Two New Species of Dactylospora (Dactylosporaceae, ), with a Key to the Known Species in Scandinavia Author(s): Per G. Ihlen, Håkon Holien, Tor Tønsberg Source: The Bryologist, 107(3):357-362. 2004. Published By: The American Bryological and Lichenological Society, Inc. DOI: 10.1639/0007-2745(2004)107[0357:TNSODD]2.0.CO;2 URL: http://www.bioone.org/doi/full/10.1639/0007- 2745%282004%29107%5B0357%3ATNSODD%5D2.0.CO%3B2

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BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. The Bryologist 107(3), pp. 357 362 Copyright ᭧ 2004 by the American Bryological and Lichenological Society, Inc.

Two New Species of Dactylospora (Dactylosporaceae, Lecanorales), with a Key to the Known Species in Scandinavia

PER G. IHLEN Department of Ecology and Environmental Science, UmeaÊ University, SE-901 87 UmeaÊ, Sweden; e-mail: [email protected]

HAÊ KON HOLIEN Faculty of Social Sciences and Natural Resources, Nord-Trùndelag University College, Serviceboks 2501, N-7729 Steinkjer, Norway; e-mail: [email protected]

TOR TùNSBERG Museum of Botany, University of Bergen, AlleÂgaten 41, N-5007 Bergen, Norway; e-mail: [email protected]

Abstract. Dactylospora aeruginosa Holien & Ihlen, sp. nov. is described from Norway and U.S.A. and D. borealis Holien & Ihlen, sp. nov. is described from Norway, Sweden, and U.S.A. Both species are lichenicolous, but D. aeruginosa is also saprobic on wood and bark of Picea abies and Juniperus communis, whereas D. borealis is only rarely growing on bark of P. abies. Notes are given on their ecology and distribution. A key to the known lichenicolous species of Dactylospora occurring in Scandinavia is included. Lichenodiplis lecanorae is suggested to be the anamorph of D. borealis.

Keywords. Dactylospora, lichenized ascomycetes, Lichenodiplis, lichenicolous fungi, Trùndelag phytogeographic element.

During ®eldwork carried out by the authors in heimleri (Zukal) DoÈbb. & Triebel on several liver- Norway (HH, TT), Sweden (PGI), and the Paci®c worts (DoÈbbeler & Triebel 1985), and D. stygia var. northwest of U.S.A. (TT), two species of Dactylos- stygia on various deciduous trees, e.g., Acer, Alnus, pora KoÈrb. (Dactylosporaceae, Lecanorales) were Carpinus, Carya, Frangula, Fraxinus, Lirioden- discovered. One contained hyaline to light brown, dron, Populus, Quercus, and Salix (see Hafellner 1-septate ascospores, and a blue-violet color of the 1979), are not included in the key. exciple and epihymenium reacting Kϩ aeruginose, and was found lichenicolous on several and saprobic on Picea abies and Juniperus communis. MATERIAL AND METHODS The other, with 3-septate ascospores and an orange brown epihymenium reacting Kϩ purple, was also In addition to the specimens of Dactylospora aerugi- found lichenicolous and once directly on bark of nosa and D. borealis collected in the ®eld, several spec- Picea abies. As these two species do not agree with imens were found by examining herbarium specimens (in any of the species included in the major taxonomic BG and TRH) of their hosts. For anatomical studies, a Zeiss treatments of Dactylospora (e.g., Hafellner 1979; light microscope with magni®cations of ϫ400, ϫ630, and ϫ1,000, equipped with a blue ®lter, was used. Microscop- Triebel 1989) or, to our knowledge, in any other ic examinations required sections, mostly 15±20 ␮m thick, paper dealing with Dactylospora, they are de- cut with a freezing microtome. The estimated values of scribed as new below. the sizes of the ascospores have been given as: (min.±) On a worldwide scale, Dactylospora contains arithmetic meanϪ1.0 SD±arithmetic mean ϩ 1.0 SD (± about 37 species (Kirk et al. 2001; Lawrey & Died- max.) where min. and max. are the extreme values and SD the corresponding standard deviation of the sample. erich 2003; SarrioÂn et al. 2002). As 20 lichenicol- All measurements were done in water, except for the sizes ous species, including the two species described of the ascospores and the thickness of the paraphyses that here, are presently known from Scandinavia (Al- were done in K (i.e., a 10% solution of potassium hy- strup 1991; Santesson et al. 2004), we found it ap- droxide). The measurements in K were done in order to propriate to include a key to these species. It should observe details in these structures more clearly, e.g., or- namentation of the ascospores and the branching of the be noted that non-lichenicolous taxa such as Dac- paraphyses. Each species keyed out has been followed by tylospora epimyces (Tobisch) Hafellner on the fun- one selected literature reference for further information on gus Tubulicrinis glebulosus (Hafellner 1979), D. morphology, distribution, and ecology.

