ARTICLE 173 CORE

, has and

as type, Phacidium Cytisporae brought to you by by you to brought (Sutton 1980, was included), 1 C. phaeocomes but that option was not Key words: Bulgariaceae coelomycete discomycete Gremmenia Helotiales LSU Phacidiales RPB2 systematics (asexual). Ceuthospora Wageningen University & Research Publications Research & University Wageningen by provided , and referred both Sphaeria , 5(2): 173–193 (2014) 173–193 · 5(2): Fungus IMA

is ) is , which is becomes , and Johannes Z. Groenewald , and Johannes a pathogen of 5 to that genus. The genus. that to and Sclerotia inclusum C. phacidioides, C. lauri, and , along with two new as type species of Phacidum Ceuthospora, was thus not clearly typified. Fries (currently Helotiales was a single fungus with two morphs (pleomorphic), and Tulasne & Tulasne (1861–65) recognised that Tulasne & Tulasne The generic name Ceuthosporawas published by in The Netherlands. The available in 1981. ilicis be representative of to the conidiomatal states (spermatial) and Ceuthosporae to Phacidium been linked as an asexual morph As with Phacidium , the however, Nag Raj 1993). typification of the asexual name was also beset by problems. Fries (1825) with two species, Sphaeria phaeocomes Sclerotium inclusum. Greville (1828) included another three species, all accepted by Fries (1832). Fries (1849) introduced the generic name Pyrenophora phaeocomes genus selected Sphaeria phaeocomes (1832) effectively original protologue (in this case Fries 1815) or the sanctioning lacerum work (in this case Fries 1823, where P. so there would no objection be now to the acceptance P.of lacerum pseudotsugae, ), as sister to Helotiales A. Phacidiaceae . as , David L. Hawksworth , David L. 4 , and , has traditionally been linked to Phacidium subsp. germanicum Phacidium , a species (Leotiomycetes . , and consequently the family Bulgariaceae Ceuthospora lacerum , Karen Hansen 1 Bulgaria 2013), al. (McNeill et was introduced Fries by is introduced to accommodate P. integerrimum P. includes . Phacidium ) are congeneric: taxonomic and taxonomic and ) are congeneric: (Phacidiaceae Ceuthospora is resurrected to accommodate the snow-blight pathogens of conifers, . Several new combinations are introduced in Phacidium

, William Quaedvlieg Phacidiaceae 1,2,3 Gremmenia was subsequently transferred to Excipula The morphologically diverse genus Trichophorum cespitosum , which occurs on Trichophorum

specified by the author or licensor (but not in any way that suggests that they endorse you or your use of the work). must attribute the work in the manner You P. pseudophacidioideson Ilex and Chamaespartium which occurs in Europe, and Phacidium , P.

1988). While the proposal to While the 1988). al. 1988 (Greuter et of

Departamento de Biología Vegetal II, Facultad de Farmacia, Universidad Complutense de Madrid, Plaza Ramón y Cajal, Madrid 28040, Spain; Cajal, Madrid 28040, de Madrid, Plaza Ramón y Facultad de Farmacia, UniversidadComplutense II, Vegetal Departamento de Biología Wageningen University and Research Centre (WUR), Laboratory of Phytopathology, Droevendaalsesteeg 1, 6708 PB Wageningen, The Droevendaalsesteeg 1, 6708 PB Wageningen, University and Research of Phytopathology, Centre (WUR), Laboratory Wageningen Box 50007, SE-104 05 Stockholm, Sweden P.O. Department of Botany, History, Swedish Museum of Natural CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands; corresponding author e-mail: [email protected] Utrecht, Fungal Biodiversity Uppsalalaan Centre, 8, 3584 CT CBS-KNAW Africa Pretoria, Pretoria, 0002, South Biotechnology Institute, University of Agricultural Forestry and Submitted: 15 April 2014; Accepted: 29 September 2014; Published: 9 October 2014. Accepted: April 2014; Article info: Submitted: 15 G. abietis, G. infestans, and G. pini-cembrae. sexual morphs via association, though molecular or cultural data to confirm this relationship have been the relationship The aim of this study was thus to resolve genera by generating of these two lacking. nucleotide sequence data for three loci, ITS, LSU and RPB2. Based on these results, Ceuthospora reduced to synonymy under the older generic name Phacidium Phacidiales suggested to constitute a separate order, species, trichophori generic name Allantophomopsiella conifers, while clearly paraphyletic. Abstract: Department of Life Sciences, The Natural History Museum, Cromwell The Natural History Museum, Road, London and Mycology SW7 5BD, UK; Section, Royal Botanic Department of Life Sciences, 3DS, UK TW9 Surrey Gardens, Kew, a synonym of Phacidiaceae Netherlands © 2014 International Mycological Association © 2014 International Mycological conditions: distribute and transmit the work, under the following are free to share - to copy, You Attribution: 3 4 5 nomenclatural implications nomenclatural 1 2 Pedro W. Crous Pedro W. and Phacidium Non-commercial: No derivative works: Any of the above conditions can be waived if you get For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. transform, or build upon this work. may not alter, You may not use this work for commercial purposes. You Nothing in this license impairs or restricts the author’s moral rights. permission from the copyright holder. described later by Fries (1818), was therefore considered the type species of the genus (von Höhnel 1917), even though the two species originally one of described by not it was Fries. Subsequent authors have accepted Arx & Müller 1954, Reid & Cain 1962, this 1942, von (Terrier typification Korf 1973, Dennis 1978). In order to regularise this situation, Hawksworth & Sherwood (1981) proposed the conservation as the conserved type species; lacerum of Phacidium with P. this proposal was accepted by the Committee for Fungi and Lichens and incorporated into the appendices of the Berlin Code conserve the type was under discussion, DiCosmo et al. lacerum (1984) followed Höhnel (1917) and accepted P. Melbourne Code Under the type. the types for sanctioned names can be selected either from the volume 5 · no. 2 The generic Phacidiumname Introduction and coronatum (1815) for P. integerrimum P. coronatum to Coccomyces P. doi:10.5598/imafungus.2014.05.02.02 View metadata, citation and similar papers at core.ac.uk at papers similar and citation metadata, View 174 ARTICLE

