GYROMITRA FR. SENSU LATO (DISCINACEAE, PEZIZALES) IN MONTENEGRO

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1 GYROMITRA FR. SENSU LATO (DISCINACEAE, PEZIZALES) IN MONTENEGRO Branislav Perić * & Olgica Perić ** * University of Montenegro, Biotechnical Faculty, Mycological center Podgorica, Mihaila Lalića 1, Montenegro branislav@mycolmonten.org **81000 Podgorica, Đulje Jovanova 16 Abstract 11 taxa of the genus Gyromytra Fr. sensu lato recorded in Montenegro are presented. These are G. esculenta (Pers.) Fr., G. gigas (Krombh.) Cooke, G. geogenia (Rahm) Harmaja, G. infula (Schaeff.) Quél., G. martinii Donadini & Astier, G. megalospora Donadini & Riousset), G. neuwirthi Velen., (= G. esculenta var. fragilis Marchand ex Réaudin, 2008), Gyromitra fastigiata (Krombh.) Rehm, G. parma (J. Breitenb. & Maas Geest.) Kotl. & Pouzar, G. perlata (Fr.) Harmaja and G. tasmanica Berk. & Cooke. The southernmost point of G. geogenia range, for now, was found in this region. G. neuwirthii, G. fastigiata and G. tasmanica are rare entities, new for Montenegro. The latter occures in Southern Hemisphere, previously was reported only in Spain in Northern Hemisphere. Key words: Discinaceae, Gyromitra, gigas, geogenia, infula, martinii, megalospora, neuwirthi, fastigiata, parma, tasmanica, Montenegro Introduction T he genus Gyromitra Fr. s.l. (Pezizales), at the present time, increasingly come to our attention. Probable reasons for that are, not only the attractiveness of these specific discomycetes and ambience in which they develop, but also a complete nice atmosphere during early spring trips. Most species mainly inhabit high mountain areas and should be sought in the time of melting snow or shortly thereafter. Otherwise, working with them requires great patience due to delayed maturation of ascomata. After collection it is necessary to be kept in the refrigerator, maintaining an adequate level of humidity, pursuant to natural conditions, in order to complete maturation.

2 Branislav Perić & Olgica Perić Material & Method The material was collected in the Mediterranean, sub-mediterranean and continental region of Montenegro. Investigation was conducted on fresh material while verification was done, on several occasions, on herbarium specimens. Macroscopic descriptions of species and photo illustrations were done in situ with Fg20 reflex Nikon camera and digital cameras Nikon 100 and C5D. Microscopic analysis was performed using the optical microscope (Leica DMLS). Microscopic observations were made on sections, prepared by hand with razors then mounted in the water, Congo Red, Melzer s reagent (to monitor amyloid reaction of ascus), Cotton blue and lacto phenol (to monitor ascospore ornamentation). Micro illustrations were made with the digital camera (DC 300). Drawings of microelements were done using pen and ink and as well as micro illustrations were processed with Adobe Photoshop CS4. 30 ascospores were measured from different apothecia. The material is stored in the herbarium of Mycological Centre, Biotechnical Faculty in Podgorica. Systematic position 120 The family Discinaceae Benedix emend. N.S. Weber, Trappe & O Donnell is characterized by the following features: Macroscopic features epigeous or hypogeous ascomata generally of large stature, sessile or stipitate, cupulate, cerebriformis or more or less correctly or incorrectly tuberiformis, hymenium in shades of whitish, through ocher-yellow to a more or less dark brown and dark, almost black and brown; outward surface smooth or downy, whitish or coloured like hymenium; stalk if present, less/more cylindrical, with ribes, full or hollow with chamber, whitish or brownish surface with a pinkish or purple reflex; flesh broadly at the intersection. Microscopic features asci operculate eightspored, uniseriate, J-, cylindrical, base type pleurorinque; paraphyses at top clavate, with brownish intercellular pigment; ascospores ellipsoidal, fusiform or spherical with characteristic extensions (apicule) at the poles, hyaline, smooth or ornamented with cyanophilous episporium in maturity, with four nuclei, containing one to three large oil drops; excipulum composed of intertwined cells in the form of textura intricata rarely of textura angularis which is in the ectal excipulum. In regards to mutual generic boundary of particularly complex entity of the family Discina Neogyromitra Maublancomyces Paradiscina Gyromitra Pseudorhizina, to date, acceptance has not been established. The discussion about relationship between Discina and Gyromitra is characterized by mutually opposing opinions. They can be divided into three groups. The first group of authors (Dennis, 1978; Breitenbach & Kränzlin, 1984; O Donnell et al., 1997; Vizzini, 2003 and Medardi, 2007) believe that Discina and Gyromitra are two autonomous genera. Another group (Eckblad, 1968; Donadini, 1984, 1987; Luchini, 1986) has the opinion that they are subgeneric entities of genus Discina. The point of view of the third group (Harmaja, 1969a, 1969b, 1973, 1976a, 1976b; Kotlaba & Pouzar, 1974; Pfister,