0007-2745/04/$0.75/0 358 THE BRYOLOGIST [VOL. 107

FIGURE 1. Dactylospora aeruginosa (holotype). Ð A. Section of ascoma (in water) with patches of pigments in the epihymenium. Ð B. Paraphyses (in LCB). Scales: A ϭ 23.0 ␮m; B ϭ 5.0 ␮m.

DACTYLOSPORA AERUGINOSA Holien & Ihlen, sp. padium disciforme (Flot.) Kullh., and Micarea pra- nov. FIG.1 sina s. lat. and saprobic on twigs, wood, and bark lichenicolus et sapro®ticus. Excipulum et epihy- of Picea abies and twice on partly decayed wood menium violaceum, K plus aeruginosum. Ascosporae, of Juniperus communis. When growing lichenicol- 1-septatae, hyalinae ad dilutus brunnescens, (9.0±) ous, the hosts were found lignicolous and corticol- 11.0±14.5(±16.0) ϫ (3.0±)3.5±5.5(±7.0) ␮m. ous on twigs and trunks of Picea abies (9 speci- TYPE: NORWAY. SùR-TRùNDELAG. Orkdal, E of Svorkmo, mens), and corticolous on Alnus incana (4), Betula N-facing slope of SprangaÊsen, UTMED: NR 40 03±04, elevation ca 260±270 m, saprobic on Juniperus com- sp. (1), Juniperus communis (5), Prunus padus (1), munis in old forest, 9 June 1993, Holien 5622 (TRH, Salix caprea (1), and Sorbus aucuparia (2). Many holotype; UPS, isotype). specimens were from old Picea forests. Closely as- Apothecia (Fig. 1A) rounded, 0.3±0.7 mm in di- sociated taxa (found on the same small piece ameter; disc ¯at, brown; margin distinct, concolo- of substrate) observed in the Scandinavian material rous or darker than disc; upper part of exciple in included for example Biatora toensbergii Holien & section (15±)30±80 ␮m thick, inner part light Printzen, Catinaria atropurpurea (Schaer.) VeÏzda brown, KϪ, outer part violet-blue, Kϩ aeruginose, & Poelt, Cavernularia hultenii Degel., Lecanora NϪ, exciple in section in lower part (8.0±)15±50 farinaria Borrer, Lobaria scrobiculata (Scop.) D. ␮m thick, cells polygonal, violet-blue, Kϩ aerugi- C., Micarea cinerea (Schaer.) Hedl., nose, NϪ. Hymenium (Fig. 1A) hyaline to light or- juvenalis Brodo, Parmeliella parvula P. M. Jùrg., ange brown, KϪ, with patches of violet-blue, Kϩ and Pseudocyphellaria crocata (L.) Vain. In aeruginose, (50±)70±120(±140) ␮m high; epihy- U.S.A., D. aeruginosa was collected on thalli of menium hyaline to light brown KϪ, with densely Biatora ef¯orescens and once on Japewia subaur- arranged patches of violet-blue, Kϩ aeruginose. ifera growing corticolous on Alnus crispa ssp. sin- Hypothecium light brown, KϪ. Paraphyses (Fig. uata. Associated lichen species included Lopadium 1B) slightly branched, 1.5±2.0 ␮m thick, usually disciforme, Micarea cinerea, Ochrolechia sp., and with swollen and pigmented apical caps or some- Ramalina roesleri (Schaer.) Hue. times with indistinct apical caps. Asci with a K/Iϩ Distribution. Dactylospora aeruginosa is so blue outer layer, 8-spored. Ascospores hyaline to far known from the provinces Hordaland, Mùre og light brown, ellipsoid, smooth, 1-septate (9.0±) Romsdal, Sùr-Trùndelag, Nord-Trùndelag, and 11.0±14.5(±16.0) ϫ (3.0±)3.5±5.5(±7.0) ␮m(n ϭ Nordland in Norway and from Alaska in U.S.A. 20); perispore present, 1.5±2.0 ␮m thick. Its elevation ranged