On the Structure, Ecology and Distribution of the Species of Mitrula S.Lat

On the structure, ecology and distribution of the species of Mitrula s.lat. (Ascomycetes, Geoglossaceae)

Esteri Kankainen The Subarctic Research Station of the University of Turku, Turku. Finland

Introduction the notes of KALLIO & KANKAINEN ( 1964, 1966) are also plotted on the maps (Figs. 1, The matter under consideration is a part 3). of my research including the larger fungi in I express my best thanks. for direction and arctic and subarctic regions. For eight years I advice of great value to my teacher Prof. Paa have also collected Mitrula material, and so vo Kallio, for valuable notes and discussion me notes are published in KALLIO & KANKAI to Phil. lie. Tauno Ulvinen, for determinat NEN (1964, 1966 ). I have had the opportunity ion of the mosses in question to Phil. mag. to participate on the excursions of Kevo, the Unto Laine, and for the linguistic revision to Subarctic Research Station of the University Mrs. Linna Miiller-Wille. I have received of Turku under the leadership of Prof. Paavo financial support from Suomen. Kulttuurira Kallio. Our Mitrula collections from these hasto. trips to Spitsbergen in 1966 and northeastern Mit r u l a Fr., Syst. Myc. 1, p. 491. 1821. Canada in 1967 are taken into consideration (sensu strict.) in the present paper. The specimens studied lMAI ( 1941 ) and MAAS GEESTERANUS are preserved in the following herbariums: ( 1964) have unravelled the history of the genus Mitrula, and we only ascertain the lar H Botanical Museum of the Univer ge variation in its content from one extreme sity of Helsinki to the other, from the wide opinion of HFR Finnish Forest Research Institute, KARSTEN ( 1871) and MASSEE ( 1897) to the Helsinki narrow one of IMAI (1941, 1956) and MAAS HPP Department of Phytopathology, GEESTERANUS ( 1964). IMAI ( 1956), divided University of Helsinki Geoglossaceae into new subfamilies. and tribes. KEVO Collections of the Subarctic Re Consequently the tribe Mitruleae includes search Station of the University of only one monotypic genus Mitrula on. the Turku, Utsjoki basis of that the ascocarp of M . paludosa is OULU Botanical Museum of the Univer hemiangiocarpic. In the tribe Gymnomitru sity of Oulu leae is included one genus Gymnomitrula s Botanical Museum of Stockholm with two species G. abietis and G. gracilis TROMS Department of Botany, Troms0 on the basis of that the ascocarp of M . pusilla Museum is gymnocarpic. According to MAAs GEESTE TUR Botanical Museum of the Univer RANUS ( 1964) the genus Mitrula is also mo sity of Turku notypic. He has renewed the old genus Only unpublished finds are included in the H eyderia which is in his opinion monotypic, enclosed lists of the studied specimens, but because the place of M. gracilis is indistinct.

23 In the present paper we have taken the genus cylindrical-spherical cap is pure yellow in Mitrula in a larger sense than the last two colour ( cf. var. cylindrica of VELENOVSKY authors, but sensu stricto compared with the 1934). opinion of e.g. FRIES (1821), NANNFELDT The ascigerous portion is connected to the ( 1942) and EcKBLAD ( 1963). stem without a sterile margin as seen in Mit rula gracilis and Heyderia abietis. The cap 1. Mitrula paludosa Fr., Sy.st. Myc. 1, p. stem relation varies very much depending on 491. 1821. the habitat. In very rotten, soft places the Helvella laricina Viii., Fl. Dauph. 3, p. 1045, Tab. 56. 1786-1789. specimens have thick, short, caespitose stems Clavaria phalloides Bull., Champ. France, p. and small caps ( cf. var. pachyceps of KAR 214, Tab. 463, Fig. 3. 179·1-179-8. STEN 1883); on a more solid substrate the Clavaria epiphylla Dicks., Plant. Crypt. 3, p. 22, stems grow slender and long, the height of Tab. 9, Fig. 10. 1793. Leotia ludwigii, dicksoni, bulliardi Pers., Syn. them depending on the depth of the substrate Fung., pp. 611, 612, Tab. 3, Fig. 13. 1801. in the water. The caps are generally large Leotia laricina ·Pers., Syn. Fung., p. 614. 1801. and variable in form. H elvella aurantiaca C.umino, Act. A cad. Sci. Tour., p. 221. 1805. Anatomy. The width of the hyphae in the Mitrula aurantia Cumino, Fung. Vall. Pi<>ii in central part of the stipe widens up to 15 fl, Act. Tur. 1806. Leotia uliginosa Pers., Myc. Eur. 1, p. 200. in the periphery of the stipe about 6 fl· The 1822. surface of the stem is smooth. Leotia uliginosa var. aurantiaca .Pers., Ibid., p. The size of the asci averages 75- 110 X 5 201. 1822 . - 6 fl (measured in five specimens). The Mitrula pha:lloides Ghev., Fl. Paris, p. 114, Fig. 185. 1826-1827. paraphyses are straight, not at all or very Leotia elegans Berk., London Journ. Bot. 5, p. 6. slightly enlarged above. The spores are hya 1846. line, not punctate, nonseptate, cylindrical to Mitrula elegans Fr., Nov. Symb. Myc., p. 103·. clavate-cylindrical, in average larger than 1851. Mitrula paludosa var. pachyceps Karst., Hedw. those in M. gracilis. The size of the spores in 22, p. 17. 1883. -Karst., Acta Soc. F. Fl. Fenn. the following specimens is: Masku 4. 6. 1959 2, p. 110. 1885. 15.3 X 3.5 fl (50 measurement); Lohja 12. Mitrula phalloides var. pa:chyc('ps Sacc., Syll. 6. 1961 12.2 X 2.5 fl (50 meas.); Kiirninki Fung. 8, p. 33. 1889. Microglossum elegans Underw., Minn. Bot. Stud. 25. 6. 1966 13.7 X 2.6 fl (50 meas.); Inari 1, p . 495. 1>896. 15. 7. 1968 11.1 X 2.6 fl (30 meas.). MitrulGJ laricina Mass., Ann. Bot. 11, p. 271. The morphological and anatomical features 1897. of the specimens studied correspond well with Mitrula norvegica Rostr., Vid.-Selsk. Skrift. 1: 4, p. 5. 1904. those recorded in the literature (FRIES 1821, Mitrula phalloides var. aurantiaca Boud., Icon. KARSTEN 1871, 1883, REHM 1896, MASSEE Myc. 3·, Tab. 427. 1909. 1897, DuRAND 1908, LLOYD 1916, VELENovsKv External morphology. The size and fo-rm 1934, IMAI 1941, MAINS 1955, BENEDIX 1962, of the studied ascocarps of Mitrula paludosa EcKBLAD 1963, MAAS GEESTERANUS 1964) . are quite variable. The stipe is 2-5 centimet Specimens belonging to var. p.achyceps Karst., res high, smooth, transparent, whitish when and possibly to var. sphaerocephala Boud. fresh, orange yellow when dry. The base of and var. cylindrica Vel. are included in the the stem is generally thickened when grown list of the studied specimens. Var. castanea in soft habitat, for instance in mud, forming Vel. is unknown to us as is f. pallens Eisfelder a large white ball of hyphae. When growing & Benedix. MAINS ( 1955, p. 873) has found on only a little decayed leaves the base of three types of ascospmes occurring in the the stem is not at all thickened or very Mitrula paludos.a collections. We have found, slightly. however, spores of very variable fo·rm and The diameter of the ascigerous portion va size in the same specimen. The uniseptate ries from 0.5 to over one centimetre. Mostly spores are very rare in our collections ( cf. the cap is uneven, folded, but also even, al MASSEE 1897, lMAI 1941, MAINS 1955). most spherical caps exist. Sometimes the caps Habitat. Mitrula paludosa grows in Fin resemble Spath ularia. The colour of the cap is land preferably on decaying leaves of birch, yellow - orange yellow when fresh becoming but it is also found on leaves of Alnus, on far darker when dried. The form with even, needles and twigs of Pinus silvestris and Picea

