in the proceedings (Fig. 1). Stafleu & Cowan (1976) and Leeusink (in Farr & al., 1979) accepted the independently paginated pamphlet as a preprint distributed in 1845, with no indication of doubt. Two facts may support this interpretation. First, De Notaris (1846) attributed the name Abrothallus to “DeNtrs” with no indication of it being in press, and “Charact. auctus” as a source of the description; in contrast, other newly introduced generic names in the same work were given as “nov. gen.” with no indication of De Notaris’s name. This suggests that De Notaris considered the name to be already validly published by at least 1846. Second, the unpaginated copy in G is bound in a volume where it was placed immediately before an article that was published in 1846. What does not support this interpretation is that no further copies of the unpaginated version could be located in the libraries of either the academy or the university in Turin, or in any other major botanical libraries we were able to check. The date of publication of Abrothallus needs to be resolved as it could affect the typification of the genus. Specifically, A. bertianus was the only species included in the genus by De Notaris in the “1845 preprint”, and that name was accompanied by a description. In contrast if De Notaris (1846) was considered to be the place of publication for Abrothallus, in that work A. bertianus was not accompanied by a separate description (it was referred to as published in “DNtrs l. ca.”), and a second name, A. buellianus, was also included. Note that although De Notaris (1846) did not include a description for A. bertianus, three differences were listed between A. buellianus and A. bertianus on page 194 and we consider this to constitute a validating diagnosis for the latter name.

Fig. 1. Excerpts from the proceedings’ protocol of the of Real Academia of Torino from 15 June 1845 showing approval of G. De Notaris’s study about the genus Abrothallus in Memorie della Reale Accademia delle Scienze di Torino (From: Archive of Real Academia of Torino, Italy).

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In the context of the current rules of nomenclature, it is currently only possible to fix the place of publication of names at the rank of family by conservation (Turland & al., 2018, Art. 14.14). In view of the uncertainty in this case, we propose to accept the valid publication of the generic name as from De Notaris (1846), and continue to accept A. bertianus De Not. as type in order to be consistent with the treatments in Farr & al. (1979) and most other authors. The presentation of the 1846 work makes clear that De Notaris intended A. bertianus to serve what we would consider today to be the type of the genus. In contrast, Clements & Shear (1931) listed A. parmeliarum (Sommerf.) Arnold as the type, but that selection cannot be correct as that species was not originally included by De Notaris. The listing of A. parmeliarum as type by those authors almost certainly followed Keissler (1930), who treated A. bertianus as a synonym of A. parmeliarum.

TYPIFICATION OF THE NAMES Abrothallus De Not. in Giorn. Bot. Ital. 2: 192. 1846 – Type: A. bertianus De Not. (implied by De Notaris, l.c.). = Pseudolecidea Marchand, Enum. Méth. Mycophyt.: 159. 1896

Fig. 2. Herbarium sheet with collection of Abrothallus bertianus from De Notaris’s collection in RO.

(“Pseudo-Lecidea”) – Type: P. parmeliarum (Sommerf.) Marchand (≡ Lecidea parmeliarum Sommerf.). = Vouauxiomyces Dyko & D.Hawksw. in Lichenologist 11: 57. 1979 – Type: V. truncatus (B.de Lesd.) Dyko & D.Hawksw. (≡ Phoma truncata B.de Lesd.). Abrothallus bertianus De Not. in Giorn. Bot. Ital. 2: 192. 1846 – Lectotype (designated here, MBT-381109): ITALY. Liguria, ad Olea, circa Porto-Maurizio, [on Melanelixia subaurifera], 1842, Berti s.n. (RO!, middle thallus). = Abrothallus glabratulae I.Kotte in Centralbl. Bakteriol. Parasitenk., 2. Abth. 24: 80. 1909. For images of the lectotype of Abrothallus bertianus, see Figs. 2 and 3; illustration of A. bertianus in De Notaris (1849: 355). Description. – Vegetative hyphae immersed, K/ILugol+ violet [I Melzer + blue]. Ascomata black, globose to slightly appressed, 0.17–(0.24 ± 0.037)–0.3 mm diam. (n = 18), 0.06– 0.1 mm tall, mainly epruinose, only a few ascomata slightly green pruinose (De Notaris, 1849 indicates the presence of green pruina on cited specimens). Epihymenial layer of dark brown granules, K+ greenish to emerald greenish, N+ violet. Hymenium hyaline, ca. 60 µm, upper part K+ greenish. Exciple ca. 55–65 µm. Hypothecium brown below the hymenium, turns gradually lighter towards the base, hypothecial cells rectangular, 4–7 × 3 –5 µm, l/w ratio = 1–1.6 (n = 10). Asci clavate, bitunicate, 47–55 × 8–12 µm, 8-spored. Interascal

Fig. 3. Lectotype of Abrothallus bertianus De Not. (the middle thallus) from De Notaris’s collection in RO.

