Mycologia ISSN: 0027-5514 (Print) 1557-2536 (Online) Journal homepage: http://www.tandfonline.com/loi/umyc20 Ascospore diversity of bryophilous Hypocreales and two new hepaticolous Nectria species Peter Döbbeler To cite this article: Peter Döbbeler (2005) Ascospore diversity of bryophilous Hypocreales and two new hepaticolous Nectria species, Mycologia, 97:4, 924-934 To link to this article: http://dx.doi.org/10.1080/15572536.2006.11832784 Published online: 27 Jan 2017. Submit your article to this journal View related articles Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=umyc20 Download by: [University of Newcastle, Australia] Date: 28 March 2017, At: 09:31 Mycologia, 97(4), 2005, pp. 924±934. q 2005 by The Mycological Society of America, Lawrence, KS 66044-8897 Ascospore diversity of bryophilous Hypocreales and two new hepaticolous Nectria species Peter DoÈbbeler1 tinct microniches on their hosts, such as ventral leaf Ludwig-Maximilians-UniversitaÈt MuÈnchen, FakultaÈt borders or perianths in Jungermanniales or adaxial fuÈr Biologie, Systematische Botanik und Mykologie, surfaces of leaves in Polytrichales. Several species reg- Menzinger Straûe 67, D-80638 MuÈnchen, Germany ularly develop ascomata on the ventral side of foliose hepatics and perforate host leaves. Ascomata are glo- bose or pyriform, setose or nonsetose, nonstromatic Abstract: Hypocreales represents one of the most perithecia of varying size, and color varies from successful orders of ascomycetes on mosses and he- orange-red to hyaline. Excipulum structures repre- patics, and more than 30 obligately bryophilous spe- sent a variety of tissue types. Hyphae offer additional cies belonging to seven genera of Bionectriaceae and important diagnostic features. They grow super®cial- Nectriaceae are known. These fungi have a remark- ly over host cells, or between or within them, some- ably wide range of ascospore types that differ in form, times forming appresoria, haustoria or conidiogen- size and septation. Especially heterogeneous are the ous cells. The ecological relationships among fungal ascospores of the six hypocrealean parasites recorded species and hosts range from biotrophic (with supra- on European populations of Frullania dilatata. Pat- or intercelluar mycelium) to necrotrophic parasitism terns of distribution of bryophilous Hypocreales on (with intracellular mycelium). At least one species this host appear to follow the principles of island bio- causes malformations of infected host organs. Speci- geography. Two new leaf-perforating species of Nectria ®city to certain host species, genera or groups of host with unusual ascospores are described. Nectria foertheri genera is pronounced in most species. Their diversity is highlighted by an array of ascospore features rep- DoÈbbeler sp. nov. grows biotrophically on Porella sp. in Guatemala, and N. lagodes DoÈbbeler sp. nov. is a resenting many types that occur within bryophilous necrotroph on Frullania dilatata in Italy and Greece. ascomycetes in general (DoÈbbeler 1997, FIG.1)and Key words: ascospore morphology, bryophilous within the numerous nonbryophilous Hypocreales. ascomycetes, Frullania dilatata, Nectria foertheri, Nec- In other cases the bryophilous Hypocreales have tria lagodes unique ascospores not recorded elsewhere within nectrioid fungi. In this paper ascospore types of bryophilous spe- INTRODUCTION cies of Nectria and related genera are described and illustrated and an attempt is made to explain patterns More than 30 species in seven genera of Bionectri- in ascospore diversity by drawing correlations be- aceae and Nectriaceae (Hypocreales, Ascomycetes) tween ascospore types and ecological aspects. In ad- are recorded on mosses and hepatics and nearly all dition two new hepaticolous species of Nectria with are obligate parasites unknown from other substrates. unusual ascospores are proposed. Hypocrealean fungi comprise the most successful fungi bryophili with regard to species number, host range and geographic distribution. Although most MATERIAL AND METHODS are poorly known from herbarium material and only Measurements and illustrations of microscopic details were single or very few collections exist, substantial infor- done in lactophenol-cotton blue (Merck 13741). Blue reac- mation on the teleomorphic stages is available (e.g. tions of hyphae or spores in lactophenol-cotton blue are von HoÈhnel 1919, Corner 1935, Hawksworth 1973, called cyanophilic or cyanophilous (Kotlaba and Pouzar Racovitza 1959, DoÈbbeler 1978, 1998, 1999, 2004, 1964, Rambold 1995, Kirk et al 2001). Delicate structures such as apical paraphyses and ascospores of some species DoÈbbeler and Carranza 1993, Rossman et al 1999). were analyzed with tap water and lactophenol-cotton blue as The bryophilous Hypocreales display a remarkably mounting media and different optical equipments (phase- variable set of morphological, anatomical and ecolog- contrast optics and differential interference contrast [DIC]). ical features indicating the group is taxonomically di- verse. The tiny ascomata usually are restricted to dis- DESCRIPTIONS OF ASCOSPORE TY PES Accepted for publication 29 June 2005. Unless otherwise stated the following species are 1 E-mail: [email protected] known only from bryophytes. Ascospores are color- 924 DOÈ BBELER:ASCOSPORE DIVERSITY OF BRYOPHILOUS HYPOCREALES 925 FIGS. 1±17. Ascospore diversity of bryophilous Hypocreales. 