24 abies, and among more rotten rests of plants writer are mostly from Finland (see Fig. 1). (in mud and slime) . Almost without excep The species has never been found in the alpi tion the species is found along springs and ne regions of Lapland, thus above the Betula spring brooks, and in bogs. Often its habitat tortuosa tree limit. In other parts of Finland is a puddle, but always a place where the wa it seems to be common. Many collections ter changes but does not flow. In such places and observations have been made along the the oxygen conditions are good. The habitat Salpausselka ridges. The species is common of our specimen collected in Canada was on this area because of the occurrence of very similar to those in Lapland. springs at the foot of the ridges. The habitats o.f the studied specimens agree Mitrula paludosa occurs in all of Europe with the notes presented in the literature (SACCARDO 1889, IMAI 1941, BENEDIX 1962), (FRIES 1821, KARSTEN 1871, 1883, CooKE in Greenland (LANGE 195 7), North America 1871, REHM 1896, MAssEE 1897, DuRAND (DURAND 1908, MAINS 1955 ), Japan (IMAI 1908, LLOYD 1916, VELENOVSKY 1934, biAI 1941 ), that is to say only in the northern 1941, NANNFELDT 1942, MAINS 1955, BENEDIX hemisphere. EcKBLAD (1963, p. 153 ) has also 1962) . Var. pachyccps of KARSTEN ( 1883 ) noticed that this species does not occur above is grown «Supra acus Pini sylvestris putres the tree limit. He says that «

25 ,. 40' M.Ltrula. ra.ludosa rr: • Colled~ons studied ,o 0 Observai~on.s

200km

Fig. I. The studied collections and observations of M i t r u l a p a l u d o sa Fr.

Saltin (TUR) ; Oripaa Myllylahde 13.7. 1956 Porraskoski 19. 8. 1952 Otto v. Schulmann (H ); L. E. Kari (TUR.), 30. 6. 1968 Martti KemiHi. Tam me I a Mustiala 30. 7. 1-8166·, 23. 7. 1869, (OUILU ); Po mark k u Uusikyla 19. 6. 1939 24. 7. 1869 P. A. Karsten (H) and 27. 6. 1867 A. Matti Laurila (HPP). L.Borenius (H ); Vanaja Isojarvi 8. 7. 1965 Ta. As i k k a I a 15. 6. 1863 ]. P. Nordin (H); Harri T. Toppari (H ). Janakkala Koljala Suurisuo 5. 7.19·68 Pertti Tb. La u k a a Seppala 14. 6. 19'15 Anne-Marie Uotila (H); Jo 'kioinen 22.6.1910S.Salmen Hayren (H), Valkola 18. 9. 1949 E. ] . Valovirta linna (H); Jams a Jamsanniemi Myllyjarvi 26 . (H). 7. 19·64 RaiUo R.uotsalo (H); K u h moine n Kol. Pari k k a I a Kasuri 21. 6. 1955 Otto Puuk!koistenkoski 22 6. 1965 Yrji:i Makinen (TUR.); v. Schulmann (HFR). L am m i Evo 24. 6. 1965 Harri T. Top pari (H), Ka. Sip poI a Enajarvi 6. 7. 1948 Viljo Ku-