3 Gyromitra Fr. sensu lato (Discinaceae, Pezizales) in Montenegro 1980; Abbott & Currah, 1997; Eriksson et al., 2004; Medel & Marmolejo, 2005; Lumbsch & Hundorf, 2007; Van Vooren & Moreau, 2009a) is that they are subgeneric entities of genus Gyromitra. For complete information of this complex issue we refer to: Benedix (1969, 1972) McKnight (1969), Harmaja [1969a, 1974 (1973)], Donadini (1987), Abbott & Currah (1997). In this paper, we rely on the infra-generic systematic proposed by Abbott & Currah (1997), followed by Van Vooren & Moreau (2009) and supplemented by Van Vooren (2009). Also, we follow the position of O Donnell et al. (1997) in terms of belonging of this genus to the family Discinaceae not Abbott & Currah (l.c.), which classify it in Helvellaceae. Subgenus Gyromitra, section Gyromitra Type: G. esculenta The results of research Basic taxonomic data: all species of subgenus Gyromitra, unlike other subgenera, have stipitate ascomata. In addition they are characterized by irregularly shaped cap with cerebriformis surface or more or less grouped irregular lobe. Ascospores are ellipsoidal or fusiform, smooth (observed by optical microscope) and generally, with two polar drops. Gyromitra esculenta (Pers.) Fr., 1849, Summa veg. Scand., Sectio post. p. 346 (1849). Basionym: Helvella esculenta Pers., Comment fung. Bav. p. 64 (1800); Helvella esculenta Pers.: Fr., Syst. mycol. II (1), p. 16 (1822). Physomitra esculenta (Pers.) Boud., Icon. mycol., liste prélim., p. [2] (1904). = Helvella mitra Schaeff., Fung. Icon., 2, pl. 160 (1763). = Gyrocephalus aginnensis Pers., Mém. Soc. linn. Paris, 3, p. 77 (1824). = Helvella sinuosa Brondeau, in Pers., Mém. Soc. linn. Paris, 3, p. 77 (1824). = Gyromitra suspecta (Krombh.) Fr., Summa Veg. Scand., sect. post., p. 346 (1849), after Nannfeldt (1932). = Helvella suspecta Krombh., Naturgetr. Abbild. Schwämme 3, pl. 16 (1834). = Gyromitra suspectus (Krombh.) Rehm, Rabenhorst s Kryptog.-Fl., 1 (3), p (1896), comb. superflua. = Gyromitra suspecta (Krombh.) J. Schröt., Schles. Kryptofl. III, 2, p. 27 (1908), comb. superflua. = Maublancomyces suspecta (Krombh.) Herter, Rev. Sudam. Bot., 10 (1), p. 17 (1951). = Gyromitra queletii (Schulzer) Sacc., Syll. fung., 8, p. 17 (1889), after Kotlaba & Pouzar (1974). = Helvella queletii Schulzer, Hedwigia, 24 (4), p. 149 (1885). 121 Selected iconography: Cooke (1878) fig. 328; Boudier ( ), pl. 224; Marchand (1971) n 97, photo above; Cetto (1976), n 362; Breitenbach & Kränzlin (1981) n 11; Lincoff (1981) n 714; Phillips (1991) p. 302; Dähncke (1993) p. 1106; Roux (2006) p Literature: Harmaja (1979); Van Vooren & Moreau (2009b); Abbott & Currah (1997). Distribution: widespread in Europe, Asia and North America.

4 Branislav Perić & Olgica Perić New findings: - Montenegro, Žabljak, Mlinski potok (Mill brook), 23. V 2010, one specimen on the ground in forest Abies alba Mill. and Picea abies (L.) H. Karst, leg. B. Perić & O. Perić, exsicc. Dgf/C5D Montenegro, Hum Orahovski, 21 V 2010, three specimens near dry stump of Pinus heldereichii Christ., leg. B. Perić & O. Perić, exsicc. Dgf/C5D Note: Cosmopolitan, polymorphic species, recognizable in the field. However microscopic inspection is necessary for reliable determination. Not recommended for consumption due to toxic and carcinogenic properties. Gyromitra neuwirthii Velen., České Houby 4-5: 894 (1922) Basionym: Gyromitra neuwirthi Velen. České Houby, p. 894 (1922).? = Gyromitra esculenta var. fragilis A. Marchand ex Réaudin, Documents Mycologiques 34 ( ): 82, = Gyromitra esculenta var. fragilis A. Marchand, Champ. du Nord et du Midi, 1, p 212, 1971, nom. inval. after Article Selected iconography: Marchand (l.c) n 97, bottom photo; Réaudin (2008) p. 78 and 80. Literature: Velenovský (1.c); Marchand (1971); Réaudin (l.c); Van Vooren & Moreau (2009b) Distribution: Czech Republic, France, Montenegro. References: Perić & Perić (2005 sub nom. G. esculenta) Our findings: - Montenegro, Hum Orahovski, 30 V 2002, above the place where is the mountain cottage, a dozen specimens on the ground in the Bosnian Pine forest, near rotten trunk, leg. B. Perić, exsicc. JF30(0-6)30. Published in Perić & Perić (2005 sub nom. G. esculenta) - Montenegro, Hum Orahovski, 27 IV 2006, five specimens on the ground in Bosnian Pine Forest, leg. B. Perić & O. Perić, exsicc. Dgf/N Montenegro, Hum Orahovski, 21 V 2010, two specimens on the ground in the Bosnian Pine forest, on the burned place, a few meters from the G. esculenta, leg. B. Perić & O. Perić, exsicc. Dgf/C5D a Note: There is no type material for this species. On the other hand, even in the collection of Marchand s corresponding dried specimens lack. Only Réaudin (l.c.) based on new findings valided its variety. Van Vooren & Moreau (2009a) express skepticism towards this variety, considering that the differences with the type are minimal to justify its creation.