in Scandinavia from about sea Ecology. In Scandinavia, Dactylospora aeru- level to 270 m, and in Alaska from sea level to ginosa has been collected on thalli of different crus- 20±39 m. tose lichens, such as Biatora ef¯orescens (Hedl.) Discussion. Dactylospora aeruginosa is dis- Erichsen, Japewia tornoeÈnsis (Nyl.) Tùnsberg, Lo- tinct by the 1-septate, hyaline to light brown as- 2004] IHLEN ET AL.: DACTYLOSPORA 359 cospores, with a thin perispore; 8-spored asci; and DACTYLOSPORA BOREALIS Holien & Ihlen sp. nov. the violet-blue color of the exciple, also present as FIG.2 patches in the hymenium and epihymenium, turn- Fungus lichenicolus in Mycoblasto af®ni et M. sanguina- ing Kϩ aeruginose. Several Dactylospora species rio. Epihymenium brunnescens, in K purpurascens. As- contain 1-septate ascospores produced in 8-spored cosporae 3-septatae, maturae verrucosae. asci (see key). Two of these, D. athallina (MuÈll. TYPE: NORWAY. NORD-TRùNDELAG. Overhalla, W of Arg.) Hafellner and D. purpurascens Triebel, might Foss, 64Њ28.8Ј N, 11Њ59.7Ј E, UTMED: UM 555 540, resemble D. aeruginosa, but they can be separated elevation ca 80 m, corticolous on Picea abies in old forest; host: af®nis, 13 March 1997, Hol- from that species by their brown epihymenium re- ien 7092 (TRH, holotype; UPS, isotype). acting Kϩ purple and the distinctly brown asco- spores (see e.g., Ihlen 1998; Triebel 1989). Other Apothecia (Fig. 2A, B) rounded, 0.2±0.8(±1.0) Dactylospora species containing a violet-blue color mm in diameter; disc ¯at, dark brown; margin dis- in the exciple and/or the epihymenium are D. de- tinct, concolorous with disc; upper part of exciple minuta (Th. Fr.) Triebel and D. urceolata (Th. Fr.) in section 20±40 ␮m thick, brown, KϪ,NϪ, some- Arnold. However, D. deminuta is easily separated times greenish brown in inner part; cells in lower from D. aeruginosa by its 3-septate and even sub- part of exciple in section polygonal, 5±15 ␮min muriform ascospores (Triebel 1989), whereas D. diameter, brown, KϪ. Hymenium (Fig. 2B) hyaline urceolata has long (15±23 ␮m), up to 7-septate as- to light orange brown, (40±)70±110(±130) ␮m cospores (Foucard 2001). high, orange pigment Kϩ purple; epihymenium Although found on several lichen species as well brown to orange brown, Kϩ purple, NϪ. Hypoth- as being found as a saprobic, Dactylospora aeru- ecium green, then Kϩ brown, or greenish brown to ginosa seems in Scandinavia to be predominantly brown or orange, then Kϩ purple. Paraphyses (Fig. restricted to the humid spruce forests of central 2C) branched in upper part, 1.5±2.0 ␮m thick, with Norway where it occurs on Picea abies as well as swollen and brown apical caps. Asci with a K/Iϩ on deciduous trees mixed in the forest. Due to its blue outer layer, 8-spored. Ascospores (Fig. 2D) presence among several oceanic lichen species e.g., brown, ellipsoid, ornamentation granulate at matu- Cavernularia hultenii, Parmeliella parvula, and rity, 3-septate, rarely 4-septate or submuriform, Pseudocyphellaria crocata, it may prove to be a (15.0±)15.5±19.5(±23.0) ϫ (4.5±)6.0±7.5(±9.0) ␮m