Introduction
The recent phylogenetic study by Baloch and co-workers (Baloch et al. Reference Baloch, Lücking, Lumbsch and Wedin2010) showed that the type species of Belonia and Pachyphiale are nested within Gyalecta. Based on these results, Baloch et al. (Reference Baloch, Lücking, Lumbsch and Wedin2010) suggested that Belonia and Pachyphiale should be synonymized with Gyalecta, but the authors did not provide formal new combinations. Here we provide the necessary new combinations for currently accepted species of Belonia and Pachyphiale that lack valid names in Gyalecta.
Former Belonia species
Belonia Körb. is a group of Trentepohlia-containing crustose lichens with perithecioid apothecia and needle-shaped, multi-septate spores. In the phylogeny presented by Baloch et al. (Reference Baloch, Lücking, Lumbsch and Wedin2010), the two Belonia species included [the type species of Belonia, B. russula, and B. herculina (as B. herculana)], do not form a monophyletic group, but both were nested within Gyalecta. Based on these findings, we here suggest new combinations for the species currently placed in Belonia.
Gyalecta calcicola (Walt. Watson) Baloch & Lücking comb. nov.
MycoBank No.: MB 803305
Belonia calcicola Walt. Watson, J. Bot. 73: 160 (1935); type: England, North Somerset, Goblin Coombe, June 1934, Watson s. n. (BM—holotypus; BM—isotypus).
Gyalecta herculina (Rehm) Baloch, Lumbsch & Wedin comb. nov.
MycoBank No.: MB 803316
Segestrella herculina Rehm in Lojka, Mathem. és Természet. Közlem. 11: 62 (Reference Lojka1876).—Verrucaria herculina (Rehm) Lojka, Mathem. és Természet. Közlem. 21: 370 (Reference Lojka1885).—Belonia herculina (Rehm) Keissler, Rabenh. Krypog. Fl. 9(1–2): 287 (1938); type: Romania (“Hungary”), “ad corticem fagorum infra sum. montem Domugled prope Thermis Herculis in Banatu,” 20 September 1872, Lojka 1048 (S L2801—lectotypus, designated here).
Belonia herculana Hazsl., Grevillea 6: 109 (1878); type: Romania (“Hungary”), “ad corticem fagorum infra sum. montem Domugled prope Thermis Herculis in Banatu”, 20 September 1872, Lojka 1048 (S L2801—neotypus, designated here).
The nomenclature of this lichen has been rather confused. The earliest name was published as Segestrella herculina Rehm., in Lojka (Reference Lojka1876). The type locality (“on bark of a beech at lower elevation of Mt Domugled”, translated from Hungarian) corresponds to Mt Domogled in present-day Romania, and “herculina” implicitly refers to the Hercules Thermes spa on the lower southern slope of this mountain. Lojka expected a more detailed description by Rehm that never appeared, but it is clear that the original description was by Rehm (Lojka Reference Lojka1885). Verrucaria hungarica Nyl. in litt. was mentioned in the original diagnosis, but never published. Later, Lojka (Reference Lojka1885) gives more detail on the original material, all of which is consistent with the original brief diagnosis. The material was collected by Lojka in 1872 and, according to Lojka (Reference Lojka1885), given the collection number 1048. It was also distributed in the Lojka, Lich. Regn. Hung. Exs. as number 115. There are two samples annotated as types in hb. Rehm (which is housed in S), one annotated Lojka 1048 and one Lojka 1049. We designate the sample Lojka 1048 from hb. Rehm in S as lectotype of Segestrella herculina Rehm.
Belonia herculana Hazsl. (Haszlinsky Reference Haszlinsky1878) was described separately, but it is based on part of the same original material as Segestrella herculina Rehm. Lojka (Reference Lojka1885) comments on the fact that Hazslinsky attributes the type of Belonia herculana to two collectors: “Vicenze Borbás was mentioned as collector, too – it is just because during the excursion in autumn 1872 he was asking for a specimen and I cut a piece of bark for him…” (translated from Hungarian). The sample studied by Hazslinsky should be housed in Budapest (hb. BP), as BP bought Hazslinsky's herbarium after his death, but no material could be located there. A number of Belonia specimens were, however, out on loan and destroyed during World War II, according to a shipping list from 1944 preserved at BP (E. Farkas & L. Lökös, in litt.). We assume that the type of Belonia herculana was among the samples destroyed. As it is clear that Hazslinsky based his description on one of the many pieces collected by Lojka during his excursion to Mt Domogled (all probably to be considered syntypes of Segestrella herculina), we neotypify Belonia herculana Hazsl. on the same sample in S from hb. Rehm as the lectotype of Segestrella herculina Rehm., to automatically make the names synonyms and Belonia herculana Hazsl. superfluous.