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filaments irregularly dichotomously branched, 1.5–2 µm wide, tip 2–3 µm. Ascospores 2-celled, asymmetric, verruculose, 11–(12.68 ± 1.19)–16 × 4 –(4.89 ± 0.46)–6 µm (upper cell) × 3.5– (3.97 ± 0.31)–4.5 µm (lower cell), l/w ratio = 2.2–3.5 (n = 20). Asexual morph not observed. Notes. – Santesson (1960) stated that isotypes of Abrothallus bertianus were preserved in VER and RO, but without any further information, and without citation of a holotype or lectotype. We studied De Notaris’s specimens in NY, PC, and RO, and above select as lectotype a specimen from RO collected in 1842 (Fig. 3: middle thallus). This was the only collection cited by De Notaris (1849: 354) in that account, and which was identical to the unpaginated “pre-print”; the collector was also the only person acknowledged at the end of that work. This collection is accompanied by a handwritten description, black and white drawings, two subsequent collections of A. bertianus (both on Melanelixia subaurifera (Nyl.) O.Blanco & al.) and in the uppermost right corner an envelope with A. welwitschii Tul. ex Mont. (Fig. 2) growing on Sticta sylvatica (Huds.) Ach. A De Notaris collection at PC in herb. Montagne (PC0068735) from Italy, Genna (Genova?) is undated, but annotated by De Notaris as Coccocarpia bertiana; the host proved to be Melanelixia subaurifera and is intermixed with Frullania sp. growing on an olive tree. We were not able to directly examine the specimen in NY (barcode 00920130), which was collected near Genova (“p[rope] Genuam”) in 1845 and on Melanohalea olivacea (L.) O.Blanco & al. The description provided for Abrothallus glabratulae by Kotte (1909) agrees well with the description of A. bertianus including the host genus – Melanelixia O.Blanco & al. We tried to locate the two collections of A. glabratulae cited by Kotte (1909: 11) but without success. The size of the ascospores (10.9–13.3 × 4.7–5.2 µm) is within the range of our measurements for A. bertianus (11–16 × 4 –6 µm), and the vegetative hyphae of both species are amyloid. Although De Notaris (1849) described the ascomata of A. bertianus as green pruinose, we found most of his specimens to be epruinose or with only slightly discernible pruina (PC0068735). Kotte (1909) did not mention pruina on ascomata of A. glabratulae. We assume that the pruina disappear or are swept off with age (Suija, 2006); the persistence may also depend on the storage conditions. We did not find any asexual morph in any of the De Notaris A. bertianus specimens in RO, but one was mentioned by Kotte (1909) for A. glabratulae, and by Diederich (1989), Clauzade & al. (1989) and Brackel (2009) for A. bertianus. Detailed descriptions were provided by Kotte (1909) and Diederich (1989), the dimensions of the conidia were given in different papers as 5.46–6.5 × 3.9–4.42 µm (Kotte, 1909), 5–8 × 3–4 µm (Clauzade & al., 1989), (5–)5.5–6.9(–8.5) × (2.8–)3.1–4.1(–4.5) µm (Diederich, 1989), and 5.5–(6.7 ± 0.68)–8 × 4 –(4.85 ± 0.63)– 6.5 µm (IVL4594; cited by Brackel, 2009: 6; measured by AS). Additional specimens examined. – ITALY. Genna, ad truncos Olea, [undated], De Notaris s.n. (PC0068735); ad olea Europea, 20 Mar 1845 (RO). GERMANY. Bavaria, Oberbayern, Kreis Eichstätt, W Wipfenberg, at the wooden tower, 480 m, on Melanelixia glabratula (Lamy) Sandler & Arup, 3 Apr 2008, Brackel s.n. (IVL4594 – asexual morph). 1172

Abrothallus buellianus De Not. in Giorn. Bot. Ital. 2: 192. 1846 – Lectotype (designated here, MBT-381110): ITALY. In montibus Bobliensibus, in thallo parmeliae tiliaceae (Parmelina tiliacea (Hoffm.) Hale) parasiticus, 1846, De Notaris s.n. (RO [specimen in separate envelope]!; isolecto types: NY barcode 00920131!, RO [the leftmost glued thallus with “a”]!). For images of the lectotype, see Figs. 4 and 5. Description. – Vegetative hyphae immersed, K/ILugol –. Ascomata black (not chestnut brown when wet according to De Notaris, 1846), epruinose, flattened to subglobose, 0.2– (0.3)–0.5 mm diam. (n = 9), ca. 0.06 mm tall. Epihymenial layer of brown granules, K+ greenish, N–. Hymenium hyaline, ca. 95 µm high, K–, N–. Exciple ca. 70 µm. Hypothecium brown below the hymenium, turning lighter towards the base, ca. 170 µm tall, hypothecial cells rectangular, 5–8 × 4 –7 µm, l/w ratio = 1–1.6 (n = 10). Asci bitunicate, elongated to subclavate, 8-spored, 53–68 × 11–15 µm. Interascal elements irregularly branched, ca. 1–1.5 µm wide, the tip only slightly widened, 1.5–2 µm. Ascospores 2-celled, brown, verruculose, asymmetric, lower cell usually elongated, 14–(16.5 ± 1.7)–20 × 5.5–(6.1 ± 0.45)–7 µm (upper cell) × 4 –(5.16 ± 0.02)–6 µm (lower cell), l/w ratio = 2.3–3.4 (n = 22). Asexual morph not observed.

Fig. 4. Herbarium sheet with collection of Abrothallus buellianus from De Notaris’s collection in RO. The separate envelope contains the lectotype (see Fig. 5); the leftmost glued thallus marked with “a” is isolectotype.