further member of the ``Trùndelag phytogeographic (n ϭ 20); perispore present. element'' (Holien & Tùnsberg 1996). The occur- Ecology. Dactylospora borealis has been found frequently in Scandinavia on thalli, rarely on apo- rence of D. aeruginosa in the humid forests of thecia, of Mycoblastus af®nis (Schaer.) T. Schauer coastal Alaska supports this view. and M. sanguinarius (L.) Norman, and rarely on Japewia tornoeÈnsis, Loxospora elatina (Ach.) A. Paratypes. NORWAY. HORDALAND. LindaÊs, Luro (Ly- gra), NW of Utluro, near hill 42, 1995, Tùnsberg 21755b Massal., and Ochrolechia androgyna (Hoffm.) Ar- (BG). SùR-TRùNDELAG. Orkdal, E of Svorkmo, N-facing nold. In one collection (Nord-Trùndelag, Grong, slope of SprangaÊsen, 1992, Holien 5496 (TRH); 1993, Hol- Holien 6869, TRH), D. borealis occurred directly on ien 6062 (TRH); SprangaÊsen, 1992, Holien 5489 (TRH); bark of Picea abies adjacent to a thallus of Biatora Holien 5497 (TRH); NE-facing slope of HaugaÊsen, 1993, ef¯orescens infected by D. aeruginosa! In Alaska, Holien 5886a (TRH). NORD-TRùNDELAG. Namsos, Sñvik, it occurred on Mycoblastus af®nis and M. sangui- by river Barstadelva, 1993, Holien 6075 (TRH). Overhalla, W of Foss, 1997, Holien 7098 (TRH). Steinkjer, S of Tùnne narius. The hosts were found on twigs, wood, and by VaÊsetbukta, 1997, Holien 7293 (TRH). bark of Picea abies, and, more rarely, corticolous Specimens examined labelled Biatora ef¯orescens. on Sorbus aucuparia and Ulmus glabra. In Alaska NORWAY. HORDALAND. Voss, Flatlandsmoen, 1987, it occurred on Betula and Picea sitchensis. Dacty- Tùnsberg 10128 (BG). MùRE OG ROMSDAL. Aure, Vinster- lospora borealis has mainly been found in old, nes, 1983, Tùnsberg 8279a (BG). SùR-TRùNDELAG. Trond- heim, Byneset, just SE of Hangran, 1980, Tùnsberg 5237 shaded Picea forests, frequently close to rivers or (BG); Hùgstein, 1980, Tùnsberg 5191 (TRH), 5196 (BG), streams. Closely associated lichen taxa (found on 5198 (BG). AÊ fjord, TùrresengaÊsen, along river Norddal- the same small piece of substrate) observed in the selva, 1988, Holien 3167 (TRH). NORD-TRùNDELAG. Fla- Scandinavian material were Biatora ef¯orescens, tanger, Gaupdalen, 1990, Holien 3731 (TRH); Mt. Bryoria capillaria (Ach.) Brodo & D. Hawksw., Rùythaugfjellet, 1981, Tùnsberg 5494 (BG). Grong, W of river Gartlandelva, 1995, Holien 6869 (TRH). Namsos, 0.5 Lecanora circumborealis Brodo & Vitik., Loxos- km E of Kjerstivik, 1980, Tùnsberg 4913a (BG). Nñrùy, pora elatina, Nephroma parile (Ach.) Ach., and along river AÊ vikelva, 1980, Tùnsberg 4657, 4665 (BG). Ochroelchia microstictoides RaÈs. In Alaska Myco- NORDLAND. Brùnnùy, Tilrem, 1985, Tùnsberg 9226 (BG); blastus sanguinarius was a close associate. Hemnes, Hemnesberget, towards Geitvika, 1984, évstedal Distribution. Dactylospora borealis is so far (BG). U.S.A. ALASKA. Kodiak Island Borough, Near Is- land, 2001, Tùnsberg 29482, 29483, 29491, 29492, 29493 known from the provinces Sùr-Trùndelag, Nord- (BG). A total of 29 specimens seen. Trùndelag, and Nordland in Norway and JaÈmtland 360 THE BRYOLOGIST [VOL. 107