5 Gyromitra Fr. sensu lato (Discinaceae, Pezizales) in Montenegro Gyromitra tasmanica (Berk.) Berk. & Cooke, in Cooke, Mycographia seu Icones Fungorum, 1: 193, tab. 90, fig. 331, Basionym: Helvella tasmanica Berkeley (herbarium). Selected iconography: Cooke (1878) fig. 331; Torre (1976) p. 33; Calonge & Torre (1977). Literature: Cooke (1.c.); Raitviir (1965); Torre (1.c.); Calonge & Torre (l.c.); Van Vooren & Moreau (2009b). Distribution: Tasmania, New-Zealand, Spain, Montenegro. Our findings: - Montenegro, Durmitor, Žabljak, Mill Brook, 01. VI 2008, a few specimens on sandy soil and moss in the A. alba and P. abies forest, leg. O. Perić & B. Perić, exsicc. Dgf/ C5D Montenegro, Žabljak, Mill Brook, 29. V 2010, tenth specimens on soil and moss in the A. alba and P. abies forest, leg. B. Perić & O. Perić, exsicc. Dgf/C5D Montenegro, Plav, Hrid, 06. VI 2010, on moist sandy soil in the A. alba and P. abies forest, ten specimens, on two sites hundred meters away from each other, exsicc. Dgf/ C5D Note: Raitviir (l.c.) thought that this is a good species which differs from G. esculenta by the free edge of the pileus, as Morchella semilibera. Species inhabit the Southern Hemisphere of Planet primarily. The first European record originated from Spain. Our records are the second for the European area. 123 Gyromitra infula (Schaeff. : Fr.) Quél., Enchir. fung.: 272, Basionym: Helvella infula Schaeff., Fungorum qui in Bavaria et, 4, ind. 105 (1774); Helvella infula Schaeff : Fr., Syst. mycol., II (1), p. 17 (1822). Physomitra infula (Schaeff.: Fr.) Boud., Icon. mycol., liste prélim., p. [2] (1904). = Phallus triceps Muell., Fl. Dan., 5, pl. 835 (1780). = Helvella mitra Afzel., Kgl. Vet. Akad. Nya Handl., 4, p. 303 (1783). = Helvella brunnea Gmel., Syst. Nat. Linn., éd. XIII, 2, p (1791). = Helvella erythrophaea Pers., Mycol. Europaea, 1, p. 211 (1822). = Helvella rhodopoda Krombh., Naturgetr. Abbild. Schwämme, 3, pl. 19, fig (1834). = Physomitra infula var. rhodopoda (Krombh.) Boud., Hist. class. Discom. Europe, p. 35 (1907). = Helvella friesiana Cooke, Mycographia, 1, p. 195 (1875). =Physomitra infula var. friesiana (Cooke) Boud., Hist. class. Discom. Europe, p. 35 (1907). Helvella infula f. friesiana (Cooke) Bres., Icon. mycol., pl (1932). = Gyromitra inflata (Cumino) Cooke, Mycographia, p. 248 (1879), nom. inval. =Helvella inflata Cumino, Act. Acad. Reg. Taurin., 8, p. 250 (1805), illeg., non H. inflata Schaeff. (1774). =Maublancomyces inflatus (Cumino) Herter, Rev. Sudam. Bot., 8 (5), p. 160 (1950), nom. inval.

6 Branislav Perić & Olgica Perić = Gyromitra infula f. gyrosa Benedix, Kulturpflanze, 14, p. 363 (1967), nom. inval.? = Gyromitra birretum Krombh., Naturgetr. Abbild. Schwämme, 3, p. 33, pl. 21, fig (1834). Selected iconography: Cooke (1.c) fig. 334; Boudier (l.c.) pl. 223; Marchand (l.c) n 98; Cetto (1976) n 364, sub nom. Helvella infula); Breitenbach & Kränzlin (l.c.) n 13; Linkoff (l.c,) n 719; Phillips (l.c.) p Literature: Abbott & Currah (1997); Harmaja (1969b). Distribution: The species is widespread in Europe. Also in Asia, Turkey (Sesli, 1998), Japan, Bhutan, North America (Kempton & Wells, 1973; Abbott & Currah, (l.c.); Medel, 2006). References: Perić & Perić (1997, 1999). New record: - Montenegro, Komovi, Lučka River, Potok od Bijele vode (Stream from the White Water), 02 X 2006, a specimen on the ground covered with sawdust after cutting, in a mixed beech and fir forest, leg. B. Perić exsicc. Dgf/C20D Note: Despite the significant morphological variability (Benedix, 1960) it is easily recognized in the field, but because of possible confusion with G. ambigua it requires a microscopic analysis. It occurs in autumn, unlike others from the same genus. Subgenus Discina (Fr.) Harmaja section Discina: Typ : G. perlata (Fr.) Harmaja Basic taxonomic data: the species are recognized as sessile or subsessile, apothecia fleshy. Only recently Abbott & Currah (l.c.) noted anatomical and microscopic affinities of these species with the species of subgenus Gyromitra. Ascospores usually are ellipsoidal, sometimes fusiform, with one or three drops, smooth or ornamented, always with apicule at the poles, which can be reduced as a blunt extension. Gyromitra perlata (Fr. : Fr.) Harmaja, Karstenia, 9, p. 11 (1969). Basionim: Peziza perlata Fr.: Fr., Syst. mycol., II (1), p. 43 (1822). Discina perlata (Fr. : Fr.) Fr., Summa veg. scand., sect. post., p. 348 (1849). = Discina venosa (Pers.) Fr., after Velenovský (1934). = Rhizina helvetica Fuckel, Symbol. mycol., 2, p. 66 (1873), after Velenovský (1922) Discina helvetica (Fuckel) Sacc., Syll. fung., 8, p. 101 (1889).? = Gyromitra macknightii Harmaja, Karstenia, 26 (2), p. 42 (1986).? = Gyromitra ancilis (Pers.: Fr.) Kreisel, Boletus, 1, p. 29 (1984). Peziza ancilis Pers., Mycol. Europ., 1, p. 219 (1822); Peziza ancilis Pers.: Fr., Syst. mycol., II (1), p. 43 (1822).