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Notes. – We examined three specimens deposited in NY, PC and RO for which locality data corresponded to that in the protologue (De Notaris, 1846). The material in PC (PC0027230, herb. Montagne) represents Nesolechia oxyspora (Tul.) A.Massal. The material in NY (barcode 00920131) was annotated as an isotype by R. Santesson, who determined it to be Abrothallus parmeliarum without further comment. We did not make microscopical sections of the ascomata in the herbarium specimen at NY (barcode 00920131) because only a few remained on the host. The material in RO comprises six pieces of infected lichen thalli mounted directly onto a handwritten sheet, plus one placed in a separate envelope (Figs. 4, 5). However, only the specimen in the envelope and the leftmost of the glued specimens (marked by Santesson with “a”) represent Abrothallus, the other thalli are instead infected by Nesolechia oxyspora. We therefore select the specimen in the envelope as lectotype (Fig. 5), and consider NY barcode 00920131 and the specimen marked with an “a” as isolectotypes. De Notaris (1846) stated that the ascomata of A. buellianus were epruinose, black but turning chestnut brown when wet and larger compared with those of A. bertianus. He also stated that the ascospores were significantly larger than those

Fig. 5. Lectotype of Abrothallus buellianus De Not. from De Notaris’s collection in RO.

of A. bertianus. The De Notaris specimen closely resembles A. parmeliarum in the sizes of the ascomata and ascospores, and the presence of non-amyloid vegetative hyphae. The synonymy of both taxa has been previously proposed (Tulasne, 1852; Rabenhorst, 1870; Saccardo, 1889; Keissler, 1930), but due to the presence of slight differences in the morpho-anatomical features, especially the pigmentation patterns within the apothecia (a darker hypothecium than seen in A. parmeliarum) and the colour reactions of the hymenial substances, we consider that synonymization is premature without a critical mophological study and molecular sequence data. Abrothallus caerulescens I.Kotte in Centralbl. Bakteriol. Parasitenk., 2. Abth. 24: 86. 1909 – Lectotype (designated here, MBT-381111): [GERMANY]. Sachsen, nordöstlich von Wurzen, [on Xanthoparmelia conspersa], Aug, Sep, von Delisch s.n. [Rabenhorst, Lich. Eur. Exs.: No. 550] (BR barcode BR5030004116426!; isolectotypes: H barcode H7028758!, UPS!). Illustration. – Kotte (1909: pl. III, fig. 31), cross-section through infected thallus of X. conspersa (Ehrh. ex Ach.) Hale, showing Abrothallus hyphae reacted with iodine. Description: Vegetative hyphae immersed, K/ILugol+ violet. Ascomata black, epruinose, mostly globose, 0.2–(0.27)– 0.3 mm diam. (n = 11), ca. 0.15 mm tall. Epihymenial layer of dark brown granules, K+ greenish, N+ violet. Upper part of the hymenium greenish, downwards hyaline, ca. 55 µm, K+ greenish, N+ violet. Hypothecium dark brown, turns gradually ligther downwards the base, hypothecial cells rectangular 6.5–11 × 6.5–8 µm, l/w ratio = 1–1.4 (n = 10). Asci bitunicate, clavate, 40–48 × ca. 16 µm, 8-spored. Ascospores 2-celled, asymmetric, brown to dark brown, 14.5–(16.16 ± 1.37)–19 × 5.5–(6.75 ± 0.84)–8.5 µm (upper cell) × 5–(5.93 ± 0.63)–6.5 µm (lower cell), l/w ratio = 2–2.9 (n = 20). Interascal filaments unequally dichotomosly branched, ca. 1.5–2.5 μm wide, tip sometimes slightly widened. Asexual morph present, pycnidial, immersed to semi-immersed, roundish to irregular in shape, Vouauxiomyces-type, black, ostiolate, globose, 0.13– (0.16)–0.2 mm diam. (n = 10). Pycnidial wall dark brown to olivaceous green, greenish parts K+ intensified, brownish parts K–, N–, textura angularis; wall widened at the ostiole, to 45 µm, lateral and lower parts, ca. 10 µm in width. Conidiophores absent. Conidiogenous cells lining the cavity of pycnidium, holoblastic, hyaline, percurrently proliferating, with 1–2(–3) annellations, 8–(11.2)–13 × 2.5–(3.2)–4 µm (n = 10). Conidia hyaline, simple, obpyriform to clavate with truncated end, in muscilage, 9.5–(12.43 ± 1.83)–14.5 × 3–(4.08 ± 0.89)–6.5 µm, l/w ratio = 2.2–4.8 (n = 20). Notes. – Abrothallus tulasnei M.S.Cole & D.Hawkws., another species described from Xanthoparmelia (Vain.) Hale species in the U.S.A., differs mainly in the non-amyloid vegetative hyphae and olive-green reaction of hymenium with potassium hydroxide (Cole & Hawksworth, 2001). Although synonymisation was proposed by Diederich (2003), we treat them as distinct pending a detailed comparative morphological study and molecular sequence data.

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Abrothallus cetrariae I.Kotte in Centralbl. Bakteriol. Parasitenk., 2. Abth. 24: 82. 1909 – Lectotype (designated here, MBT381112): GERMANY. [on Platismatia glauca], 1854, 1856, Bail s.n. [Rabenhorst, Lich. Eur. Exs.: No. 90 sub A. smithii Tul.] (H barcode H7028760!; isolectotypes: O!, UPS!). Illustration. – Kotte (1909: 14, fig. 1; pl. III, figs. 26–27). Description. – Vegetative hyphae K/I Lugol –. Ascomata black, globose, epruinose, 0.17–(0.28)–0.45 mm diam. (n = 8), ca. 0.25 mm high. Epihymenial layer of brown granules, K+ olive-green, N–. Hymenium hyaline, K–, N–, ca. 70 µm. Exciple ca. 70 µm. Hypothecium light brown, ca. 170 µm tall, hypothecial cells rectangular, 5–8 × 4 –7 µm, l/w ratio = 1–1.3 (n = 10). Asci bitunicate, clavate, 8-spored, 47–61 × 10–12 µm. Ascospores 2-celled, asymmetric, brown to dark brown, 13– (14.6 ± 1.27)–17 × 5 –(6 ± 0.52)–7.5 µm (upper cell) × 4.5–(5 ± 0.28)–5.5 µm (lower cell), l/w ratio = 2.15–2.8 (n = 20). Asexual morph not observed. Notes. – In introducing this species name, Kotte (1909) cited A. smithii Tul. (Tulasne, 1852) as a synonym. That name is, however, superfluous and illegitimate as the protologue included the earlier name A. bertianus as a synonym which consequently should have been adopted, and so Tulasne’s name must be automatically typified by the type of the earliest legitimate name he cited as a synonym, Endocarpon parasiticum Ach. (see under A. parmeliarum below) (ICN Art. 52.2). We speculated whether that meant Kotte’s name should also be regarded as illegitimate, but concluded that not to be the case and that ICN Art. 52.1 did not apply as Tulasne’s name was illegitimate and so not an “epithet [that] ought to have been adopted”. The Rabenhorst exsiccate (Rabenhorst, Lich. Eur. Exs.: No. 90) which was also cited by Kotte (1909) is designated as lectotype for A. cetrariae above.