FIGURE 2. A. Mycoblastus af®nis (large ascomata), Dactylospora borealis (arrow), and Lichenodiplis lecanorae (dark area on the host thallus) (Ihlen 1443). Ð B±D. Dactylospora borealis (holotype). Ð B. Section of ascoma. Ð C. Paraphyses with pigmented end cells. Ð D. Ascospores. Scales: A ϭ 1.0 mm, B ϭ 18.0 ␮m, C and D ϭ 5.0 ␮m. and AÊ sele Lappmark in Sweden, at elevations rang- 3.5±5.0 ␮m (Hafellner 1979). Dactylospora pseu- ing from 80 m in Overhalla (Norway) to 600 m in dourceolata SarrioÂn & Hafellner (see SarrioÂn et al. Krokom (Sweden). In U.S.A., it has been found in 2002), might because of the reddish brown epihy- the coastal lowlands of SE Alaska. menium and ornamented ascospores, also resemble Note. In eight out of the 28 specimens of Dac- D. borealis. However, in that species the epihy- tylospora borealis examined, the pycnidia of Lich- menium and hypothecium are KϪ, the apothecia enodiplis lecanorae (Vouaux) Dyko & D. Hawksw. are slightly lichenized, and it is saprobic. Dactylos- (Hawksworth & Dyko 1979) were observed on the pora crassa SarrioÂn & Hafellner (see SarrioÂn et al. same discolored spots as, and close to, the ascomata 2002) is also similar to D. borealis, but it is distinct of D. borealis (Fig. 2A). in having smooth and narrower ascospores (3.5±4.5 Discussion. Dactylospora attendenda (Nyl.) ␮m). Dactylospora rostrupii Alstrup, described Arnold, D. borealis, and D. parasitica (FloÈrke) from the Faroe Islands on Pertusaria dactylina Zopf, all occurring in Scandinavia, can easily be (Ach.) Nyl., can be separated from D. borealis by recognized by having predominantly 3-septate as- its smooth and larger ascospores, measuring 19±22 cospores. However, D. borealis is distinct from D. ϫ 5.0±8.0 ␮m, and a brown epihymenium (Alstrup attendenda and D. parasitica by the brown to or- et al. 1994). ange brown epihymenium reacting Kϩ purple (in Dactylospora borealis appears to be a species of D. attendenda it is brown and KϪ or Kϩ dull humid Picea-dominated forests, and like D. aeru- brown). An examination of the isotype of D. par- ginosa, it may prove to be a further member of the asitica revealed that it has a dark brown epihymen- ``Trùndelag phytogeographic element'' (Holien & ium reacting KϪ. Furthermore, D. borealis has Tùnsberg 1996). The presence of D. borealis in the larger ascospores, in D. attendenda they are (7.0± humid forests of coastal Alaska supports this view. )9.5±12.5(13.0) ϫ (3.0±)4.0±6.5(8.5) ␮m (Ihlen However, D. borealis is probably wider distributed 1998) and in D. parasitica they are 9.0±15.0 ϫ than D. aeruginosa, as it has also been found in the 2004] IHLEN ET AL.: DACTYLOSPORA 361 more eastern and drier provinces of Scandinavia 2. Ascospores 1-septate, sometimes 3-septate, 7±11 (JaÈmtland and AÊ sele Lappmark). ϫ 4±6 ␮m. On Pertusaria aspergilla ------D. pertusariicola (Tuck.) Hafellner 1979 The observation of the pycnidia of Lichenodiplis 3. Ascospores always 1-septate ------4 lecanorae on the same discolored spots as, and 3. At least some ascospores more than 1-septate or close to, the ascomata of Dactylospora borealis submuriform ------14 (Fig. 2A), indicates an anamorph-teleomorph rela- 4. Ascospores hyaline to light brown, epihymenium tionship. Consequently, we propose that L. leca- with patches of violet-blue pigment turning Kϩ norae is the anamorph of the teleomorph Dactylos- aeruginose ------D. aeruginosa Holien & Ihlen pora borealis. 