7 Gyromitra Fr. sensu lato (Discinaceae, Pezizales) in Montenegro Aleuria ancilis (Pers.: Fr.) Gillet, Champ. Fr., Discom., p. 36 (1879). Acetabula ancilis (Pers.: Fr.) Lambotte, Fl. mycol. belg. 2, p. 573 (1880). Helvella ancilis (Pers.: Fr.) Quél., Enchiridion fung., p. 275 (1886). Discina ancilis (Pers.: Fr.) Sacc., Syll. fung., 8, p. 103 (1889). Selected iconography: Boudier (l.c.) pl. 252; Marchand (l.c.) n 99; Cetto (1976) n 822; Breitenbach & Kränzlin (l.c.) n 31; Lucchini (198) p. 45; Phillips (l.c) p. 304; Medardi (2006) p. 53. Literature: Frencken (1985 ); Abbott & Currah (l.c.). Distribution: Cosmopolitan species, widely distributed in mountain areas in Europe. Present also on the North American continent, North Africa and Asia (Japan and India). New findings: - Montenegro, Trešnjevik, 03 V 2006, two specimens on tree stump of fir, leg. B. Perić & I. Perić, exsicc. Dgf/N Montenegro, Jezerine, on the place which is left of the ski resorts, along the creek that leads to Vranjak, 09 V 2006, more specimens (at 5 sites) on trunks and stumps of fir and spruce, that have begun to rot but otherwise on bare sandy ground, leg. B. Perić, exsicc. Dgf/ C20D Montenegro, Žabljak, Mill Brook, on the three sites from the Mill to Zminje Lake, 29 V 2010, on the ground and rotten stumps of fir and spruce, leg. B. Perić & O. Perić, exsicc. Dgf/C5D Montenegro, Hrid, above Plav, from katun Bajrović (dwelling for farmers during the summer gasing) to the Hrid lake, 06 VI 2010, a few specimens in three localities on the ground and spruce stumps, leg. B. Perić & O. Perić, exsicc. Dgf/C5D Note: The determination of this species is quite easy, but requires microscopic examination of mature ascospores due to possible confusion with related species. Gyromitra geogenia (E. Rahm ex Donadini) ) Harmaja, Karstenia 26 (2), p. 42 (1986). Basionym: Discina geogenia E. Rahm ex Donadini, Mycol. Helv., 1 (4), p. 254 (1985). = Discina perlata var. geogenia E. Rahm, Schweiz. Z. Pilzk., 48 (7), p. 75 (1970), after Article Paradiscina geogenia (E. Rahm) Benedix, Kulturpflanze, 19, p. 178 (1972), nom. inval. Gyromitra geogenia (E. Rahm) Harmaja, Karstenia, 15, p. 30 (1976), nom. inval. = Gyromitra albofuliginosa Donadini, Bull. Féd. mycol. Dauphiné- Savoie, 76, p. 29 (1980), nom. inval. Selected iconography: Rahm (1970); Roffler (2004) p ; Van Vooren (2006) p. 13; Perić (2009) p

8 Branislav Perić & Olgica Perić Literature: Rahm (l.c.); Donadini (1985); Harmaja (l.c.); Perić (2009). Distribution: Switzerland, France, Montenegro. Our findings: see Perić (l.c.). Note: At first glance it is similar to G. perlata, but differs by shade of olive color and conical shape. It occurs along the edge of melting snow. Until recently it was known only in the area of the Alps and the Pyrenees and more recently in Montenegro, in the area of Mt. Bjelasica and Massif Žijevo. 126 Gyromitra megalospora Donadini & Riousset, Bull. trimest. Soc. mycol. Fr. 92(3): 314, Basionym: Gyromitra megalospora Donadini & Riousset, Bull. trimest. Soc. mycol. Fr., 92 (3), p Discina megalospora (Donadini & Riousset) Donadini & Riousset, Mycol. Helv., 1 (4), p. 254 (1985). = Gyromitra megalospora Donadini, Bull. Soc. linn. Provence, 28, p. 71 (1975), nom. inval.? = Discina radiosensilis Falck, Mykol. Untersuch. Ber., 2, p. 119 (1916).? = Discina pseudospora Monti, Tomasi & Maccioni, Micol. Ital., XXX (1), p. 24 (2001), nom. inval. Selected iconography: Donadini & Riouset (1976) p. 313; Perić (2000) p Literature: Donadini & Riousset (1976); Perić (l.c); Van Vooren & Moreau (2009b). Distribution: France, Montenegro. Our findings: see Perić (l.c.). New findings: - Montenegro, Podgorica, 14 II 2007 artificial Pinus halepensis forest stands along the road towards Petrovac, on the ground near by rotten stumps or on them, leg. B. Perić, exsicc. Dgf/C5D Montenegro, 03 III 2007 Podgorica, Golubovci in a stand of P. halepensis. On the ground next to stumps, leg. B. Perić, Dgf/C5D Note: This is a typical Mediterranean species which develops in the winter, on the stumps and humus of Aleppo pine (Pinus halepensis) and on the sandy soil near by them. It has an extremely large ascospores (over 40 μm) ornamented surface with large apicules. It is often replaced with G. perlata, which requires revision of previous collections.

9 Gyromitra Fr. sensu lato (Discinaceae, Pezizales) in Montenegro Subgenus Discina (Fr.) Harmaja, section Pseudogyromitrae Van Vooren Typ : G. gigas (Krombh.) Quél. Syn: genus Gyromitra, section Cerebriformae Cooke, Mycographia, p. 248 (1879), pro parte. Gyromitra gigas (Krombh.) Quél., Mém. Soc. émul. Montbéliard, sér. 2, 5, p. 338 (1873). Basionym: Helvella gigas Krombh., Naturgetr. Abbild. Schwämme, 3, p. 28 (1834). Mitrophora gigas (Krombh.) Léveillé, Ann. sci. nat., sér. 3, botanique, 5, p. 250 (1846). Neogyromitra gigas (Krombh.) S. Imai, Bot. Mag. (Tokyo), 52, p. 358 (1938). Maublancomyces gigas Herter, Rev. Sudam. bot., 8 (5), p. 161 (1950). Discina gigas (Krombh.) Eckblad, Nytt Mag. Bot., 15 (1-2), p. 99 (1968). = Gyromitra gigas var. pumila Velen., Monogr. Disc. Bohem., p. 389 (1934). = Gyromitra curtipes Fr., Ätl. Sw., 34, pl. 56 (1861). = Maublancomyces curtipes (Fr.) Herter, Rev. Sudam. bot., 10 (1), p. 17 (1951). = Gyromitra ussuriensis Lj. N. Vassiljeva, Notulae Syst. Sect. Crypt. Inst. Bot. Acad. Sci. U.S.S.R., 6 (7-12), p. 189 (1950), after Raitviir (1970). = Neogyromitra ussuriensis (Lj. N. Vassiljeva) Raitv., Soobsch. Akad. Nauk. Soyuza SSR, Sibirsk. Otdel. Dal nevost Fil., 23, p. 53 (1964), nom. inval. Selective iconography: Cooke (l.c.) fig. 327; Cooke (l.c.) fig. 329, sub nom. G. curtipes; Boudier (l.c) pl. 221; Boudier (l.c) pl. 222, sub nom. G. curtipes; Cetto (1976, n 809); Cetto (1987) n 2086, sub nom. G. curtipes; Breitenbach & Kränzlin (l.c) n 12; Lucchini (1986) p. 47); Medardi (2006) p. 313 sub nom. Discina gigas. 127 Literature: Raitviir (1970); Torre (l.c); Abbott & Currah (l.c); Benkert (2000); Medardi (2006). Distribution: Widespread species recorded in Switzerland, France, Spain, Italy, Netherlands, Poland, Hungary, Estonia, Russia, Sweden, Finland, Norway, Latvia, Germany, Turkey, Montenegro etc.. References: Perić & Perić (1997 sub nom. G. esculenta) Our findings: - Montenegro, Mojkovac, Minići, in the property Minić, 10 V 1995, on the ground under Populus tremula near by fir stump and trunks, gregarious, leg. Milorad Budiša, exsicc. Bf7(16-21, 26-28). - Montenegro, NP Biogradska gora, near by the ascent to Goleš, 02 V 2006, one specimen on the ground next to rotten fir trees in the mixed forest of beech and fir, leg. B. Perić, exsicc. Dgf/C20D Montenegro, Žabljak, along the left bank of Otoka, 22 V 2009, four specimens on the ground in the spruce forest, leg. B. Perić. & O. Perić, exsicc. Dgf/C5D Montenegro, Žabljak, Mlinski potok (Mill Brook), 23 V 2009, a specimen on the ground in the spruce forest, leg. B. Perić. & O. Perić, exsicc. Dgf/C5D