The hyphae of the host species, Platismatia glauca (L.) W.L.Culb. & C.F.Culb. turn light blue with iodine solutions (Culberson & Culberson, 1968). This reaction can be easily misinterpreted as the reaction of the Abrothallus. In this case, however, the hyphae of A. peyritschii (Stein) I.Kotte and A. bertianus turn violet in response to treatment with Lugol’s solution, while those of A. cetrariae are unchanged. The sizes of the ascospores in Rabenhorst, Lich. Eur. Exs.: No. 90 deviate slightly from those stated by Kotte (1909): 12.5–13.8 × 3.9– 6.2 µm versus 13–17 × 5 –7 µm measured by us. Our measurements are closer to those of A. buellianus and A. parmeliarum as reported elsewhere in the present paper. Despite this similarity, we refrain from treating the species as synonyms pending molecular sequence data and morpho-anatomical study. Kotte (1909) mentioned the presence of pycnidia with coni dia 2.08–3.12 µm in length, but we did not find any pycnidia in the specimens we examined. The conidia of Abrothallus santessonii, a species which occurs on the same host lichen, are much longer (see below). Instead, we found that Platismatia W.L.Culb. & C.F.Culb. specimens with A. cetrariae were additionally infected by Nesolechia oxyspora both forming ascomata on the irregular galls on the lichen thallus. Abrothallus parmeliarum (Sommerf.) Arnold in Flora 57: 102. 1874 ≡ Lecidea parmeliarum Sommerf., Suppl. Fl. Lapp.: 176. 1826 (“parmeliorum”), nom. cons. prop. – Lectotype (designated here, MBT-381221): NORWAY. Nordland, on Parmelia saxatilis, May 1823, Sommerfelt s.n. (O No. L-171771!, left uppermost thallus). = Endocarpon parasiticus Ach., Syn. Meth. Lich.: 100. 1814, nom. rej. prop. ≡ Abrothallus parasiticus (Ach.) Sacc. in Michelia 2: 332. 1881 (“(DeNtrs.) Nyl.”) – Lectotype

Fig. 6. A, Herbarium sheet from S.C. Sommerfelt’s collection with lectotype specimen of A. parmeliarum (the uppermost left thallus; O-L-171771); B, Thallus of Parmelia saxatilis with ascomata of A. parmeliarum, lectotype. The arrow indicates the region with abundance of ascomata.

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(designated here, MBT-383467): [illustration in] Smith & Sowerby, Engl. Bot.: t. 1866. 1808. = Lichen parasiticus Sm. in Smith & Sowerby, Engl. Bot.: t. 1866. 1808, nom. illeg., non Lichen parasiticus Hoffm., Enum. Lich.: 39. 1784 – Holotype: UNITED KINGDOM. Wales, “first sent some years ago by Mr. Griffith from N. Wales & from Cribbath Hill near Ystrad Gynlais, S. Wales” [on Parmelia omphalodes] (BM-Sm 26.1866). = Abrothallus smithii Tul. in Ann. Sci. Nat., Bot. 17: 113. 1852, nom. illeg. See Keissler (1930) for further isonyms and possible additional later synonyms. For images of the lectotype of Abrothallus parmeliarum, see Fig. 6A, B). Illustrations (selected). – Smith & Sowerby (1808: t. 1866); Hertel (1971: 229), ascospores of Sommerfelt’s material deposited in UPS. Description. – Vegetative hyphae immersed, K/ILugol –. Ascomata plane to globose, up to 0.4 mm diam., ca. 0.25 mm tall, slightly green pruinose (the material in UPS mostly epruinose, a few younger ascomata slightly green pruinose); some ascomata on black necrotic spots. Epihymenial layer of brown granules. Hymenium colourless to slightly greenish in upper part, K+ green, N+ violet, ca. 80–90 µm. Exciple 50–80 µm. Hypothecium light brown to brown, ca. 150 µm high, hypothecial cells rectangular, 6.5–9.5 × 6.5–9.5 µm, l/w ratio = 1–1.5 (n = 10). Asci bitunicate, subclavate to clavate, 8-spored, 47–64 × 8–17 µm. Interascal elements irregularly branched, ca. 1.5 µm wide, tip ca. 2.5 µm. Ascospores 2-celled, asymmetric, verruculose, brown, 13–(15.4 ± 1.14)–16.5 × 5 –(6.1 ± 0.58)–7 µm (upper cell) × 4 –(5.35 ± 0.65)–6.5 µm (lower cell), l/w ratio = 2–3.1 (n = 20) [14.5–(16.33 ± 0.88)–17.5 × 5.5–(6.53 ± 0.55)–7.5 × 5–(5.73 ± 0.55)–6.5, l/w ratio = 2.13–3.18 (n = 20) in specimen in UPS]. Asexual morph not observed. Notes. – Alstrup & Hawksworth (1990) and Hawksworth (2003) pointed out that the correct name for this species could be Abrothallus parasiticus, based on the name Endocarpon para siticum Ach. (Acharius, 1814: 100). As Acharius only referred to the published illustration of Lichen parasiticus Sm., and did not see any specimen, the illustration he cited is selected as lectotype above. The name Lichen parasiticus itself, however, is typified by fragmentary material protected in celophane and glued (unfortunately with the lower surface of the lichen upwards) to the original drawing of the published plate in BM. Although four localities were cited in the original account, and two from Wales are handwritten on the sheet, it is unclear which of these the fragment came from. This single fragment was accepted as the holotype by Laundon (2005) as that was the single element evidently used in preparing the drawings, even though Smith & Sowerby were aware of reports from other sites. Sommerfelt (1826: 176) listed the name Endocarpon para siticum as a synonym of Lecidea parmeliarum, and further referred to the English Botany illustration. However, the name proposed by him was not superfluous when published as there was an earlier homonym in existence, Lecidea parasitica Flörke, so the epithet “parasitica” was pre-occupied in Lecidea. When the species is placed in Abrothallus, however, the epithet