4. Ascospores distinctly brown, epihymenium with- out patches of violet-blue pigment, KϪ or Kϩ purple ------5 Paratypes. NORWAY. SùR-TRùNDELAG. Bjugn, S-fac- 5. Epihymenium brown, Kϩ purple 6 ing slope by end of lake Skjervatnet, Snùholet, 1994, Hol------5. Epihymenium brown, not Kϩ purple 7 ien 6385 (TRH). Meldal, Urvatnet forest reserve, 1991, ------6. Holien 4743 (TRH). Orkdal, SE of Svorkmo, NE-facing Ascospores 9±13 ϫ 4±7 ␮m, hymenium 55±65 slope of HaugaÊsen, 1993, Holien 5887 (TRH); E of Svork- ␮m high. On Baeomyces rufus ------mo, N-facing slope of SprangaÊsen, 1993, Holien 6063 ------D. athallina (MuÈll.Arg.) Hafellner 1979 (TRH). Rissa, Stadsbygd, on the southern side of lake Vat- 6. Ascospores 12±19 ϫ 4±8 ␮m, hymenium 55±90 nagaÊrdsvatn, 1961, Santesson 14299b (UPS). NORD- ␮m high. On Amygdalaria, dovrensis, TRùNDELAG. Overhalla, W of Foss, 1995, Holien 6973 and Porpidia glaucophaea ------(TRH); 1997, Holien 7645 (TRH). Steinkjer, Strukstadmyra, ------D. purpurascens Triebel 1989 2004, Holien 9862 (TRH). SWEDEN. JAÈ MTLAND.AÊ re, west 7. On thalli of foliose lichens (Lobaria, Pannaria,or È side of lake KallsjoÈn, by river Oster-Kjolan, 2003, Ihlen Parmeliella) ------8 1443 (UPS); near waterfall TaÈnnforsen, 2003, Ihlen 1447 7. On thalli and ascomata of crustose lichens ------9 (UPS). Krokom, near lake LaÊngtjaÈrn, NW of place Gra- 8. Excipulum reddish brown, ascospores with gran- UPS vbraÈnna, 10 km S of FoÈllinge, 2003, Ihlen 1450 ( ), ular ornamentation. On Lobaria pulmonaria ------1451 (UPS); between place Lakavattnet and Mt SkakafjaÈl------D. lobariella (Nyl.) Hafellner 1979 Ê let, 2003, Ihlen 1455 (UPS). ASELE LAPPMARK. BorgafjaÈll, 8. Excipulum brown, ascospores smooth. On Par- LillfjaÈllet, 2004, Hagner (UPS); Vilhelmina, Borkan, vid meliella and Fuscopannaria praetermissa ------nordvaÈstra sidan av BorkasjoÈn, 2004, Hagner (UPS). ------D. protothallina (Anzi) Hafellner 1979 U.S.A. ALASKA. Haines Borough, NW of Haines, Chilkat Peninsula, along and E of main road to Chilkat State Park, 9. Apothecia without stalk. Hypothecium not black at Mount Riley Trail trailhead, 2003, Tùnsberg 33212 ------10 (BG). 9. Apothecia slightly stalked (ca 50 ␮m high). Hy- Specimens examined labelled Biatora ef¯orescens. pothecium dark brown to almost black. On Amyg- NORWAY. NORD-TRùNDELAG. Grong, W of river Gartlan- dalaria panaeola, Lecanora, and Pleopsidium delva, 1995, Holien 6869 (TRH). chlorophanum ------Specimens examined labelled Loxospora elatina. ------D. rimulicola (MuÈll. Arg.) Hafellner 1979 NORWAY. SùR-TRùNDELAG. Malvik, Hùybydalen, SlaÊtt- 10. Hypothecium hyaline to light brown or olivae- myrdalen, 1987, Holien 2620 (TRH). Trondheim, Bymarka, cous green ------11 E-facing slope by lake Skjellbreia, 2000, Holien 8473 10. Hypothecium medium to dark brown, reddish (TRH). brown or almost black 12 Specimens examined labelled Mycoblastus af®nis or M. ------11. Ascospores Ͼ5±6 ␮m broad, on Pertusaria and sanguinarius. NORWAY. SùR-TRùNDELAG. Bjugn, Ochrolechia ------Stjùrna, Fisdal, Brandhaug, 1952, Rui (M. sanguinarius; ------D. saxatilis (Schaer.) Hafellner 1979 TRH). Klñbu, Moan, 1979, James & Tùnsberg (M. af®nis; BG). Rissa, E of Osavatnet, 1980, Tùnsberg 4615 (M. san- 11. Ascospores Ͻ5±6 ␮m broad, on Lecanora, Pro- guinarius; BG). Trondheim, Bymarka, Geitkind, 1936, toparmeliopsis muralis, and Tephromela atra --- Hùeg (M. sanguinarius; TRH). NORD-TRùNDELAG. Leksvik, ------D. homoclinella (Nyl.) Hafellner 1979 S slope of Tinghaugen, 1981, Tùnsberg 5866 (M. sangui- 12. Ascospores usually 1-septate, sometimes 3-sep- narius; BG). Namdalseid, S of Altvatn, 1981, Tùnsberg tate, hyaline to brown, 6±8 ␮m broad. On Bry- 5434a (M. sanguinarius; BG). Namsos, Almdalen, 1981, odina rhypariza ------Tùnsberg 5581 (M. af®nis; BG). NORDLAND. Grane, NE of ------D. rhyparizae Arnold, see Keissler (1930)