Gyalecta incarnata (Th. Fr. & Graewe) Baloch & Lücking comb. nov.
MycoBank No.: MB 803306
Belonia incarnata Th. Fr. & Graewe, in Th. Fr., Öfvers. K. Svensk. Vetensk.-Akad. Förhandl. 21: 274 (1865).—Gongylia incarnata (Th. Fr. & Graewe) Zahlbr., in Engler & Prantl, Nat. Pflanzenfam. 1: 57 (1903); type: Sweden, Västergötland, Baljefors, near Främmestad, 1863, Graewe s. n. (UPS—holotypus).
Belonia terrigena Eitner, Jahresber. Schlesisch. Ges. Vaterldnd. Kultur 99: 34 (1910, ‘1911’); type: Poland, Schlesien, Riesengebirge, “um die Veilchensteine auf alten Rasenausstichen neben dem Kammweg”, June 1906, Eitner s. n. (W—holotypus).
This species is commonly cited as “ex” rather than “in”, but it is clear from the text in Fries' paper (Fries Reference Fries1865) that the intention of Fries and Græwe was to describe this species together, based on material collected by Græwe.
Gyalecta lumbrispora (Etayo) Baloch & Lücking comb. nov.
MycoBank No.: MB 803307
Belonia lumbrispora Etayo, Öst. Z. Pilzk. 5: 151 (1996); type: Spain, Canary Islands, Gomera, La Meseta, barranco de la Cueva Encantada, on Ocotea foetens, 720 m alt., 3 August 1994, Etayo 58(3) (MA-Lich.—holotypus; BM, W and hb. Etayo—isotypi).
Gyalecta lyngei Baloch & Lücking nom. nov.
MycoBank No.: MB 803517
Belonia arctica Lynge, Lich. Nov. Zemlya: 39 (1928); type: Novaya Zemlya, Matochkin Shar Distr., Chalhonik Valley, 13 July 1921, Lynge s. n. (O—holotypus); non Gyalecta arctica Malme, Ark. Bot. 25A(2): 7 (1932).
This species is known only from the type collection. The taxonomic status and its distinction from G. russula, which has somewhat larger ascospores, require additional studies.
Gyalecta mediterranea (Nav.-Ros. & Llimona) Baloch & Lücking comb. nov.
MycoBank No.: MB 803308
Belonia mediterranea Nav.-Ros. & Llimona, Lichenologist 29: 16 (1997); type: Spain, Catalonia, prov. Girona, Medes Islands, 29 August 1981, Llimona s. n. (BCC Lich. 9309— holotypus).
Gyalecta nidarosiensis (Kindt) Baloch & Lücking comb. nov.
MycoBank No.: MB 803309
Microglaena nidarosiensis Kindt, Kgl. norske vidensk. Selsk. Skr. 1884: 4 (1885).—Belonia nidarosiensis (Kindt) P. M. Jørg. & Vězda in Jørgensen, Vězda & Botnen, Lichenologist 15: 54 (1983); type: Norway, 14 August 1884, Kindt s. n. (BG—holotypus; TRH, UPS—isotypi).
Clathroporina calcarea Walt. Watson, J. Bot., Lond. 63: 131 (1925); type: England, V.C.33, Gloucestershire, Winchcombe, 17 May 1924, Knight s. n. (BM—lectotypus).
Clathroporina caudata Vězda & Vivant, Bull. Soc. Bot. Fr., Let. bot. 118: 288 (1972) [‘1971’]; type: France, Gallia, Montes Pyrenaei Occident., St.-Just-Ibarre, 17 VII 1970, Vivant s. n. (Vězda: Lich. sel. exs. no. 1051; BCC Lich. 368, G—isotypi).
Gyalecta pellucida (Coppins & Malcolm) Baloch & Lücking comb. nov.