“parasiticus” again becomes available and should be taken up under the ICN. Laundon (2005) argued that conservation was unnecessary because of the existence of the combination A. parasiticus which he considered was based on a different type; Saccardo (1881: 332) had used the authority “(DNtrs.) Nyl.” which Laundon took to mean that the epithet “parasiticus” was preoccupied in Abrothallus as based on A. bertianus De Not. and so was a synonym of that species and not of A. parmeliarum. This is questionable, and it seems more probable that Saccardo had realized that the earlier epithet “parasiticus” was available for the broad concept of the species, especially as the only host he mentioned was the species now known as Flavoparmelia caperata (L.) Hale and not any brown parmelioid lichen. As such the author citation should be corrected to “(Ach.) Sacc.” in accordance with Art. 41.4, as it appears in Index Fungorum. In order to retain the well-known epithet “parmeliarum” and maintain nomenclatural stability, we have submitted a formal proposal to conserve Lecidea parmeliarum against Endocarpon parasiticum in order to maintain use of the binomial Abrothallus parmeliarum (Hawksworth & al., 2018). Material of Søren Christian Sommerfelt is preserved in UPS and O, and we considered it fitting to select one of his collections as the conserved type. The collection in UPS was labelled by Sommerfelt as “Lecidea parmeliorum Somf.” but without date and locality. That specimen was studied and illustrated by Hertel (1971), who incorrectly regarded it as a syntype, overlooking the fact that the name was superfluous and automatically typified. There is material dated 1823 in O which is a mixture of two host species, P. saxatilis (L.) Ach. and P. omphalodes (L.) Ach., glued onto the same sheet and collected from two different localities in Nordland, Norway, on different dates. We decided to designate the uppermost left thallus of Parmelia saxatilis as lectotype of L. parmeliarum (Fig. 6A, B) as permitted by Art. 7.5(b). The names A. parmeliarum and A. buellianus were synonymized with A. smithii by Tulasne (1852), however, as the synonymized names are older, the latter name is superfluous and therefore illegitimate (Laundon, 2005). The ascospores dimensions reported by Tulasne (1852) for A. smithii are slightly shorter (12.8–16 × 6.5 µm), but still within the range of A. parmeliarum and A. buellianus, as well as for A. cetrariae. Tulasne (1852) noted the rare occurrence of semi-immersed pycnidia in A. smithii with conidia ca. 6.5 × 5 –6.5 µm; Kotte (1909) reported conidia for A. parmeliarum with dimensions of 7.8–9.7 × 4.9–6.2 µm. Neither the original material of A. buellianus nor of A. parmeliarum are accompanied by the asexual morph. Diederich (2011) hypothesized that the species of Nesolechia A.Massal., a lichenicolous genus of the Parmeliaceae in the Punctelia clade (Divakar & al., 2015), could be the host of Abrothallus parmeliarum, facilitating the presence of A. parmeliarum in other hosts different than Parmelia s.str. Although Sommerfelt (1826) pointed out that the thalli of the hosts P. saxatilis and P. omphalodes were deformed by the presence of the parasite, we have not observed Nesolechia in any of Sommerfelt’s specimens neither macroscopically (presence of ascomata) nor microscopically (presence of ILugol+ hyphae). More research is needed to determine if there is any biological