Table 1. Collection details and GenBank accession numbers of isolates included in this study. Species Isolate no.1 Host Location Collector GenBank accession no.2 LSU ITS RPB2 Allantophomopsiella pseudotsugae CBS 288.37 Picea abies United Kingdom – KJ663863 KJ663824 KJ663904 CBS 320.53 Pseudotsuga menziesii Norway H. Robak KJ663864 KJ663825 KJ663905 CBS 321.53 Picea abies Norway H. Robak KJ663865 KJ663826 KJ663906 CBS 437.71 Pinus sylvestris Netherlands J. Gremmen KJ663866 KJ663827 KJ663907 CBS 562.63 Pinus sylvestris Norway F. Roll-Hansen KJ663867 KJ663828 KJ663908 CBS 841.91 Pinus sp. Germany P. Schumacher KJ663868 KJ663829 KJ663909 Allantophomopsis cytisporea CBS 262.85 Roots of Conifer species Germany H. Courtois KJ663869 KJ663830 KJ663910 ‘Allantophomopsis’ sp. CBS 109.22 Oxycoccus macrocarpos USA C.L. Shear KJ663861 KJ663822 KJ663902 Allantophomopsis sp. CBS 322.36 Pinus radiata New Zealand P.C. Birch KJ663880 KJ663839 KJ663921 Bulgaria inquinans CBS 118.31 Forest floor Germany BIHM KJ663870 KJ663831 KJ663911 CBS 129.58 Forest floor Germany – KJ663871 KJ663832 KJ663912 CBS 145.55 – Germany H. Lyr KJ663872 KM216394 KJ663913 CBS 315.71 Quercus robur Switzerland E. Müller KJ663873 KJ663833 KJ663914 Crous etal. Coleophoma sp. CBS 449.70 Liriodendron tulipifera Netherlands H.A. van der Aa KJ663874 KJ663834 KJ663915 Gremmenia infestans CBS 396.48 – Sweden E. Björkman KJ663876 KM216393 KJ663917 Hyaloscypha sp. CBS 109453 Miconia sp. Venezuela I. Hernandez KJ663875 KJ663835 KJ663916 Neofabraea sp. CBS 135481 = CPC 22154 Polygonatum sp. Netherlands U. Damm KF251745 KF251242 KJ663942 Phacidium calderae CBS 287.72 Arbutus unedo Italy W. Gams & J.A. Stalpers KJ663878 KJ663837 KJ663919 CBS 372.81 Pistacia terebinthus Spain H.A. van der Aa KJ663879 KJ663838 KJ663920 Phacidium fennicum CBS 457.83 Pinus sylvestris Finland H. Butin KJ663881 KJ663840 KJ663922 Phacidium lacerum CBS 130.30 Pinus sylvestris Netherlands J.Y. van Vliet KJ663882 KJ663841 KJ663923 CBS 338.70 Ilex aquifolium Netherlands H.A. van der Aa KJ663883 KJ663842 KJ663924 CBS 400.81 Juniperus communis France O. Petrini KJ663884 KJ663843 KJ663925 CBS 540.70 Pinus sylvestris Netherlands H.A. van der Aa KJ663885 KJ663844 KJ663926 CBS 557.70 Sciadopitys verticillata Netherlands H.A. van der Aa KJ663886 KJ663845 KJ663927 CBS 761.73 Pinus sylvestris France M. Morelet KJ663887 KJ663846 KJ663928 Phacidium lauri CBS 198.68 Vinca minor Netherlands H.A. van der Aa KJ663890 KJ663849 KJ663930 CBS 308.68 Prunus laurocerasus Netherlands H.A. van der Aa KJ663891 KJ663850 KJ663931 CBS 443.71 Ilex aquifolium Netherlands L. Marvanová KM216397 KM216392 KM216396