Trofors, E-facing slope S of lake Almvatnet, 2001, Holien 12. Ascospores 1-septate, brown, 3±6 ␮m broad ----- 13 8784 (M. af®nis; TRH). U.S.A. ALASKA. Kodiak Island 13. Ascospores 9±13 ϫ 4±6 ␮m. On Porpidia con- Borough, Kodiak Island E, Middle Bay, 1991, Tùnsberg traponenda and P. macrocarpa ------15343 (M. af®nis; BG). A total of 28 specimens seen. -----D. australis Triebel & Hertel, see Triebel (1989) Exsiccate specimen of Dactylospora parasitica used for 13. comparison. Floerke, Deutsche Lichenen, 101 (UPS!, Ascospores 7±11 ϫ 3±6 ␮m, On Amygdalaria lectotype). ------D. amygdalariae Triebel 1989 14. Ascospores up to 3-septate ------15

14. Ascospores 3 to 7-septate or submuriform ------19 KEY TO THE LICHENICOLOUS SPECIES OF 15. Hypothecium medium brown to reddish brown, DACTYLOSPORA IN SCANDINAVIA orange, or greenish brown ------16

1. Asci multi-spored ------2 15. Hypothecium light brown. On Ochrolechia par- 1. Asci 8-spored ------3 ella ------2. Ascospores 1-septate, 4±5(±7) ϫ 2±3 ␮m. On Me- D. parellaria (Nyl.) Arnold, see Clauzade et al. galaria pulverea ------D. microspora Etayo 1991 (1989) 362 THE BRYOLOGIST [VOL. 107