26 jala (HIFR), Kaipiainen 26 . 6. 1949 Viljo Kujala St. K an k a an p a a Kuninkaanlahde 18. 6. 428 (HFR); Ve!hkalahti Pyhalto 19. 6.19·47 1957. Lars Fagerstrom (H). Ta. Hausjarvi churoh village 16.6. 1958 ; Kb. Pie I is jar vi Koli 24. 8. 19-56 Viljo Ku J a a I a churdh village Iso Ruhmasjarvi 30. 7. jala (HFR); R aut a v a a r a Suo•jarvi 8. 7. 1952 1956, Retusjarvi 31. 7. 1967, «Ritti« 7. 7. 1963, Albert Korhonen (H); To h m a j a r vi Herne Hartola 6. 7.1963; Kuusankoski Keltti 1G. vaara 27. 6. 1966 Esteri Kankainen (TUR). 6.1959'; Riihimaki 21. 7. 1956 ; Ruovesi Sa. Lappe e Korkeamaki 8. 6. 1906 Hans Buch Jiiminkipohja 30. 6. 1957; Somer n i em i Miiy (H), Vehkasalo 26. 6. 1957 collector unknown (H) ramaki 13 . 6. 1957. and 20. 6. 1959 Kirsti Sandberg (H); R u o k o Ka. Sip poI a Kaipiainen 19. 6. 1953, Pajari I a h t i Inkilanmaki 24. 6. 1967 Terho Poutanen 29. 6. 1953. (TUR) ; Taipalsaari Halila 1. 7.1966 Orvo Kb. Kite e Juurikka-aho along Kaarlola road Vitikainen 206 (H) . 18.7.19·57; Kontiolahti Kulho 15. 7.1957, Sb . K u o pi o Pitkalahti 30. 6. 1949 Ritva Paihola 14. 7.1957 ; Liperi Vaivio 12. 7.1957. Ruotsalo (HFR), SarkilaJhti 9. 6. 1909 K . Linko.Ja Sa. Kangasniemi Paappala .3. 7. 1963 ; (TUR) ; Sii-Jinjarvi Kasurila 14. 6.1912 Lappe e Myllyla 20. 8. 1953; Lei von m a k i Lauri Korhonen (HFR) ; Tervo Talluskyla, in churah village 1. 7. 19•6·3; L u u m ii 'k i Somerhar 193•8 Anu .Malinen (H). ju 22 . 6. 195·3, between Somerharju and Taavetti Oa. ·Lap a art t i Pyhavuori 1. 7. 1948 Art in 195•3, Uro 16. 8. 1963 ; M ii n t y h a r j u Part turi Railonsala (TUR) and 4. 7. 1957 G. Nord simaa 8. 7. 1963; T a i paIs a a r i Karhula strom (H). 12. 8. 195·6, Levainen 12. 8. 1956; Va Ike a I a Om. Lappajarvi Savo 21.7.1904 A.L. Anttila 4. 8. 195•6, Hevosoja 5. 8. 195·6, Mankki Backman (HFR); P u I k k i I a Niskankyla 5. 8. 2. 7. 1953, Utti 8. 7. 1957. 1966 Tauno Ulvinen (OULU); Vim p e I i Kos Ob. K i i mink i Joutenojanpalo 19·. 6. 1963. kela 17.6.1904 A. L.Backman (HFR), PY'ha Lkem. Savukoski Martti in 1961 (cf. UL vuori 22 . 7. 190:3 A. L . Backman (HFR). VINEN 196>3, p. 54) . Ob. Hau 'kipudas Martimo 10.7.1966/ XXVI Martti Ohenoja (OULU); K i i mink i Observations of Lauri Teivainen in HFR Haara 25. 6. 1966/III Martti Ohenoja (OULU), (marked with rings on the enclosed map, Juuans; danmaa 25 . 6. 1967/V Martti Ohenoja Fi<;. 1) : (OULU). Kn. Sot k am o Lontta 29. 6. 1·945 Jaakko ]a Ta. Korpi I a h t i Oittila, in 19·46. las (H). Tb. Jyvaskyla Vesanka, in 1945. Ks. K u us am o Juuma 6. 8. 1948 L. E. Kari Ob. Rovaniemi Pisavaara (two finds), in (TUiR) . 1946-47. Lkem. Pelkosenniemi Kairala15.6.19•37 Lkem. K i t t i I a - E n on t e k i o Pallas-Ounas Richard Frey (H); Sodankyla 24.8.1947 tunturi (three finds), in 19·48-49. Viljo Kujala (HFR). All observations of Lauri Teivainen are from Li. In a r i Kyro, in about 19·20 A. R. Ruoranen wet copse fen. We must, however, be critical with (H), 5 km west of Ivalo 15. 7. 19'68 Yrjo Maki the observations in Ob and Lkem, because there nen ( 68~262) & Marja~Lee n a Halme & Leena is a possibility of confusing Mitrula paludosa and Laitinen (TUR, KEVO), along Ivalo-Inari road M. gracilis. south of Myosajarvi 9. 8. 19·68 Yrjo Makinen (TUR), 4 km south of Tolonen 20·. 7. 1968 Yrjo 2. Mitrula gracilis Karst., Hedwigia 22, Makinen & Esteri Kankainen (TU'R ), Tsharmi p. 17. 1883. tunturit Laanaselka 1. 8. 1968 Esteri Kankainen (TUR) ; U t s j o k i Jesnalvaara 3. 9. 1968 Paa Mitrula muscicola E. Henn., bfvers. Kong!. Vet. vo Kallio (KEVO ). Akad. FoPhandl. 1•88·5, p. 71, Tab. 8, Figs. 6- 8. 1885. USISR Mitrula rehmii Bres., Fung. Trid. 2, p. 41, Tab. Lim. Kant a I a h t i in 1843 Fr. Nylander (H). 147, Fig. 2. 1892. Lpets. Pets am o Alakongas 13 . 7. 1932 E. Mitrula gracilis var. flavipes Peck, Ann. Rep. Hayren (H) . N.Y. State Mus. 49•, p. 32. 1•8·96. Prov. Petropolitanae. Julkki prope Pargala 26. 5./ Gymnomitrula gracilis Imai, J ourn. Fac. Agric. 7. 6. 189>8 legit Tranzschel (S) . (See NAuMov 1964, Hokkaido Imp. Univ. 45, p. 175. 19•41. p . 230) . Gymnomitrula rehmii Favre, Ergebn. wiss. Un CANADA ters. schweiz. Nationalparks 5:42, p. 383. 1960. Ncwfoundland-Labmdor. E s ·k e r east shore of Menihek Lakes west of Esker, 53°5·2'n.lat., 66°26' Typus: In Herb. KARSTEN, Mus. Bot. Univ. w.long., elevation 1600 ft. 23 . 7. 1967 Esteri Kan Helsingfors. Knasa-guba 5. 8. 1961 P. A. Kar kainen (TUR) . sten. List of the observations of Tauno Ulvinen Exsiccate: Fungi exs. Suec., No. 1784. (marked with rings on the enclosed map, Fig. 1): External morphology. Mitrula gracilis IS Ab. Bromarv Hankoniemi 7.8. 1955; Vih a rather inconspicious species in so far as it t i Numrnela Kotokorpi 11. 6. 1960. N. Nurmi jar vi Noppo Perkelsuo 16. 6. does not grow in abundance. It differs from 1958. M . paludosa and H eyderia abietis chiefly in