ima fUNGUS CBS 589.67 Ilex aquifolium Netherlands H.A. van der Aa KJ663892 KJ663851 KJ663932 Phacidium mollerianum CBS 365.72 Eucalyptus sp. Italy W. Gams KJ663888 KJ663847 KJ663929 CBS 574.66 Polygonatum odoratum Netherlands H.A. van der Aa KJ663889 KJ663848 KM216395 Phacidium pseudophacidioides CBS 497.72 Chamaespartium sagittale Switzerland E. Müller KJ663893 KJ663852 KJ663933 Phacidium and Ceuthospora

when he excluded Sclerotium inclusum from the genus. ARTICLE However, Pyrenophora phaeocomes was selected as

2 type species of Pyrenophora by Shoemaker (1961), which effectively leaves Pyrenophora as a later nomenclatural KJ663918 KJ663941 KJ663940 KJ663903 KJ663939 KJ663938 KJ663937 KJ663936 KJ663935 KJ663934 R PB2 synonym of Ceuthospora. To resolve this situation, Sutton (1972) proposed the conservation of Ceuthospora Grev. 1826 with C. lauri as type, over Ceuthospora Fr. 1825 with C. phaeocomes as type; this was accepted by the KJ663836 KJ663860 KJ663859 KJ663823 KJ663858 KJ663857 KJ663856 KJ663855 KJ663854 KJ663853 I TS Committee for Fungi and Lichens and the name included in the appendices to the Leningrad Code (Stafleu et al.

G enBank accession no. 1978). The family name Phacidiaceae was introduced by Fries (1849) and accepted by Karsten (1871) who stressed that KJ663877 KJ663901 KJ663900 KJ663862 KJ663899 KJ663898 KJ663897 KJ663896 KJ663895 KJ663894 L SU members had a particular apothecium type with a reduced exciple. Nannfeldt (1932) recognised 14 genera in the family, but Terrier (1942) felt that the family contained different elements, and probably should only include five genera. Von Arx & Müller (1954) regarded eight genera as belonging to the family. The circumscription remained a bone of contention, as Kreisel (1969) recognised four genera, Korf (1973) nine, Dennis (1978) 13, and Lanier et al. (1978) seven. DiCosmo et al. (1984) listed 24 genera W.G. Ziller & A. Funk Ziller & W.G. J. Gremmen G.S. Roosje Zweede G. Verkley G. Verkley G. Verkley L. Marvanová W. Quaedvlieg W. H.A. van der Aa H.A. van der C ollector in Phacidiaceae, stating that it was heterogeneous and in need of further study. In a recent phylogenetic study of Leotiomycetes (Wang et al. 2006b), Helotiales (incl. Phacidiaceae) was left untreated as a large, polyphyletic assemblage, due to inadequate molecular data and sampling. However, they did refer to Phacidiaceae, based British Columbia Switzerland Netherlands Netherlands Czech Republic Czech Republic Czech Republic Netherlands Netherlands Netherlands Location on analyses including a LSU sequence of Phacidium lacerum (data not shown), which suggested a sister group relationship with Phacidiopycnis pyri (sexual morph: Potebniamyces pyri), previously questionably placed in Rhytismatales). In the SSU-LSU-5.8S phylogeny (Wang et al. 2006b), P. pyri formed a strongly supported monophyletic group with Bulgaria inquinans (Bulgariaceae), and P. pyri together with Holwaya mucida (although without support) was moved to Bulgariaceae. Bulgaria has large, brown- Pinus contorta Picea abies Pyrus communis Pyrus communis Vincetoxicum officinale Vincetoxicum Vincetoxicum officinale Vincetoxicum Vincetoxicum officinale Vincetoxicum Vaccinium vitis-idaea Vaccinium Trichophorum cespitosum Trichophorum subsp. germanicum Ilex aquifolium Host black to black, turbinate, gelatinous apothecia with brown-walled ascospores, and is thus quite different from Phacidium and Potebniamyces in morphology. In the LSU & SSU phylogeny generated by Lantz et al. (2011), however, these three genera clustered together in a well- supported clade. 1 Given the recent decision to abolish dual nomenclature for fungi (Hawksworth 2011, Hawksworth et al. 2011, Wingfield et al. 2012), we wanted to resolve the issue CBS 273.74 CBS 377.59 CBS 322.63 CBS 282.55 CBS 123743 CBS 123727 CBS 123726 CBS 444.71 CBS 138246 = CPC 22952 CBS 590.69 Isolate no. of whether Phacidium and Ceuthospora were really congeneric (Johnston et al. 2014) and fix the application of these names using genetic data, and further to determine the phylogenetic position of Phacidiaceae within Helotiales.