MycoBank No.: MB 803310
Belonia pellucida Coppins & Malcolm, Lichenologist 30: 563 (1998); type: New Zealand, North Island, Wellington, Scorching Bay, 10 October 1995, Malcolm 2490 (CHR—holotypus; E— isotypus).
Gyalecta russula (Körb. ex Nyl.) Baloch, Lumbsch & Wedin comb. nov.
MycoBank No.: MB 803311
Belonia russula Körb. ex Nyl., Act. Soc. Linn. Bordeaux 21: 346 (1857); type: Poland, “ad rupes basalticas faucis ‘Klein Schneegrube' Sudetorum”, Körber s. n. (L. Körber Typenherbar—lectotypus, designated by Navarro-Rosinés & Llimona Reference Navarro-Rosinés and Llimona1997: 25, as “holotype”; and L Körber Stammherbar—isolectotypus; Körber, Lich. Sel. Germ. no. 79—isolectotypi).
Belonia fennica Vain., Meddeland. Soc. Fauna Fl. Fenn. 10: 196 (1883); type: Russia (formerly part of Finland), Kuusamo, 30 July 1877, Vainio s. n. (TUR-31141, 31142—syntypi).
Beloniella cinerea Norman, Kongl. Vetensk.-Akad. Förhandl. 8: 35 (1884); type: Norway, Holmestrand, Nyveien, Norman s. n. (H and S L2798—isolectotypi).
The citation of this species has been confusing in the literature. The name was first published in Körber's exsiccate Lichenes Selecti Germaniae no. 79 (Körber Reference Körber1856), but without a valid description and diagnosis. Nylander (Reference Nylander1857) then provided a description and has to be considered the validating author. Yet, both Körber (Reference Körber1863) and Garovaglio (Reference Garovaglio1873) subsequently intended to publish the species validly, Garovaglio (Reference Garovaglio1873) apparently having overlooked Körber's (Reference Körber1863) treatment in his Parerga. In the latter, Körber (Reference Körber1863: 322) dismissed Nylander's (Reference Nylander1857: 322) view and description of the species, stating that “Nylander konnte keinen schlagenderen Beweis für die bodenlose Oberflächlichkeit seiner Untersuchungen geben, als diesen Nonsens!” [Nylander could not give a clearer proof of the bottomless superficiality of his research than this nonsense, translated from German]. Nylander (Reference Nylander1857) had stated that the species lacks paraphyses and that because of its widened epithecium was related to Gyalecta. So while both Körber (Reference Körber1863) and Nylander (Reference Nylander1857) were right and wrong with regard to particular details of the species, Nylander's (Reference Nylander1857) first published description, even if rudimentary, must be considered a validation of the name Belonia russula, thus taking the credit from Körber (Reference Körber1856, Reference Körber1863) who had collected and thoroughly studied the taxon. Adding to the confusion is that Rabenhorst (Reference Rabenhorst1867) published an edited version of a work by Garovaglio, who had offered validation of both the genus and the species name, with Körber as sole author, but Garovaglio's original work was not published until six years later (Garovaglio Reference Garovaglio1873).
Gyalecta uncinata (P. M. McCarthy & Kantvilas) Baloch & Lücking comb. nov.
MycoBank No.: MB 803312
Belonia uncinata P. M. McCarthy & Kantvilas, Lichenologist 29: 489 (1997); type: Australia, Tasmania, Gordon Road, 8 April 1997, Kantvilas 96/97 (HO 320762—holotypus).
Gyalecta vezdana (Malcolm & Coppins) Baloch & Lücking comb. nov.
MycoBank No.: MB 803313
Belonia vezdana Malcolm & Coppins, Australas. Lichenol. 41: 30 (1997); type: New Zealand, South Island, Sharland Creek, Hira Forest, 60 m alt., 6 February 1997, W. Malcolm 2074 (CHR 489027—holotypus; E—isotypus).