16. Epihymenium orange brown, Kϩ purple. On CLAUZADE, G., P. DIEDERICH &C.ROUX. 1989. Nelikeni- Mycoblastus ------D. borealis Holien & Ihlen gintaj fungoj likenlogaj. Bulletin de la SocieÂte lin- 16. Epihymenium light to dark brown or reddish neÂenne de Provence Numero Special 1: 1±142. brown, KϪ,Kϩ dull brown, or Kϩ greenish DOÈ BBELER,P.&D.TRIEBEL. 1985. Hepaticole Vertreter brown ------17 der Gattungen Muellerella und Dactylospora (Asco- 17. Ascospores brown, sometimes hyaline, 5±7 ␮m mycetes). Botanische JahrbuÈcher fuÈr Systematik 107: broad. On Bryodina rhypariza ------503±519. ------D. rhyparizae Arnold, see Keissler (1930) ETAYO, J. 1991. Dactylospora microspora spec. nov., nue- 17. Ascospores brown, never hyaline, Ͻ5±6 ␮m vo hongo liquenicola de la ¯ora espanÄola. Candollea broad ------18 46: 391±393. 18. Epihymenium brown to reddish brown. On FOUCARD, T. 2001. Svenska skorplavar och svampar som Ochhrolechia and Pertusaria ------vaÈxer paÊ dem. Interpublishing, Stockholm. --- D. parasitica (FloÈrke) Zopf, see Hafellner (1979) HAFELLNER, J. 1979. Karschia. Revision einer Sammel- 18. Epihymenium light brown to dark brown. On gattungen der Grenze von lichenisierten und nichtli- Amygdalaria consentiens, A. pelobotryon, Ic- chenisierten Ascomyceten. Beiheft zur Nova Hedwigia madophila, Pilophorus cereolus, P. dovrensis, 62: 1±248. Porpidia melinodes, and P. ochrolemma ------ÐÐÐ. 1985. Studien uÈber lichenicole Pilze und Flechten ------D. attendenda (Nyl.) Arnold, see Triebel (1989) IV. Die auf Brigantiaea-Arten beobachteten Ascomy- 19. Apices of paraphyses 3±5 ␮m broad. Ascospores ceten. Herzogia 7: 163±180. 5 to 7-septate to submuriform, on Brigantiaea HAWKSWORTH, D. L. & B. J. DYKO. 1979. Lichenodiplis fuscolutea and Protothelenella sphinctrioides ---- 20 and Vouauxiomyces: two new genera of lichenicolous 19. Apices of paraphyses 2±3 ␮m broad. Ascospores coelomycetes. Lichenologist 11: 51±61. 3 to 7-septate to submuriform, on various lichens HOLIEN,H.&T.TùNSBERG. 1996. Boreal regnskog i Nor------D. deminuta (Th.Fr.) Triebel 1989 geÐhabitatet for trùndelagselementets lavarter. Blyttia 20. Epihymenium brown, often with a violet tinge, 54: 157±177. ascospores up to 7-septate. On Protothelenella IHLEN, P. G. 1998. The lichenicolous fungi on species of sphinctrioides ------the genera Baeomyces, Dibaeis, and Icmadophila in ----- D. urceolata (Th.Fr.) Arnold, see Triebel (1989) Norway. Lichenologist 30: 27±57. 20. Epihymenium greenish brown, ascospores 7-sep- KEISSLER,K.VON. 1930. Die Flechtenparasiten, pp. 1±712. tate to submuriform. On Brigantiaea fuscolutea In L. Rabenhorst (ed.), Kryptogamen-¯ora von ------D. frigida Hafellner 1985 Deutschland, OÈ sterreich und der Schweiz, Band 8. Akademische Verlagsgesellschaft, Leipzig. KIRK, P. M . , P. F. C ANNON,J.C.DAVID &J.A.STALPERS. ACKNOWLEDGMENTS 2001. Ainsworth & Bisbys Dictionary of the Fungi. Per G. Ihlen acknowledges the ®nancial support pro- CAB International, Oxon. vided by the program The Swedish Initiative LAWREY,J.D.&P.DIEDERICH. 2003. Lichenicolous fungi: (Svenska artprojektet) funded by the Swedish Species In- Interactions, evolution, and biodiversity. THE BRYOL- formation Centre (ArtDatabanken). We thank Mats Wedin OGIST 106: 80±120. (UmeaÊ), Rolf Santesson (Uppsala), and Thorsten Lumbach SANTESSON, R., R. MOBERG,A.NORDIN,T.TùNSBERG & (Chicago) for valuable comments on the manuscript, An- O. VITIKAINEN. 2004. Lichenforming and Lichenicol- ders Nordin (Uppsala) for help with the Latin texts, and ous Fungi of Fennoscandia. Museum of Evolution, AÊ sa Hagner (UmeaÊ) for placing two specimens of Dac- Uppsala University, Uppsala. tylospora borealis at our disposal. Tor Tùnsberg has re- SARRIO N, F. J., J. HAFELLNER &A.R.BURGAZ. 2002. Three ceived travel grants from the Grolle Olsens Fund. new species of the Dactylospora in Spain. Li- chenologist 34: 361±368. TRIEBEL, D. 1989. Lecideicole Ascomyceten. Eine Revi- LITERATURE CITED sion der obligat lichenicolen Ascomyceten auf leci- ALSTRUP, V. 1991. Lichens and lichenicolous fungi from deoiden Flechten. Bibliotheca Lichenologica 35: 1± the TornetraÈsk area. Graphis Scripta 3: 54±67. 278. ÐÐÐ, S. N. CHRISTENSEN,E.S.HANSEN &S.SVANE. 1994. The lichens of the Faroes. FroÂdskaparrit 40: 61± 121. ms. received March 24, 2004; accepted April 30, 2004.