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relationship beyond the co-occurrence of both taxa. In our experience it is not uncommon for several lichenicolous fungi to occur on a single host thallus. Additional specimens examined. – NORWAY. Nordland, Saltdalen, in Parmelia saxatili et omphalodi in saxis, May 1823, Sommerfelt s.n. & Bodø, in Parmelia saxosum, Sep 1823, Sommerfelt s.n. (O No. L-171771); Hordaland, Østersjøen, P. saxatilis, Sep 1827, Sommerfelt s.n. (O No. L-17180); sine loco, on P. saxatilis, Sommerfelt s.n. (UPS No. L-127813). Abrothallus peyritschii (Stein) I.Kotte in Centralbl. Bakteriol. Parasitenk., 2. Abth. 24: 76. 1909 ≡ Abrothallus parmeliarum var. peyritschii Stein in Cohn, Krypt.-Fl. Schlesien 2(2): 211. 1879 – Lectotype (designated here, MBT-381113): [ITALY]. An Larixrinde am Hühnerspiel bei 1400 m oberhalb Gossensass [= Colle Isarco] in Tirol, [on Vulpicida pinastri], Oct 1878, Stein s.n. [Arnold, Lich. Exs.: No. 780, as Abrothallus parmeliarum var. peyritschii] (H barcode H7028747!; isolectotype: K!). Illustrations: Kotte (1909: pl. 1, figs. 1–24). Description. – Vegetative hyphae immersed, K/ILugol+ violet. Ascomata black, epruinose, globose and somewhat constricted at the base to slightly flattened, 0.2–(0.29)–0.38 mm diam. (n = 11); a few ascomata surrounded by a black necrotic rim. Epihymenial layer dark brown to violet-brown, K+ greenish, N–. Hymenium hyaline, upper part K+ greenish, N–, ca. 65 µm. Hypothecium dark brown to violet-brown, ca. 80 µm tall; hypothecial cells rectangular 5–8 × 4 –7 µm, l/w ratio = 1–1.4 (n = 10). Exciple ca. 80 µm. Asci bitunicate, clavate, 8-spored, 43–50 × 12–16 µm. Interascal filaments irregularly branched, ca. 1–1.5 µm, tip not or only slightly widened. Ascospores 2-celled, asymmetric, verruculose, brown, 10–(11.8 ± 0.8)–13 × 4.5–(5.3 ± 0.4)–6 µm (upper cell) × 4 –(4.3 ± 0.35)–5 µm (lower cell), l/w ratio = 1.6–2.8 (n = 20). Asexual morph not observed. Notes. – Abrothallus peyritschii was first mentioned by Stein (1879) who treated it as a variety of A. parmeliarum differing from the type variety in the ascomatal pigments (violet blackish and brown in A. peyritschii instead of almost colourless to light brown in A. parmeliarum) and different host (Vulpicida pinastri (Scop.) J.-E.Mattsson & M.J.Lai). Kotte (1909) raised it to the rank of species but pointed out that the dimensions of the ascospores of A. peyritschii correspond almost exactly to those of A. parmeliarum (10.4–13 × 4.7–6 µm and 12.5–14.3 × 4.5–6 µm, respectively). Although these measurements alone show distinction between these two species, the difference is more pronounced in our own measurements, viz. 10–13 × 4.5–6 µm in A. peyritschii and 13–16.5 × 5 –7 µm in A. parmeliarum. Kotte (1909) reported the presence of an asexual morph of A. peyritschii, characterized by the presence of pear-shaped conidia 5.2–6.76 × 3.9–5.2 µm. The presence of pycnidia with conidia roughly of this size has been mentioned in several papers (e.g., Diederich, 2004; Suija, 2006). We did not find an asexual morph on any specimen from the Arnold exsiccate collection (Arnold, Lich. Exs.: No. 780). An example from this exsiccate was selected for lectotypification because it was referred to by both Stein (1879) and Kotte (1909). 1176

Abrothallus santessonii (D.Hawksw.) Suija, D.Hawksw. & Pérez-Ort., comb. nov. (MycoBank no. MB824720111) ≡ Vouauxiomyces santessonii D.Hawksw. in Bull. Brit. Mus. (Nat. Hist.), Bot. 9: 69. 1981 – Holotype: SWEDEN. Uppland, Alsike s:n, Lunsen, vid åkertegen SV om Hjortronmossen, på grenar på marken i granskog, in thallo Platismatiae glaucae, 23 Sep 1947, Santesson s.n. (UPS No. F-000228!; isotypes: UPS No. F-015653!, K No. K(M)-24331!). Illustration: Hawksworth (1981: 68), conidia, conidiogenous cells. Description. – Vegetative hyphae immersed, K/ILugol –. Ascomata black, heavily green pruinose (Fig. 7A, B), flattened with wide base, then almost globose, constricted at the base to slightly stipitate, 0.35–(0.4)–0.5 mm diam. (n = 19), ca. 0.1– 0.15 mm tall, some arising on black necrotic spots on the lichen thallus (black rim surrounding the ascomata). Epihymenial layer of brown granules (Fig. 7E), K+ greenish, N+ violet. Hymenium hyaline, upper part slightly greenish (Fig. 7C, E) ca. 60–70 µm. Exciple (Fig. 7C) ca. 50–60 µm. Hypothecium light brown to brown (Fig. 7C, D), becomes lighter towards the base, ca. 110–140 µm; hypothecial cells rectangular (Fig. 7D), 6.5–9 × 4.5–7.5 µm, l/w = 1–2.1 (n = 10). Asci clavate, 8-spored, 34.5–46 × 10.5–15 µm. Ascospores 2-celled, light brown (when young) to brown (mature), asymmetric, verruculose (Fig. 7F), 12.5–(13.6 ± 0.9)–15 × 3.5–(5.1 ±0.7)–6 µm (upper cell) × 3–(4 ± 0.6)–5 µm (lower cell), l/w ratio = 2.2–3.8 (n = 20). Interascal filaments irregularly branched (Fig. 7F), ca. 2(–3) µm, tip slightly widened, ca. 3 µm. Asexual morph common, black, pycnidial, ostiolate, roundish; immersed to semi-immersed, ca. 0.15–(0.2)–0.35 mm diam. (n = 20), usually surrounded by a black necrotic rim formed on the lichen thallus, globose to pyriform in shape (Fig. 7A, B, G, H); the pycnidial wall dark brown, sometimes greenish near pycnidial cavity, textura angularis, K+ greenish, N–; the wall ca. 15 µm, but widened at the ostiolum, ca. 40 µm. Conidiophores absent. Conidiogenous cells holoblastic, percurrently proliferating, 1–2 annellations seen (to 4 fide Hawksworth, 1981), hyaline, lining the pycnidial cavity, 5.9–(11.2)–15.6 × 2.5–(3.2)–4.9 µm (8–12 × 3–4 µm fide Hawksworth, 1981). Conidia hyaline, obpyriform (Fig. 7I), in mucilage, 5–(7.4 ± 1.0)–9 × 3 –(4.7 ± 0.8)–6 µm ((7–)7.5–10.5 (–11.5) × (5–)5.5–7(–7.5) µm fide Hawksworth, 1981), l/w ratio = 1.5–2.1 (n = 20). Notes. – With the abolition of dual nomenclature, the generic name Vouauxiomyces has to be synonymized with Abrothallus (Rossmann & al., 2016), and a new combination is required for V. santessonii. We examined the holotype and isotype collections of V. santessonii in UPS and K. In all three specimens, we found at least one or two green pruinose ascomata typical for Abrothallus intermixed with abundantly occurring pycnidia. These were not described by Hawksworth (1981) who took them to be A. parmeliarum, a name then being used in a broad sense. We also did not examine these ascomata in detail in order not to destroy the material, and the above description of the sexual morph is based on examination of unmistakably similar material with both morphs on the same lichen thallus. This species is more frequently found as the asexual morph, and ascomata are rare or dispersed amongst the pycnidia. Based