10 Branislav Perić & Olgica Perić Note: Spring species, with characteristic appearance, with a protruding ribbed stem. It can only be replaced with G. fastigiata (Krombh.) Rehm and G. ticiana Littini, which are not so widespread. Subgenus Caroliniana Abbott, section Caroliniana Typ: G. caroliniana (Bosc : Fr.) Fr. Syn.: genus Gyromitra, section Cerebriformae Cooke, Mycographia, p. 248 (1879), pro parte. Basic taxonomic data: stipitate and non stipitate species. Ascospore are generally ornamented with network, pseudo network or warty ornamented. 128 Gyromitra fastigiata (Krombh.) Rehm, Die Pilze, in Rabenhorst Kryptogamen Flora von Deutschland, Österreich und der Schweiz, 2nd ed 1: 1194, (1896). Basionim: Helvella fastigiata Krombh., Naturgetr. Abbild. Schwämme, 3, p. 28 (1834). Physomitra infula var. fastigiata (Krombh.) Boud., Hist. class. discom. Europe, p. 35 (1907). Maublancomyces fastigiatus (Krombh.) Herter, Rev. Sudam. bot., 10 (1), p. 17 (1951). Discina fastigiata (Krombh.) Svrček & J. Moravec, Česká Mykol., 26 (1), p. 5 (1972). Neogyromitra fastigiata (Krombh.) McKnight, Mycologia, 60, p. 725 (1968). = Gyromitra krombholzii Bezděk, Houby jedlé a jim podobné jedovaté, p. 199 (1905), nom. inval. = Gyromitra pratensis Velen., Mon. Discom. Bohemiae, 1, p. 389 (1934). =Maublancomyces pratensis (Velen.) Herter, Rev. Sudam. bot., 8 (5), p. 160 (1950). = Gyromitra brunnea Underw., sensu auct. Selected iconography: Svrček & Moravec (1972) pl. 81; Kotlaba & Pouzar (1974) pl. I et II; Cetto (1983) n 1648; Cherubini & Perrone (1992) p. 35; Röllin (1995) p. 9. Literature: McKnight (1968); Kotlaba & Pouzar (1974); Abbott & Currah (1997). Distribution: Czech Republic, Germany, Hungary, Sweden, Denmark, Norway, France, Italy, Montenegro. Our findings: - Montenegro, NP Biogradska gora, 24. IV 2006, near the water station bungalows, in mixed deciduous forest on the humus rich soil, in dense colony of Allium ursinum L., leg. B. Perić, exsicc. DgfN/ Montenegro, Hum Orahovski, 08 V 2007, in the beech forest (Fagus moesiaca), by the path horse, on the humus substrate, leg. B. Perić, exsicc. Dgf/C Note: Very rare species, occurring in spring as G. gigas of which is easily distinguished by ornamented ascospores.

11 Gyromitra Fr. sensu lato (Discinaceae, Pezizales) in Montenegro Subgenus Caroliniana Abbott, section Parmae Typ: G. parma (J. Breitenb. & Maas G.) Kotlaba & Pouzar Gyromitra parma (J. Breitenb. & Maas Geest.) Kotl. & Pouzar, Česká Mykol. 28(2): 91 (1974). Basionym: Discina parma J. Breitenb. & Maas G., Proc. Konink. Nedel. Akad. Wetensch., ser. C, 76, p. 103 (1973). Gyromitra parma (J. Breitenb. & Maas G.) Donadini, Bull. Féd. mycol. Dauphiné-Savoie, 76, p. 29 (1980), nom. inval. after Article Gyromitra parma (J. Breitenb. & Maas G.) Bozonnet, Bull. ann. Féd. Centre-est hist. nat. et mycol., 1, p. 9 (1980), comb. superflua. Selected iconography: Hočevar (1978) p. 171; Breitenbach & Kränzlin (l.c.) n 30; Cetto (1993), n 822; Röllin (1993) p. 5; Tentori (1999) p. 218; Perić & Perić (2005a) p. 12; Riva (2005) p ); Medardi (l.c.) p. 52. Literature: Breitenbach & Maas Geesteranus (1973) Bregazzi 1978 (1978); Dissing (l.c.). Distribution: France, Spain, Germany, Denmark, Switzerland, Italy, Slovenia, Croatia, Hungary, Montenegro. References: Perić & Perić (l.c.). 129 Note: it develops on rotten stumps and trunks of deciduous trees (Fraxinus, Ulmus, Corylus, Alnus and Fagus) and spruce. It is a characterised by distinctive ornamentationed ascospores. Gyromitra martinii Donadini & Astier, Bull. Soc. mycol. Fr., 90 (3), p. 193 (1974). Basionym: Gyromitra martinii Donadini & Astier, Bull. Soc. mycol. Fr., 90 (3), p. 193 (1974). Discina martinii (Donadini & Astier) Donadini & Astier, Mycol. Helv., 1 (4), p. 254 (1985). Selected iconography: Donadini & Astier (1974) p. 191; Perić (2000) p Literature: Donadini & Astier (l.c.); Perić (l.c.); Van Vooren & Moreau (2009b). Distribution: France, Montenegro. Our findings: see Perić (l.c.). New data: - Montenegro, NP Biogradska gora, in two sites, 24 IV 06, near the water supply station, a lot of samples on the fallen rotten beech trunk, leg. B. Perić, O. Perić & D. Vincek, exsicc. DgN/