Former Pachyphiale species
Pachyphiale Lönnr. has traditionally been used for Gyalecta-like lichens with multi-spored asci (Vezda Reference Vězda1958, Reference Vězda1969). Baloch et al. (Reference Baloch, Lücking, Lumbsch and Wedin2010) included the type species, Pachyphiale fagicola, in their phylogeny, which was nested in Gyalecta, suggesting that ascospore number cannot be used as a genus-level character. For two Pachyphiale species to be treated in Gyalecta, valid names in Gyalecta are already available: Gyalecta carneola (Ach.) Hellb. [syn: P. carneola (Ach.) Arnold; P. cornea (With.) Poetsch] and Gyalecta fagicola (Hepp ex Arnold) Kremp. [syn.: P. fagicola (Hepp ex Arnold) Zwackh.; P. corticola Lönnr.]. Ten further species have been treated in Pachyphiale. Of these, P. cornea (Tuck.) Poetsch and P. corticola Lönnr. are synonyms of Gyalecta carneola and G. fagicola, respectively, whereas P. carneolutea (Turner) Samp. and P. geoicoides (Vain.) Vězda [syn.: G. geoicoides Vain.] have been transferred to Cryptolechia (Kalb Reference Kalb2007). Pachyphiale arbuti (Bagl.) Arnold, P. gyalizella (Nyl.) S. Ekman, P. himalayensis Vězda & Poelt, and P. ophiospora Lettau, appear to be good species (Vězda Reference Vězda1958; Poelt Reference Poelt1969; Vězda & Poelt Reference Vězda and Poelt1975; Clauzade & Roux Reference Clauzade and Roux1985; Ekman Reference Ekman1996) and are recombined in Gyalecta below. The status of P. lojkana (Nyl.) Keissl. is unclear; Vězda (Reference Vězda1958) included it in Pachyphiale following Keissler (Reference Keissler1933), although the black ascomata and cyanobacterial photobiont do not agree with typical species of Pachyphiale or Gyalecta; Vězda (Reference Vězda1968), Poelt (Reference Poelt1969) and Clauzade & Roux (Reference Clauzade and Roux1985) retained the species in Thelopsis. We believe that this taxon requires a separate phylogenetic study to clarify its relationships. Pachyphiale lecanorina J. Steiner, described from Portugal (Steiner Reference Steiner1918), has never again been mentioned in the literature (Vězda Reference Vězda1958; Poelt Reference Poelt1969; Clauzade & Roux Reference Clauzade and Roux1985) and the name does not appear in any checklist; its taxonomic status is unknown.
Gyalecta arbuti (Bagl.) Baloch & Lücking comb. nov.
MycoBank No.: MB 804287
Bacidiopsis arbuti Bagl., Comm. Soc. Crittog. Ital. 1: 22 (1861).—Pachyphiale arbuti (Bagl.) Arnold, Flora 54: 50 (1871); type: Italy, Baglietto s. n. (TO—holotypus).
Gyalecta gyalizella (Nyl.) Baloch & Lücking comb. nov.
MycoBank No.: MB 803314
Lecidea gyalizella Nyl., Sert. Lich. Trop. Labuan Singapore: 38 (1891).—Pachyphiale gyalizella (Nyl.) S. Ekman, Op. Bot. 127: 130 (Reference Ekman1996); type: USA, Massachusetts, New Bredford, Willey s. n. (H-NYL 21313—lectotypus, selected by Ekman Reference Ekman1996).
Gyalecta himalayensis (Vězda & Poelt) Baloch & Lücking comb. nov.
MycoBank No.: MB 803315
Pachyphiale himalayensis Vězda & Poelt, Khumbu Himal 6: 130 (1974); type: Nepal, Himalaya, Khumbu, “Abies-Rhododendron Wald südlich Kunde”, 3900–4000 m, October 1962, Poelt 1554 (M—holotypus).
Gyalecta ophiospora (Lettau) Baloch & Lücking comb. nov.
MycoBank No.: MB 804288
Pachyphiale ophiospora Lettau, Feddes Repert., Beih. 69: 222 (1937); type: Switzerland, Lettau s. n. (M—holotypus).
This work was funded by The Swedish Taxonomy Initiative (Svenska Artprojektet administered by ArtDatabanken). Additional funding was received from the Swedish Research Council grant VR 621-2009-537, to MW. Kind assistance with translation of Hungarian texts was given by Eva Csesznegi, László Lökös, and Edit Farkas. Edit and Lászlo also very helpfully commented on the details surrounding Lojkás collection trip to Mt Domogled, on the fate of the Haszlinsky's Belonia specimens, and searched for original material in BP. We finally want to thank Harrie Sipman and Fred Barrie for helpful discussions on nomenclatural matters. Any error, however, is entirely our own.