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on our observations, A. santessonii is known with certainity from northern Europe, the United States and Canada; the only known host is Platismatia glauca. Despite the similarity in host, we do not synonymize A. santessonii with A. cetrariae, because the size of the conidia given by Kotte (1909) for A. cetrariae (4.9–6.5 × 3.9–4.9 µm) clearly deviates from that given for A. santessonii (7.5–10.5 ×

5.5–7 µm) by Hawksworth (1981) and measured by us (5–9 × 3–6 µm). Also, the sexual morph of the type collection of A. cetrariae differs from A. santessonii in many aspects, notably in the sizes of the ascomata and ascospores, and pruinosity of the ascomata. We have not examined the material described as Epicoc cum parmeliarum H.Olivier (= Phoma parmeliarum (H.Olivier)

Fig. 7. Abrothallus santessonii (D.Hawksw.) Suija, D.Hawksw. & Pérez-Ort. Holomorph. A & B, Thallus of Platismatia glauca with ascomata and pycnidia of A. santessonii; C, Cross section of ascomata, detail of exciple, hymenium and hypothecium (in water; TU65898), D, Detail of hypothecium (in water; TU65898); E, Section of hymenium after treatement with KOH (TU65894); F, Ascospores and interascal elements (in KOH, TU45804), G, Pycnidia on thallus of Platismatia glauca (TU65895); H, SEM of the pycnidium with conidia streching out from the opening of the pycnidium; I, Conidia (both TU78950). — Scales: A, B, G: 1 mm; C, E: 50 µm; D, F: 10 µm; H: 100 µm; I: 5 µm. The arrowheads in figures A and B indicate green pruinose ascomata (left side) and conidomata (right side).

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Vouaux) which has been suggested to be a possible earlier name for A. santessonii (Hawksworth, 1981). The measurements given for the conidia of 6–9 × 5 –7 µm (Vouaux, 1914: 196) agree with the concept of A. santessonii, however, that fungus was described from Flavoparmelia caperata and Parmelia sulcata Taylor. The conidia of another species occurring predominantly as the asexual morph and growing on F. caperata, A. microspermus Tul. (= V. truncatus (B.de Lesd.) Dyko & D.Hawksw.), have been reported as shorter and narrower: 6.5 × 3.5 µm (Tulasne, 1852) and 6.5–8.5 × 4–5.5 µm (Hawksworth & Dyko, 1979). The extent to which these small differences may have been due to the use of different mounting media is unclear. Additional selected specimens examined (all on Platismatia glauca). – CANADA. New Brunswick, Albert County, Fundy National Park, Dickson Falls Trail, ca. 85 m (45.58667°N, 63.02778°W), 23 Sep 2013, Harris 58889 (NY barcode 01818015). ESTONIA. Ida-Viru County, Iisaku community, near Pootsiku village, 19 Sep 2007, Tõrra s.n. (TU65894); Põlva County, Vastse-Kuuste comm., Kiidjärve (58.14194°N, 27.03917°E), on twigs of Picea abies, 28 Sep 2007, Marmor & Tõrra s.n. (TU65898); Tartu County, Vara forestry (58.53023°N, 26.95730277°E), on Betula pendula, 2015, Liira s.n. (TU78950); Võru County, Roosa forestry (57.80083°N, 26.84778°E), on log, 12 Aug 2005, Suija s.n. (TU-45804); Võru County, Antsla comm., Karula National Park, Ähijärve village (57.72214°N, 26.54961°E), 11 Oct 2016, Randlane 098Karula (TU84566). FINLAND. Etelä-Häme, Rajakallio (61.25444°N, 25.10778°E), 11 Aug 2007, Palice s.n. (TU65896).