12 Branislav Perić & Olgica Perić - Montenegro, NP Biogradska gora, near by the Biograd Lake, 02 V 2006, more specimens on the fallen rotten beech trunks with the bark, leg. B. Perić & O. Perić, exsicc. DgN/ Montenegro, Kolašin, Jezerine, above streams flowing from the direction of Vranjak, 15 IV 2007, on beech tree trunks, in two sites. The material was not mature enough, leg. B. Perić & M. Prijović. exsicc. Dgf/C Montenegro, NP Biogradska gora, 27. IV 2010, many specimens on the flattened rotten beech trunks with the bark, leg. O. Perić & B. Perić, exsicc. Dgf/C5D Montenegro, Komovi, Kurlaj, above the River Lučka, 09 V 2010, two specimens on the beech tree trunks in the snow, leg. B. Perić & I. Perić, exsicc. Dgf/C5D Figs. 1. Ascospores Gyromitra s.l. in Montenegro (Photos and arrangement B. Perić) 1. G. esculenta (in water); 2. G. neuwirthii (a- in Cotton blue, b- in water); 3. G. tasmanica (in water); 4. G. infula (in KOH 3%); 5. G. perlata (a- in Cotton blue, b- in water 6. G. geogenia (in Cotton blue); 7. G. megalospora (a- in Cotton blue, b-in water); 8. G. gigas (in water); 9. G. fastigiata (in Cotton blue); 10. G. parma (in water); 11. G. martinii (a- in Cotton blue, b- in water); (bar = 20 μm)

13 Gyromitra Fr. sensu lato (Discinaceae, Pezizales) in Montenegro 131 Figs. 2. Gyromitra species in Montenegro (Photos and arrangement B. Perić) 1, 3. G. tasmanica, 2. G. geogenia, 4,6. G. martinii, 5. G. neuwirhii, 7, 10. G. gigas, 8. G. esculenta, 9. G. parma, 11. G. fastigiata, 12. G. megalospora, 13. G. infula, 14. G. perlata

14 Branislav Perić & Olgica Perić Note: It develops on dry beech stumps and trunks. A similarity with G. parma, which also develops in hardwoods, has led to the hypothesis that it may be a unique species (Baute, 2006; Van Vooren & Moreau 2009b). According to current insights, it looks that in Montenegro is not so rare. Discussion taxa from the genus Gyromytra Fr. sensu lato presented in the paper were collected in Montenegro and studied during over the past ten years. During the preparation of this paper, we revised some of the earlier records and now we give updates. The material previously reported as Gyromitra esculenta is shown to be G. gigas due to as immature ascocarp as our incorrect determination. That was at the beginning of our work when we had less experience. As regards current taxonomic knowledge of the genus, the material previously reported as G. esculenta according to Van Vooren & Moreau (2006) now is recognized as G. neuwirthii. Due to limited space here, detailed macro and micro descriptions that we have done for all the species, are not included. This will be done in a forthcoming paper. We provide the Figure 1. with microphotos of ascospores of all taxa and Figure 2. with photos of all ascomata. At first, the material was collected during the usual spring trips to the mountain sites of Montenegro, and only recently, we have taken systematic visiting certain mountain areas searching for species from the genus Gyromitra. We have visited localities in Continental mountain region Hum Orahovski, National Park Durmitor, National Park Biogradska gora, Hridsko lake above Plav, Jezerine on Mt. Bjelasica, and in Massif Komovi the northern slopes (Trešnjevik and Štavna) and southern slopes Planinica, Bijela voda and Kurlaj. Mediterranean forest in site above Budva was viseted only once, after that it was burned. Sub Mediterranean sites in and near Podgorica (Park Forest: Ljubović, Gorica, Zlatica; forest stands toward Agricultural combines, Tuzi and Golubovci) were also visited. Seasonalan visits to Mt. Durmitor, Mt. Bjelasica and recently Hrid above Plav throught years did not yield the expected results. It appears that the species associated to the genus are not widely distributed in our mountains as it is the case with the Central European and North European regions. The majority of species was collected in the conifer stands of Pinus halepensis, P. heldreichii, Abies alba and Picea abies. Several were found in mixed stands and only two species were found exclusively on the beech. It is interesting to note that G. gigas, which is usually found in coniferous forest, we found in the clearing of beech forest. Regarding to substrate, species occur on rich humus soil, usually near trees, stumps or lodged trunks, on sandy soil and some of them are lignicolous. G. neuwirthii, G. fastigiata and G. tasmanica are rare entities, new for Montenegro. The latter is a mycological element of the Southern Hemisphere. It was observed on the Northern part of our Planet only in Spain and Montenegro. Our records of G. megalospora and G. martinii are notable because these species were recorded only in France. For G. geogenia we found the southernmost point of its range. According to our current insights, G. perlata and G. martinii are widely distributed