27 Fig. 2. A circu.laT population of Mit r u l a gracilis Karst. on the Cal lie r E: on sa r m e n to sum mat. Spitsbergen, Kongrcssdalen, in 1966. colour, in SIZe ol asci and spores, and in kinds of specimens are as a rule smaller and habitat. paler, and the stipe is more scaly than in The height of the studied ascocarps varies the former type. They ha e features, even from 0. 7 to five centimetres. The stipe is when adult, similar to the juvenile stage o.f yellowish or ochraceous in hesh material, but M. gracilis. in some specimens it is whitish because of Though the extreme specimens among the the presence of white, partly hyaline scales studied material differ very conspicuously and hyaline hairs. The scales are poorly seen from each other it is not necessary to attach in dry material. The base of the stem is great importance to them as separate species, rugged, somewhat thickened, and divided into or e\·en as variations, because mediator speci many fibres crossing the rhizoids o.f the host mens exist in this material. For instance most moss. The form of the ascigerous portion and specimens from Canada indicate an infra its connexion with the stipe varies between specific series. Some of these are also from that of M itmla paludosa and H eydcria abie Lapland, although it is yery difficult to distin tis. The head and stipe are, however, almost guish this group. Uost material from north the same colour when fresh. ern Fennoscandia is of the capitate form. Two extreme morphological forms can be Some specimens from the northernmost distinguished in Mitrula gracilis material. In Fennoscandian research area. and all mate some specimens the fertile, c a p i t a t e, more rial from Spitsbergen are of the extreme pi or less folded head continues to the stipe leate form. without a distinct sterile zone. In very young Particularly in dry material from Canada, ascocarps of this type, however, a free mar but also in the pileate specimens rests of a gin, filled with mucilaginous matter and han veil between the stem and the head can be ging over the apex, can be seen (Tauno Ulvi seen. When dried or preserved in alcohol the nen, by letter) . colours of M. gracilis, like M. p.aludosa be The specimens o.f the other type have a come far darker, and the stipe and the head pi 1 eat e, unfolded head with a sharp over are no longer unifo-rmly coloured the stipe hanging margin. The surface of the broad being (lemon) yellow, and the head reddish sterile area is somewhat reticulated. These brown.

28 Anatomy. The anatomy does not vary like terial correspond to those of M . rehmii of the external morphology of the species in BRESADOLA ( 1892, p. 41) . HEIM & REMY question. The hyphae in the axis of the stipe ( 1932, p. 68 ) widen M . rehmii and describe are up to 8 f-l wide, in the periphery 4- 5 f,l the «forme alpine«, which according to NANN wide. The hairs are hyaline, one- and two FELDT ( 1942, p. 50 ) is M. gracilis, while the celled, up to 30 f-l long. The scales extend «forme sylvatique subalpine« - Bresadola's upward, and they turn dark blue with cotton original type - should perhaps be different. blue. According to BErM & REMY both forms have The asci average 65- 85 X 6- 7 f-l (mea much larger asci and spores than the type sured in five specimens ). The tips of the asci described by BRESADOLA. The spores are also turn blue with iodine. The paraphyses are very irregular in form. numerous and slightly thickened above. The NANNFELDT (1928) has studied herbarium spores are hyaline, somewhat punctate (best material of M . rehmii, and he says that it is seen with oil immersion), often uniseptate, a good deal larger than M. gracilis. The au more or less straight, fusiform. The size of thor has seen a specimen of the exsiccate the spores in the following collections is: Ra material of LuNDELL & NANNFELDT, and it nua 22. 7. 1943 9.2 X 2.4 f-l (50 measure corresponds well with the capitate type des ments); Kuusamo 6. 8. 1948 9.8 X 2.0 f,l (50 cribed above. meas. ); Kiirninki 21. 7. 1966 11.2 X 2.8 f,l IMAI ( 1941 ) on the basis of DITTRICH's (30 meas. ); Utsjoki 14.8.1963 12.5 X 2.7[-l ( 1898 ) and CoRNER's ( 1930 ) investigations (30 meas.) ; Spitsbergen Kongressdalen 16. 8. separated M. gracilis and M . cucullata to 1966 11 .1 X 2.6 f,l (50 meas.) ; Canada: Beam his own genus Gymnomitrula instead of Hey Lake 6. 8. 1963 11.6 X 2.6[-l (25 meas.), deria, which according to IMAI is already Attikamagen Lake 5. 8. 1967 11.7 f-l X 2.6 f,l occupied. Thus also M . gracilis should be (30 meas.), Highfall Creek 2. 8. 1967 11.2 X gymnocarpic like M. pusilla. But the develop 2.5 f,l ( 25 meas.). The specimens from Ranua, mental phases of M. gracilis have not yet been Kuusamo and Kiiminki are of the capitate clarified, and the possibility that M. gracilis form, the specimens from Utsjoki and Spits might be angiocarpic is not excluded. That is bergen of the pileate form. why we for the present place the species in Mitrula. For instance rests o.f a veil in our Many different observations and opinions specimens may resemble those seen in M. pa have been presented on Mitrula gracilis and ludosa (DURAND 1908, p. 389, NANNFELDT the related species. The notes on the ana 1932, p. 317) . tomical features do not vary greatly, but the In 1949 FAVRE has united M. rehmii in morphology has caused much confusion. M . gracilis and M . muscicola. In 1960 FAVRE KARSTEN ( 1883 ) considers this species to be recorded Gymnomitrula rehmii as synonym between M. elegans and M . paludosa. Du of Gymnomitrula gracilis. MAINS ( 1955, p. RAND ( 1908 ) however, has combined the 8 71 ) has observed some variation in M . gr a mentioned two species as one species M . phal cilis in America. He writes that «