AUTHOR CONTRIBUTIONS AS performed researches in most of the herbaria, except BM; DLH and SPO performed searches of De Notaris publications and archival materials; all authors wrote the manuscript. — ORCID: AS, https:// orcid.org/0000-0003-3784-9414; DLH, https://orcid.org/0000-00029909-0776; SPO, https://orcid.org/0000-0002-5411-3698

ACKNOWLEDGEMENTS The curators of BR, H, M, NY, PC, RO, and UPS are thanked for granting loans. We especially thank Damien Ertz (BR), Lionel Kervan, Bernard Duhem (both from PC), Begoña Aguirre-Hudson and Heidi Döring (K), Einar Timdal (O), Anders Nordin and Stefan Ekman (UPS) for their hospitality during the stay of AS in these herbaria, Anna Millozza (RO) for her kindness for seeking out the herbarium material of G. De Notaris, and Paola Bonfante, Laura Guglielmone, Elena Borgi (Torino) and Graciano García Meyer-Doehner and Félix Alonso Sánchez (RJB, Madrid) for looking for archival materials concerning De Notaris’s publications. We thank Wolfgang von Brackel (Germany) for allowing to use his material, Yolanda Ruíz León (RJB, Madrid) for performing Scanning Electron Microscopy, and Jaan Liira (Tartu) for taking macrophotographs. The curator of RO is also acknowledged for allowing us to publish scanned herbarium images. Reviewers are thanked for their constructive criticism which improved the quality of the manuscript. The work of AS is financed by institutional research

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funding project IUT20-30, SYNTHESYS project ES-TAF-5485 and the European Regional Development Fund (Centre of Excellence EcolChange). SPO is supported by the grant RYC-2014-16784 from the Spanish Ministry of Economy, Industry and Competitiveness.

LITERATURE CITED Acharius, E. 1814. Synopsis methodica lichenum. Lundae [Lund]: litteris et sumtibus Svanborg et Soc. https://books.google.at/books? id=QqI_AAAAcAAJ Alstrup, V. & Hawksworth, D.L. 1990. The lichenicolous fungi of Greenland. Meddel. Grønland, Biosci. 31: 1–90. Brackel, W. von 2009. Weitere Funde von flechtenbewohnenden Pilzen in Bayern – Beitrag zu einer Checkliste IV. Ber. Bayer. Bot. Ges. 79: 5–55. Clauzade, G., Diederich, P. & Roux, C. 1989. Nelikenigintaj fungoj likenlogaj. Ilustrita determinlibro. Bull. Soc. Linn. Provence Num. Special 1: 1–142. Clements, F.E. & Shear, C.L. 1931. The genera of Fungi. New York: Hafner. Cole, M.S. & Hawksworth, D.L. 2001. Lichenicolous fungi, mainly from the USA, including Patriciomyces gen. nov. Mycotaxon 77: 305–338. Culberson, W.L. & Culberson, C.F. 1968. The lichen genera Cetrelia and Platismatia (Parmeliaceae). Contr. U.S. Natl. Herb. 34: 449–558. De Notaris, G. 1846. Frammenti Lichenografici di un lavorno inedito. Giorn. Bot. Ital. 1: 174–224. De Notaris, G. 1849. Abrothallus novum lichenum genus. Mem. Reale Accad. Sci. Torino, ser. 2, 10: 351–355. Diederich, P. 1989. Les lichens epiphytiques et leurs champignons lichenicoles (macrolichens exceptes) du Luxembourg. Travaux scientifiques de Musee national d’Histoire naturelle de Luxembourg. No. 14. Luxembourg: Ministere des Affairs Culturelles. Diederich, P. 2003. New species and new records of American lichenicolous fungi. Herzogia 16: 41–90. Diederich, P. 2004. Abrothallus. Pp. 626–630 in: Nash, T.H., III, Ryan, B.D., Diederich, P., Gries, C.A. & Bungartz, F. (eds.), Lichen flora of the Greater Sonoran Desert Region, vol. 2. Tempe: Lichens Unlimited, Arizona State University. Diederich, P. 2011. Description of Abrothallus parmotrematis sp. nov. (lichenicolous Ascomycota). Bull. Soc. Naturalistes Luxemb. 112: 25–34. Divakar, P.K., Crespo, A., Wedin, M., Leavitt, S.D., Hawksworth, D.L., Myllys, L., McCune, B., Randlane, T., Bjerke, J.W., Ohmura, Y., Schmitt, I., Boluda, C.G., Alors, D., Roca-Valiente, B., Del-Prado, R., Ruibal, C., Buaruang, K., Núñez-Zapata, J., Amo de Paz, G., Rico, V.J., Molina, M.C., Elix, J.A., Esslinger, T.L., Tronstad, I.K.K., Lindgren, H., Ertz, D., Gueidan, C., Saag, L., Mark, K., Singh, G., Dal Grande, F., Parnmen, S., Beck, A., Benatti, M.N., Blanchon, D., Candan, M., Clerc, P., Goward, T., Grube, M., Hodkinson, B.P., Hur, J.-S., Kantvilas, G., Kirika, P.M., Lendemer, J., Mattsson, J.-E., Messuti, M.I., Miadlikowska, J., Nelsen, M., Ohlson, J.I., Pérez-Ortega, S., Saag, A., Sipman, H.J.M., Sohrabi, M., Thell, A., Thor, G., Truong, C., Yahr, R., Upreti, D.K., Cubas, P. & Lumbsch, H.T. 2015. Evolution of complex symbiotic relationships in a morphologically derived family of lichen-forming fungi. New Phytol. 208: 1217–1226. https://doi.org/10.1111/nph.13553 Farr, E.R., Leussink, J.A. & Stafleu, F.A. 1979. Index nominum genericorum (Plantarum), vol. 1. Regnum Vegetabile 100. Utrecht: Bohn, Scheltema & Holkema. Greuter, W., Brummitt, R.K., Farr, E., Kilian, N., Kirk, P.M. & Silva, P.C. 1993. NCU-3: Names in current use for extant plant genera. Regnum Vegetabile 129. Königstein: Koeltz Scientific Books.

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