15 Gyromitra Fr. sensu lato (Discinaceae, Pezizales) in Montenegro species in the area of Montenegro. That could be said as for G. gigas. We assume that G. megalospora occurs in our coastal forests of pine trees, although we have not visited these sites during the spring. Special attention was devoted to visiting the vicinity of the Black Lake in Mt. Durmitor searching for Gyromitra macknightii Harmaja, where is its locus classicus. All our collections from that localities were determinated as D. perlata. Abbott & Currah (1997) reduced G. macknightii to synonym of G. perlata. Therefore the localities on Mt. Durmitor merit further monitoring. The mutual generic boundary of particular complex entity of the family Discinaceae, Discina Neogyromitra Maublancomyces Paradiscina Gyromitra Pseudorhizina, has been intractable problem for decades. We expect that it should be resolved soon by biomolecular research. Therefore, our collections of dried specimens will be proccesed by this analyzes. Literature cited Abbott S. P. & R. S. Currah, The Helvellaceae: systematic revision and occurrence in Northern and North-western North America. Mycotaxon, 62: Baute M.-A., A propos d un Discina de notre région. Bulletin de la Société mycologique de Béarn, 112: 5-7. Benedix E. H., Helvella (Gyromitra) infula Schaeff. im sächsisch-bömischen Grenzgebiet. Česká Mykologie, 14 (1): Benedix E. H., Art- und Gattungsgrenzen bei höheren Discomyceten. III. Die Kulturpflanze, 17: Benedix E. H., Art - und Gattungsgrenzen bei höheren Discomyceten. IV. Die Kulturpflanze, 19: Benkert D., Die Helvellaceen von Brandenburg und Berlin: Erkennung, Ökologie, Verbreitung. Verhandlungen des Botanischen Vereins von Berlin und Brandenburg, 133: Boudier E., Histoire et classification des Discomycètes d Europe. Paris, Éd. Paul Klincksieck, 222 p. Boudier E., Icones mycologicæ, ou Iconographie des champignons de France. Paris, Éd. Paul Klincksieck. 4 vol. Breitenbach J. & R. A. Maas Geesteranus, Eine neue Discina aus der Schweiz. Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen, series C., 76 (1): Breitenbach J., & F. Kränzlin, Champignons de Suisse. Tome 1. Les Ascomycètes. Lucerne, Mykologia, 310 p. Bregazzi R., Discina parma Breitb. & Maas G. in Kassel gefunden. Zeitschrift für Mykologie, 44 (2): Calonge F. D. & M. (de la) Torre, Gyromitra tasmanica (Berk.) Berk. & Cooke, new to Europe. Transactions of the British Mycological Society, 68: Carlsen T. A. & Ø. Stensrud, Hattmorkelen Gyromitra longipes Harmaja funnet 133

16 Branislav Perić & Olgica Perić 134 i Norge. Blyttia, 61: Cherubini A. & L. Perrone, Gyromitra fastigiata (Kromb.) Rehm. Bollettino dell Associazione micologica ed ecologica Romana, 9 (27): Cetto B., I funghi dal vero. 1. Trento, Arti Grafiche Saturnia, 635 p. Cetto B., I funghi dal vero. 2. Trento, Arti Grafiche Saturnia, 728 p. Cetto B., I funghi dal vero. 4. Trento, Arti Grafiche Saturnia, 690 p. Cetto B., I funghi dal vero. 5. Trento, Arti Grafiche Saturnia, 722 p. Cooke M. C., Mycographia, seu icones fungorum. Vol. I. Discomycetes. London, Williams and Norgate. 6 fasc. Dähncke R. M., Pilze in Farbfotos. Aarau, AT Verlag, 1179 p. Dennis R.W.G., British Ascomycetes. Revised Edition. Vaduz, J. Cramer, 585 p. Deparis L., Journée des espèces rares et intéressantes. Bulletin mycologique et botanique Dauphiné-Savoie, 167: Dissing H., The genus Helvella in Europe, with special emphasis on the species found in Norden. Dansk Botanisk Arkiv, 25 (1): Donadini J.-C. & J. Astier, Un champignon méconnu : Gyromitra (Discina) martinii nov. sp. Bulletin de la Société mycologique de France, 90 (3): Donadini J.-C. & L. Riousset, Un nouveau gyromitre: Gyromitra megalospora nov. sp. Bulletin de la Société mycologique de France, 92 (3): Donadini J.-C., 1985 [1984]. Étude des discomycètes IV. Le genre Discina (1). Mycologia Helvetica, 1 (4): Donadini J.-C., 1987 [1986]. Le genre Discina (Gyromitra) (2). Les espèces connues Variabilité des caractères taxonomiques Scanning (Ascomycètes, Pezizales). Bulletin de la Société linnéenne de Provence, 38: Eriksson O. E., Baral H.-O., Currah R. S., Hansen K., Kurtzman C. P., Rambold G. & T. Laessøe (eds), Outline of Ascomycota. Myconet, 10: [htp:// se/myconet/curr/current.htm.] Frencken J Discina perlata: Tweede vondst voor Nederland. Coolia, 28 (2): 1-2. Harmaja H., 1969a. A wider and more natural concept of the genus Gyromitra. Karstenia, 9: Harmaja H., 1969b. A neglected species, Gyromitra ambigua (Karst.) Harmaja, n. comb., and G. infula s. str. in Fennoscandia. Karstenia, 9: Harmaja H., 1974 [1973]. Amendments of the limits of the genera Gyromitra and Pseudorhizina, with the description of a new species, Gyromitra montana. Karstenia, 13: Harmaja H., 1976a. New species and combinations in the genera Gyromitra, Helvella and Otidea. Karstenia, 15: Harmaja H., 1976b. Scanning electron microscopy of the spores of Gyromitra subg. Gyromitra and. subg. Discina (Pezizales). Karstenia, 16: 6-9. Harmaja H., Notes on Gyromitra esculenta coll. and G. recurva, a noteworthy species of western North America. Karstenia, 19: Harmaja H., Studies on the Pezizales. Karstenia, 26: Hočevar S., Discina parma Breitenb. & Maas G. (Helvellaceae, Ascomycetes) in