29 BENEDIX (1962) and SvRCEK (1962 ) record tum, H elodium blandowii, H. lanatum, and from the Tatra Mitrula material the place with Philonotis tomentella. The gra.cile, of which in their opinion is uncertain as pileate form in our collections is connected regards M. rehmii and M. gracilis. According also with Calliergon sarmentosum, Oncopho to BENEDIX (p. 404) M. gracilis has a shorter rus wahlenbergii, and with Tomentypnum stipe than M. rehmii. About his own Tatra nit ens. material BENEDIX says that those specimens Very often brown dead patches on moss have no sharp overhanging margin, but they mats around M. gracilis populations have have the uneven head typical of M. rehmii. been found. The phenomen is most obvious We assume that M. rehmii and M. gracilis when a bright green moss species (for instan sensu BENEDIX perhaps correspond in some ce Paludella squarros.a or Calliergon sar degree to the variation described above. mentosum) forms a more or less pure grow BENEDIX does not exclude the standpoint of ing unit, and when the fungus grows in REHM ( 1896) who considers M. gracilis and a!bundance. The most regular growing units M. rehmii variants of the same species. BENE we have seen in Spitsbergen (Fig. 3) DIX, however, placed M. rehmii together with where such brown rings are very conspi M. paludosa and M. omphalostoma in the cuous in some bogs (KANKAINEN & KARL subgenus Physomitrula and M . gracilis toget STROM & HEIKKILA 196 7) . The fungus widens her with M. abietis in the subgenus H eyderia. its growth area centrifugally while in the EcKBLAD (1963) and MAAS GEESTERANUS centre of the dead ring new, vigorous, bright ( 1964) have studied the anatomy of the stipe green moss begins its growth. of Mitrula species. MAAS GEESTERANUS (p. 89) In the literature M. gracilis is recorded says that «the very different structure of the growing on Aulacomnium palustre, Bryum stipe in itself is sufficient proof that Mitrula pseudotriquetrum, Calliergon sarmentosum, gracilis is not a H eyderia. I am not sure, C. stramineum, Dicranum majus (! ), Dre however, as to its relation to Mitrula.« The panocladus badius, Dr. exannulatus, Helo anatomy of the stipe in our specimens nearly dium lanatum, Hylocomium splendens, Palu corresponds to that presented by EcKBLAD della sq;,arrosa, Philonotis fontana, Ph. to ( 1963) and MAAS GEESTERANUS ( 1964) ex mentella, Rhacomitrium canescens, Rh. fasci cept that neither author has observed any culare, Tomentypnum nitens, and on Webi'Ta hairs or scales on the stem. ULVINEN (by nut.ans (HENNING 1885 concerning M. musci letter) has seen scales in his M. gracilis mate cola, BRESADOLA 1892 concerning M . rehmii, rial from En on tekio. Though the existence of DuRAND 1908, HEIM & REMY 1932 con such special structures brings M. gracilis in cerning M . rehmii, NANNFELDT 1942, MAINS some way closer to Heyderia (sensu 1'v1AAS 1955, BENEDIX 1962 concerning M . rehmii GEESTERANUS), it is not justifiable to place it and M. gracilis, EcKBLAD 1963, KALLIO & in Heyderia. In our opinion it is thus rea KANKAINEN 1964) 0 sonable to keep M . gracilis in Mitrula, but Some authors have reported that the moss the study of the juvenile development is underneath the fungus is dead (KARSTEN necessary. 1883, EcKBLAo 1963, KALLio & KANKAINEN 1964). Some authors assume that the fungus Habitat. Mitrula gracilis grows on wet is a parasite (BRESADOLA 1892, DuRAND 1908, mossy places between hummocks in bogs. The IMAI 1941, MAINS 1955). NANNFELDT (1942) studied specimens have been found on many had seen only healthy moss below the fungus. different moss species. It is surprising that EcKBLAD ( 1963 ) writes that he and Mr. S. Si the extreme morphological forms presented vertsen have observed almost circular brown above have no notable preference concerning patches on the moss mats. ULVINEN (by let to the mosses on which they live. Both forms ter) has found M. gracilis (the pileate form are found on Aulacomnium palustre, Dre from Enontekio) killing Aulacomnium panocladus revolvens, and on Paludella palustre. squarrosa. In Lapland Paludella seems to be EcKBLAD (1963, p. 152 ) says that «

30 JZ" ,......

Marula qrocals Karst.

50· 100 200km

Fig. 3. The studied collections of Mit r u l a gracilis Karst. that we should expect a usual host - parasite effect which does not necessarily have to do relation«. Further on EcKBLAD reports he has with the nutrition of the fungus ( cf., too, not seen any fungal hyphae within the cells EcKBLAD 1963, p. 152) . On the other hand of the moss. Neither have we observed fungal the toxic effect may be also a primary effect hyphae penetrating the moss cells. We are increasing the permeability of the cell mem of the opinion that the fungus gets nutrients branes in the host moss. diffusioned from the exuding liquid of the As seen above there is very large variation moss. The toxic effect of the fungus is an which does not seem to depend noteworthy axiom but it can be a secondary handicap on the moss substrate. The variation does not