1. Bryocentria brongniartii (cyanophilic part stippled). 2. Bryocentria metzgeriae (cyanophilic part stippled). 3. Bryonectria callicarpa. 4. Bryonectria sp. 5. Calonectria frullaniae. 6. Nectria brenesii (disarticulating at the septum). 7. Nectria byssophila. 8. Nectria contraria. 9. Nectria foertheri (cell rich in plasmatic content stippled). 10. Nectria gynophila. 11. Nectria lagodes (cell with plasmatic content stippled). 12. Nectria lankesteri. 13. Nectria muscivora (epispor with cyanophilic warts). 14. Nectria sanramonensis (epispor with cyanophilic striations). 15. Thy- ronectria hyperantarctica. 16. Ticonectria perianthii. 17. Trichonectria hirta. All ascospores are drawn at the same magni®cation. An asterix indicates ascospores of species growing on Frullania dilatata. Bar 5 15 mm. less and formed eight per ascus. Specimens exam- ranged in asci but generally orientated parallel to ined and their herbarium location is given in paren- long axis of ascus (GERMANY, BAVARIA: between St theses. Ottilien and TuÈrkenfeld, 1 May 1999, P. DoÈbbeler 7091, GZU). Bryocentria brongniartii (P. Crouan et H. Crouan) DoÈbbeler, Mycol Progr 3:248 (2004) FIG.1 Bryonectria callicarpa DoÈbbeler, Sendtnera 6:93 Biotrophic leaf-perforating parasite on Frullania di- (1999) FIG.3 latata (L.) Dumort.; Europe. Biotrophic parasite on the ventral side of Frullania Ascospores dumbbell-shaped, rarely almost cylin- dilatata; Europe. drical, two-celled but seemingly one-celled, with a Ascospores ellipsoid to fusiform, two-celled, not central cyanophilic band-like part, (6±)6.5±7.5(±8) 3 constricted at the septum, (11±)12±14(±15) 3 4±5 2±2.5 mm; irregularly arranged in asci but generally mm, colorless but with a large yellow lipid body in orientated parallel to long axis of ascus, sometimes each cell and often an additional smaller one, epis- in pairs of two spores lying side by side (GERMANY, pore slightly cyanophilic (SPAIN: Catalonian Pyre- BAVARIA: Kohlgraben southwest of Dieûen, 4 May nees, between Figueres and Olot, 20 Aug 1999, P. 2003, P. DoÈbbeler 7741, M). DoÈbbeler 7094, HOLOTY PE M). Bryocentria metzgeriae (Ade et HoÈhn.) DoÈbbeler, Bryonectria sp. FIG.4 Mycol Progr 3:252 (2004) FIG.2 Biotrophic parasite on Frullania dilatata; Romania. Necrotrophic leaf-perforating parasite on hepatics in- Ascospores cylindrical to narrowly ellipsoid, two- cluding Frullania dilatata; Europe. septate, 18.5±23 3 4.5±5 mm, epispore minutely Ascospores fusiform, two-celled but seemingly one- rough and cyanophilic, 4 ascospores formed per as- celled, with a central cyanophilic band-like part, cus (ROMANIA: ``Centralcarpathen, an den StaÈm- (4.5±)5.5±7.5(±8.5) 3 1.5±2(±2.5) mm, irregularly ar- men der NadelbaÈume'' (without further details), 926 MYCOLOGIA Kalchbrenner, Rabenhorst, Hep. Eur. 156, sub F. di- Ascospores spherical to broadly ellipsoid, often at latata, HBG). one or both ends impressed and then almost cube- shaped, two-celled, 4±5(±5.5) mm diam, epispore smooth, cyanophilic (IVORY COAST: Parc National Calonectria frullaniae Racovitza, MeÂm Mus Natl Hist de Tai, 18 Dec 1975, L. Ake Assi 13160/A, HOLO- Nat, SeÂr B, Bot 10:28, 91 (1959) FIG.5 TY PE EGR). Necrotrophic parasite within perianths of Frullania dilatata; Europe. Ascospores cylindrical, straight or slightly bent, Nectria lagodes DoÈbbeler, this publication FIG.11 rounded at both ends, with up to seven transverse walls, 42±57 3 2.5±3 mm (ITALY, TUSCANY: San Polo in Chianti, 6 Sep 2001, P. DoÈbbeler 7377, M). Nectria lankesteri DoÈbbeler et J. Carranza, Rev Biol Trop 41:204 (1993) FIG.12 Biotrophic parasite on epiphyllous Lejeuneaceae; Nectria brenesii DoÈbbeler et J. Carranza, Rev Biol Costa Rica. Trop 41:203 (1993) FIG.6 Ascospores ellipsoid, two-celled, with thick walls Biotrophic parasite on epiphyllous Lejeuneaceae; and subglobose lumina, 5±6.5 3 3.5±4.5(±5) mm, ep- Costa Rica. ispore smooth or rough, slightly cyanophilic (COSTA Ascospores bacilliform, bicellular when immature, RICA: JardõÂn Lankester southeast of Cartago, 11 Feb at maturity disarticulating at the