17 Gyromitra Fr. sensu lato (Discinaceae, Pezizales) in Montenegro some virgin forest nature reserves of Slovenia, Yugoslavia. Acta botanica Croatica, 37: Kempton P. E. & V. L. Wells, Studies on the Fleshy Fungi of Alaska. VI. Notes on Gyromitra. Mycologia, 65 (2): Kotlaba F. & Z. Pouzar, Dalši lokality svazčitého Gyromitra fastigiata (Krombh.) Rehm Čechaách spoznaámkami k rodové příslušnosti uchačú a destic. Česka Mykologie, 28 (2): Lincoff G. H., The Audubon Society field guide to North American mushrooms. New York, Alfred A. Knopf, 928 p. Lucchini G., Studi preliminari sul genere Discina Fr., sottogenere Discina sensu Eckblad. In: Alcune specie interessanti e nuove proposte di studio. Secondo contributo. XIX Comitato scientifico nazionale. Consorzio Valle Serina, Serina (Bergamo): planches p LumbschT.H. & S. M. Hundorf, Outline of Ascomycota. Myconet, 13: (htp:// Marchand A., Champignons du Nord et du Midi. Vol. 1. Perpignan, Société mycologique des Pyrénées méditerranéennes, 282 p. McKnight K. M., Artifacts on spores of Discinaceae induced by common reagents. Mycologia, 60, p. 725 (1968 McKnight K. M A note on Discina. Mycologia, 61 (3): Medardi G., Atlante fotografico degli Ascomiceti d Italia. Trento, A.M.B., 675 p. Medardi G., 2007 [2006]. Il genere Discina in Italia. Rivista di micologia, 39 (4): Medel R., 2006 [2005]. A review of the genus Gyromitra (Ascomycota, Pezizales, Discinaceae) in Mexico. Mycotaxon, 94: Medel R. & J. Marmolejo, Micromorfología de esporas en algunos especies de Gyromitra s.l. (Ascomycotina, Pezizales, Discinaceae). Revista mexicana de micologia, 21: Nannfeldt J. A Bleka Stenmurklan, Gyromitra gigas (Krombh.) Cke. Friesia, 1: O Donnell K., Cigelink E., Weber N. S. & Trappe J. M Phylogenetic relationships among ascomycetous truffles and the true and false morels inferred from 18S and 28S ribosomal DNA sequence analysis. Mycologia, 89 (1): Perić B., Deux espèces nouvelles pour le Monténégro dans le genre Discina: Discina martinii (Donad. et Astier) Donad. et Astier, et Discina megalospora (Donad. et Riousset) Donad. et Riousset. Mycologia Montenegrina, III (1): Perić B., Gyromitra geogenia, un discomycète rare, nouveau pour la flore mycologique du Monténégro. Ascomycete.org, 1 (3): Perić B. & O. Perić, 2005a. Discina parma, nouvelle espèce de la flore mycologique du Monténégro. Revue du Cercle de mycologie de Bruxelles, 5: Perić B. & O. Perić, 2005b. Makromicete Crne Gore, 46 prilog proučavanju. Mycologia Montenegrina VIII: Perić B. & O. Perić, Diverzitet makromiceta u Crnoj Gori. Glasnik odjeljenja prirodnih nauka CANU 11:

18 Branislav Perić & Olgica Perić 136 Perić B. & O. Perić, Prilog proučavanju makromiceta Crne Gore. Mycologia Montenegrina 2 (1): Phillips R., Mushrooms of North America. Boston, Little Brown and Company, 319 p. Pfister H. D., On Peziza melaleucoides - A species of Gyromitra from the western United States. Mycologia, 72: Rahm E., Über einige Rhizinaceae aus dem Hochtal von Arosa. Schweizerische Zeitschrift für Pilzkunde, 48 (7): Raitviir A., Taxonomical notes on the genus Gyromitra. Eesti NSV Tead. Akad. Toim., biol. seeria, 14 : Raitviir A., Once more on Neogyromitra caroliniana. Botaanika - alased tood, 9: Réaudin D., Macromycètes rares ou nouveaux des Côtes d Armor (Bretagne, France) 1. Documents mycologiques, XXXIV ( ): Riva A., Gyromitra parma, un ascomicete primaverile poco conosciuto. Bollettino del Gruppo micologico G. Bresadola, n. s., 48 (1): Roffler U., Gyromitra geogenia (Rahm) Harmaja. Erdige Lorchel. Schweizerische Zeitschrift für Pilzkunde, 82 (5): Röllin O., Discina parma Breitenbach & Maas Geesteranus, trouve en Haute- Savoie. Bulletin trimestriel de la Fédération mycologique Dauphiné-Savoie, 136: Roux P., Mille et un champignons. Sainte-Sigolène, Pierre Roux, 1224 p. Sesli E., Four interesting records of Pezizales of the macrofungal flora of Turkey. Turkish journal of botany, 22, p Svrček M. & J. Moravec, O druhu Helvella fastigiata Krombholz. Česka Mykologie, 26 (1): pl. 81. Tentori A., Su un ritrovamento di Discina parma. Rivista di Micologia, 42 (3): Torre M. (de la), Aportacion al estudio del genero Gyromitra Fr. em. Harmaja en España. Boletin de la Estacion central de ecologia, 5 (10): Van Vooren N., Ascomycètes, saison Bulletin mycologique et botanique Dauphiné-Savoie, 183: Van Vooren N., Nouveautés taxonomiques. Bulletin mycologique et botanique Dauphiné-Savoie, 193: 28. Van Vooren N. & P.-A. Moreau, 2009a. Essai taxinomique sur le genre Gyromitra Fr. sensu lato (Pezizales), 1. Introductionet systématique. Ascomycete.org 1 (1): 3-6. Van Vooren N. & P.-A., Moreau, 2009b. Essai taxinomique sur le genre Gyromitra Fr. sensu lato (Pezizales), 1. Le genre Gyromitra Fr., sous-genre Gyromitra Ascomycete.org 1 (1): Van Vooren N. & P.-A., Moreau, 2009c. Essai taxinomique sur le genre Gyromitra Fr. sensu lato (Pezizales), 2. Le genre Gyromitra Fr., sous-genre Caroliniana. Ascomycete.org 1 (2): Van Vooren N. & P.-A. Moreau, 2009d. Essai taxinomique sur le genre Gyromitra Fr. sensu lato (Pezizales), 3. Le genre Gyromitra Fr., sous-genre Discina. Ascomycete.org 1 (2): 3-13.

19 Gyromitra Fr. sensu lato (Discinaceae, Pezizales) in Montenegro Velenovský J., České Houby. Prague, Nakladem Ceské Botanické Spolecnosti, 950 p. Velenovský J., Monographia Discomycetum Bohemiae. Pars 1. Prague, 436 p. Vizzini A La disposizione spaziale dei nuclei nelle ascospore di Discina e Gyromitra. Bollettino del Gruppo micologico G. Bresadola, 46 (2):

20