31 obviously correspond to the «biologic vari collections in Canada are made m the lati ation« of Mitrula abietis (VELENOVSKY 1934, tudes of 54° and 58° n.lat. ULVINEN 1966) because the nutrition is different in M. gracilis and M. abietis. It is Mitrula gracilis is a subarctic-arctic-alpine necessary to know more about the nutrition fungus. It is recorded from the Kola Penin in M. gracilis and about the influence of sula (KARSTEN 1883), from northern Fenno e.g. the temperature on the morphogenesis of scandia (HENNING 1885: M. muscicola, Ro this species before the problem of variation is STRUP 1904, NANNFELDT 1928, 1942, IMAI solved. 1940, EcKBLAD 1963, KALLio & KANKAINEN 1964, 1966), from Spitsbergen (SKrRGIELLO Phenology. Mitrula gracilis is a species 1961' KANKAINEN & KARLSTROM & HEIKKILA between M. paludosa and H eyderia abietis 1967) , Jan Mayen (LARSEN 1924), Iceland as regards the period of growth, for it is (LARSEN 1932), Greenland (RosTRUP 1891 , a species. of summer and autumn. The 1894, LANGE 195 7), from North America capitate specimens in the studied material (DuRAND 1908, MAINS 1955 ), Japan (IMAl seem to be earlier than the pileate types. 1941 ), from the Alps (BRESADOLA 1892: The former is found to occur from the middle M. rehmii, HEIM & REMY 1932: M. rehmii, of July to August 21, the latter from Au FAVRE 1949, 1955, 1960, SKIRGIELLO 1961, gust 6 to September 12. The specimens from ErsFELDER 1962 ), and from the Tatra (BE Canada are collected between July 31 and NEDIX 1962, SVRCEK 1962 ). August 6. The climate fluctuates very much from year to year. For instance the summer List of the specimens studied (marked with and autumn in Finland in 1967 was wanner dots on the map, Fig. 3, together with the than average. In 1968 the summer was excep collections reported by KALLIO & KANKAINEN tional cold in Lapland. 1964, 1966): In Fennoscandia the peak of occurrence HNLAND of M. gracilis is clearly in August, as seen in Ob. K i i mink i rich fen between old Kuu the following table, which presents the samo road and Jolosjoki river (on Aulacomnium collections. reported by KANKAINEN in this ar palustre, Bryum pseudotriquetrum, Drepanocladus revolvens etc. ) 21. 7. 1966 Tauno Ulvinen (OU ticle ( abbrev. K), by KALLIO & KANKAINEN LU), and 2. 8. 19•66 Tauno Ulvinen (OULU, 1964, 1966 (K & K), NANNFELDT 1942 (N) TUR), (on Bryum pseudotriquetrum) 5. 9. 19·68 and ECKBLAD 1963 (E). M. Ohenoja & T. Ulvinen (OULU), southwest of the central ·Swamp of Murtoinsaaret 8. 9. 1968/ XXXV Martti Ohenoja (OULU); Ran u a !so Number of collections palo (on H elodium blandowii, Paludella squarrosa) K K&K N E 22. 7. 1943 A. V. Auer (TUR). During June 1 Ks. Kuus amo Juuma (on Paludella squar rosa) 22. 8. 19'23 Harry Waren, det. Matti Lau 4 2 4 « July 2 rila 1938: Mitrula gracilis Karst. = M. muscicola (( August 16 26 8 10 E. Henning (TUR) . (( September 6 Le. Enontekiii Hirvasvuopio (on Helodium blandowii) 12. 8. 19'61 H. Roivainen (H) and (on Also the notes of the other authors nearly Paludella squarrosa) 10. 8. 1961 Laila & H. Roi vainen (H), Markkina (on Aulacomnium palustre) conform to that reported above as regards 12. 9. 19•67 and 16. 9. 1967 Tauno Ulvinen (OU the period of growth in M. gracilis (KARSTEN LU), Toskaljarvi (on Calliergon sarmentosum, 1883, IMAI 1941, MAINS 1955 ). DuRAND's Drepanocladus exannulatus) Aug., 19·67 A.]. note ( 1908, p. 403) «March (?) - Septem Huuskonen (TUR), Kilpisjarvi Saana (on Bryum) 18. 9. 196-8 Tauno Ulvinen (OULU). ber« is really doubtful. In the Alps and in Li. U t s j o k i Pulmanki Ruossavaara (on the Tatra the species is found in September Calliergon stramineum) 14. 8. 1963 Yrjii Makinen (HErM & REMY 1932 concerning M . rehmii, (TUR), east side of the lake Kevujarvi (Kevo jar\'i) (on Paludella squarrosa) 21. 8. 1965 P. Kal BENEDIX 1962). lio & E. Kankainen & 0. Skifte (TROMS), south west slope of Kenishpakte (Kenespahta) cliff 18. Distribution. (Fig. 3) The distribution of 8. 19-65 Ola Skifte (TROMS). the different forms of M. gracilis is already treated above. The capitate form is found NORWAY Troms. L y n g s e i d e t Rattenvikfjellet (on to occur from 63 o20'to 70° n.lat. and the Helodium lanatum, Paludella squarrosa) 25 . 7. pileate form from 68° to 78° ri.lat. The 1968 Reino Alava & Kalevi Alho (TUR).

32 Finnmark. Po I m a k paisa bog west of Varan only list the unpublished notes. For the infor gerbotn (on Paludella squarrosa, Drepanocladus) mation as regards the nomenclature of this 12. 8. 1968 Esteri Kankainen (TUR) and 20. 8. 1968 Anja Niskanen (TUR, KIEVO). genus we thank Dr. R. A. Maas Geesteranus Svalbard. V e s t s pi t s berg e n Isfjorden and Phil. lie. Tauno Ulvinen. southwest of Festningen 14. 8. 1966 Paavo Kallio & Esteri Kankainen (TUR), near the Isfjord Ra H eyderia abietis is a species of the autumn. dio Station (on Drepanocladus revolvens, Onco It can be found even during November. It phorus wahlenbergii) 13. 8. 1966 Heli Heikkila is recorded to occur except in the northern (TUR, KEVO) and (on Calliergon sarmentosum) 17. 8. 1966 Paavo Kallio (TUR), Kongressdalen hemisphere also in Australia (IMAI 1941). (on Calliergon sarmentosum) 1·6. 8. 196•6 Esteri List of the specimens studied: Kankainen (TUR, KEVO), souvhwest side of Linnevatnet (on Drepanocladus) 12. 8. 1966 Heli H eyderia abietis var. abietis Heikkila (TUR), west side of Linne,·atnet (on Calliergon sarmentosum) 13 . 8. 1966 Heli Hei'k FINLAND kiHi. (TUR), Vardasen (on Tomentypnum nitens) St. Lappi Lapinkyla 20.10. 1957 Holger 18. 8. 1966 ·Esteri Kankainen (TUR). Saltin (TUR), Simasalo 8. 11. 1954 Holger Sal tin (TUR). SWEDEN Ta. J o k i o in en Minkii:i Nummela (fqq) 22. Torne Lappmark. J u k k as jar v i Abisko (on 10. 1967 Esteri Kankainen (TUR); Y I i:i jar vi Paludella, Drepanocladus?) 5. 6. 1943 Rolf San Makkyla 15. 9. 1968 Esteri Kankainen (TUR), tesson ( S), Kopparil.sen (on Paludella, Drepano Teivaala 15. 9. 1968 Yrji:i Makinen (TUR). cladus) 6. 8. 1943 Rolf Santesson (S). Tb. To iva k k a Huikko Riihimaki (also on pineneedles) 19·. 9·. 1968 Esteri Kankainen (TUR) . USSR Sa. Taipalsaari Karhunpaa 19.9. 1965 Kpoc. P a ate n e (on Philonotis tomentella) Orvo Vitikainen (TUR). 15. 7. 1•8•69 J. Sahlberg (TUR). Ob. K i i mink i Kalkkialue 5. 9·. 1968 Tauno CANADA Ulvinen (OULU). N ewfoundland-Labrador. S chef fer vi II e CANADA are a, soutiheast of Mike Lake 54°40' n.lat., Newfoundland-Labrador. Scheff e r vi II e 66°37' w.long., elevation 165.0 ft. 4. 8. 19·67 Esteri are a, southeast o.f Mike Lake (on rhe needles Kankainen (TUR), southwest shore o.f Attikama of Picea mariana and/or P. glauca) 54°40.' n.lat., gen Lake 54°54' n.lat., ·66°3T w.long. (on Aula 66°37' w.long., elevation 1650 ft. 4. 8. 1967 Este comnium palustre) 5. 8. 1967 ·Esteri Kankainen ri Kankainen ( TUR) . (TUR), northwest shore of Petitsikapau Lake (on Paludella squarrosa) 6. 8. 1967 Paavo Kallio (TUR, KEVO), norvh shore of Beam Lake 54°45' H eyderia abietis var. pusilla (Fr.) Ulv. n.lat., 66°49' w.long., elevation 1700 ft. 6. 8. 1963 FINLAND Paavo Kallio (TUR) . St. K o k em a k i Kuivasaari 2-8. 9·. 1967 Aira Quebec-Labrador. F or t C him o are a, & Kalevi Pohjola (TUR) . Highfall Creek, nort!h of the falls 58°01' n.lat., Ta. Tam me I a Saloisjarvi 15. 10. 19·67 Esteri 68 ° 29' w.long. (on Drepanocladus revolvens) Kankainen ( TUR). 2. 8. 19·67 Esteri Kankainen (TUR), Old Fort Ob. Kiiminki Juuansydanmaa (aisoonspru Chimo 5·8°09, n.lat., 68° 18' w.long. (on Paludella ce needles) 28. 9. 1968/XII Martti Ohenoja (OU squarrosa) 31. 7. 1967 Esteri Kankainen (TUR) . LU); M u h o s Muhospera 14. 9. 1968/XV and Heyde ria Weinmann, Syll. Pl. nov. 2, 14. 9. 19·68/XLII Martti Ohenoja (OULU) . Le. En on t e k i i:i 12 km northeast of Palo p. 108. 1828. joensuu 12. 9. 1967 Tauno Ulvinen (OULU). 11his is the northernmost collection and it is made 3. Heyderia abietis (Fr.) Weinmann, Syll. in !!he area where the spruce is completely lacking. Pl. nov. 2, p. 108. 1826. - Link, Handb. Erkenn. Gewachse 3, p. 311. 1833. As seen above there is hardly a basis for In 1966, ULVINEN wro-te about this species the separation of different taxa. The culti and renewed FRIES' ( 1828) «Mitrula Hey vation of the species in different circum deria Abietis« and M. Heyderia pusilla«. We stances is needed. SUMMARY Mitrula paludosa Fr. is a species of spring forms exist. The basis of the variation is not and early summer. It is common in Finland yet solved. up to the birch forest limit. H eyderia abietis (Fr.) Weinm. grows on Mitrula gracilis Karst. is a subarctic pine and spruce needles, but the separation arctic-alpine species. Its external morphology of var. abietis and var. pusilla is not fully varies very much. Except the extreme capi founded. The species is found in Finland up tate and pileate forms also infraspecific to the limits of the pine and spruce forest.

33 REFIERENGES BENEDIX, E. H ., 1962 : Neues tiber die Geoglossa 1885: Revisio monographica a tque synopsis. ceen: Coelotiella, Mitrula. - Kulturpfl. Ascomycetum in Fennia hucusque detec Beih. 3, 3 · 89~410. torum. - Acta Soc. F. Fl. Fennica II: 6, BRESADOLA, AB. ]., 189•2: Fungi tridentini novi, vel 1-174. nondum delineati, descripti, et iconibus LANGE, M., 1957 : Macromycetes 3. - Medd. illustrati II. - 46 pp., 150 tab. Tridenti. Gmnland 148: 2, 1-125. CooKE, M. C., 1871 : Handbook of British fungi II, LARSEN, P., 19·24: Fungi, in]. Gandrup: A botani 48 · 9~ 9•81. London. cal trip to Jan Mayen. - Dansk Bot. Ark. CoRNER, E.]. H., 19·3·0: Studies in the morpho 4, 23-24. logy of Discomycetes III The Clavuleae. - LLOYD, C. G., 1916 : The Geoglossaceae.- Mycol. Trans. Brit. Myc. Soc. 15, 107- 120. Writings 5, 1- 24. DITTRICH, G., 1898: Zur Entwickelungsgeschichte LUNDELL, S. & ]. A. NANNFELDT, 19·34-59: Fungi der Helvellineen. - Cohn's Beitdige Biol. exsiccati suecici, praesertim upsalienses. - Pfl. 8, 17-52. Uppsala. DURAND, E. ]., 1908: The Geoglossaceae of North MAAS GEESTERANUs, R. A., 19•64 : On some white- America. - Ann. Myc. £, 387- 477. spored Geoglossaceae. Persoonia 2, EcKBLAD, F.-E., 19·63: Contributions to the Geo 81 - 9·6. glossaceae of Norway.- Nytt Mag. Bot. 10, MAINS, E. B., 1955: North American hyaline 137-158. spored species of the Geoglosseae. - Mycol. EISFELDER, I., 1962 : Hohere ·Pilze a us dem Pitztal 47, 846~877. (Tirol). - Ber. Bayer. Bot. Ges. 35, 28- MAssEE, G., 1897: A monograph of the Geoglosseae. 3-8. - Ann. Bot. 11, 225-306. FAVRE, ]., 1949: A propos d'un Mitrula arctico NANNFELDT, ]. A., 19•2.8: Contributions to the alpin (Mitrula gracilis Karst. = M. musci mycoflora of Sweden. 1. Discomycetes from cola E. Henn. = M. Rehmi Bres.).- Schw. Tome Lappmark. - Sv. Bot. Tidskr. 22, Zeitschr. Pilzk. 19'49, 143- 147. 115- 139. 196'5: Les champignons superieurs de Ia 1932: Studien tiber die Morphologie und zone alpine du Pare National suisse. - Er Systematik der nicht-lichenisierten inoper gebn. wiss. Unters. schweiz. Nationalparks 5 culaten Discomyceten. - N. Acta Reg. Soc. (Neue Folge) : 33, 1-212. Sci Upsal. IV 8: 2, 1- 3·68. 1960: Cataloque descriptif des champig 1942: The Geoglossaceae of Sweden. - nons superieurs de Ia zone subalpine du Ark. Bot. 3.0 A: 4, 1- 67. Pare National suisse. - Ibid. 6:42, 323- NAUMOV, H. A. , 19·64: Flora gribov Leningradskoi 610. oblasti II Diskomitseti. - 258 pp. Moskva~ FRIES, 'E., 18'21: Systema mycologicum I. - 520 Leningrad. pp. Lundae. PERSOON, C. H., 1801: Synopsis methodica fungo HEIM, R. & L. REMY, 193•2: Especes rares ou rum. - 708 po. Gottingen. nouvelles de Discomycetes des Alpes Brian REHM, H., 1896: Die Pilze Deutschlands, Oester ~;onnaises. - Soc. Myc. France 48 , 53- 75 . reichs und der Schweiz 3. - 1275 pp. HENNING, E., 18•85: Bidrag till svampfloran i Norges Leipzig. sydligare fjelltrakter.- Ofvers. Kong!. Vet. RosTRUP, E., 1904: Norske Ascomyceter. - Vid. Akad. Forhandl. 5, 49~ 76. Selsk. Skrift. 1. Math.-naturv. Kl. 1904, 4, IMAI, S., 1940: The Geoglossaceae of Norway. - 1- 44. Ann. Myc. 38, 26-8- 278 . SACCARDO , P. A., 1889: Sylloge fungorum 8. - 1941: Geoglossaceae Japoniae. - Joum. Patavii. Fac. Agric. Hokkaido Imp. Univ. 45, 155- SEAVER , F. ]., 19-11: Studies in Colorado fungi I. 264. Discomycetes. - Mycol. 3, 57~6 · 6. 1956: Contributiones ad studia mono SKIRGIELLO, A., 196·1: De quelques champignons graphica Geoglossacearum. III. - Sci. superieurs recoltes par M. Kuc au Spitsberg Reports Yokohama Nat. Univ. II: 5, 1-8. en 1958. - Bull. Res. Counc. Israel 10D, KALLIO, P. & E. KANKAINEN, 1964: Notes on the 287-29·3. macromycetes of Finnish Lapland and SvRCEK, M., 1962: Diskomycety z Nizkyoh Tater, adjacent Finnmark. - Ann. Univ. Turku nalezene behem posjezdove exkurze II. A II: 32 (Rep. Kevo Subarctic Sta. 1), SEM. 1960. - Ces. Mykol. 16, 87-114. 178- 235 . ULVINEN , T., 1963 : Beobachtungen tiber die Pilze 19•66: Additions to the mycoflora of north on Savukoski, Nord£innland. - Aquila 1, enrmost Finnish Lapland. - Ibid. A IT: 3·6 53-58. (Rep. Kevo Subarctic Sta. 3), 177- 210. 1966 : Uber die auf Koniferennadeln !eben KANKAINEN, E. & P. KARLSTROM & H . HEIKKILA , den Arten der Gattung Mitrula s. lat. (Asco 19•67: Kevolaisten retki Huippuvuorille mycetes, Geoglossaceae). - Ann. Bot. Fenn. v. 1966. - Luonnon Tutkija 71, 118- 123. 3, 433- 438. KARSTEN, P. A., 1871: Myco.Jogia Fennica 1. Pars WEINMANN , ]. A., 1828 : Enumeratio fungorum in prima Discomycetes. Bidr. Kanned. agm Pawlowskiensi praecipue crescentium. Fin!. Natur och Folk 19·, 1- 264. - Syll. Pl. nov. 2, 82- 118. 1883·: Fragmenta mycologica. - Hedwigia VELENOVSKY, ] ., 1934: Monographia Discomycetum 22, 17-18. Bohemiae 1-2. - 436 pp, 31 tab. Pragae.