Materials and methods

Isolates Isolates used in this study were obtained from the culture collection of the CBS-KNAW Fungal Biodiversity Centre (CBS). Utrecht, The Netherlands (Table 1). Colonies were established on Petri dishes containing 2 % malt extract able 1. (Continued). CBS: CBS-KNAW Fungal Biodiversity Centre, Utrecht, The Netherlands; CPC: Culture collection of Pedro Crous, housed at CBS. Fungal Biodiversity Centre, Utrecht, CBS: CBS-KNAW polymerase II second largest subunit gene. gene; ITS: internal transcribed spacers and intervening 5.8S nrDNA; RPB2: partial RNA LSU: large subunit (28S) of the nrRNA Sarcotrochila longispora Pseudophacidium ledi Potebniamyces pyri Phlyctema vincetoxici Phacidium vaccinii Phacidium trichophori T S pecies 1 2 agar (MEA), potato-dextrose agar (PDA), and oatmeal volume 5 · no. 2 175 176 ARTICLE

of the individual datasets, a 70 % neighbour-joining (NJ) of datasets, a70%neighbour-joining the individual in BioEditv. 7.0.5.2 (Hall1999). To checkthecongruency if necessary,and html; Katohetal.2002) improved manually MAFFT v. 7 (http://mafft.cbrc.jp/alignment /server/index. alignment of the obtained sequence data was first done using (RPB2). locus largest subunit II second polymerase A basic (Quaedvlieg region (ITS), fRPB2-5F (Liuetal.1999),andfRPB2-414R spacer transcribed the internal used toamplify were al. 1990) the partial 28SrRNA gene (LSU), ITS5 and ITS4 (White and LR5(Vilgalys2009a) & Hester1990)wereusedtoamplify protocol. The primersLSU1Fd(Crousetal . manufacturer’s to the Beach, CA)according Solana (MoBio Laboratories, on MEADNA usingtheUltraCleanTMMicrobial IsolationKit Genomic DNAgrowing wasextractedfromfungalcolonies DNA isolation,amplificationandanalyses continuous near-ultravioletlighttopromotesporulation. agar (OA) (Crous et al. J01355). The scalebarindicates0.1expectedchangespersite. wasusedfortheBayesiananalysis. diagnostic convergence for thetopological The treewasrootedtoSaccharomycescerevisiae support Bayesian posterior probabilitiesvaluesfor the respective nodesaredisplayedinthe tree. A stop rule(set to 0.01) for the criticalvalue Fig. 1. Ato Leotiomycetes belonging isolates representative tree containing consensus rule LSU % majority 50 Bayesian

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Vibrisseaceae (continued) Loramycetaceae Rhytismataceae Geoglossaceae Dermateaceae Dermateaceae Incertae sedis Incertae Erysiphaceae Helotiaceae Helotiaceae and related classes. related and Rhytismatales Geoglossales Erysiphales (continued) Helotiales Helotiales Helotiales ima fUNGUS ). (GenBank Phacidium and Ceuthospora

DQ257352 Chlorovibrissea sp. 1 Vibrisseaceae ARTICLE AY789383 Vibrissea albofusca 0.63 AY544674 Lachnum bicolor cf. bicolor Hyaloscyphaceae AY544646 Lachnum virgineum 0.65 1 AY789341 Cudoniella sp. 1 AY789365 Ombrophila violacea Helotiaceae 1 AY789373 Cudoniella clavus AY789431 Hymenoscyphus scutula 1 Hyaloscypha sp. CBS 109453 Hyaloscyphaceae AY789415 Hyaloscypha daedaleae 0.97 1 AY616237 Lambertella tubulosa Rutstroemiaceae AY616238 Spirosphaera floriformis Incertae sedis GU727559 Catenulifera brachyconia Helotiales AY789420 Bryoglossum gracile Helotiaceae FJ176871 Bisporella citrina 1 AY789322 Ciboria sp. 1 AY789347 Sclerotinia sclerotiorum Sclerotiniaceae AY544683 Monilinia fructicola 1 0.97 AY544670 Monilinia laxa 1 AY789407 Scleromitrula shiraiana Rutstroemiaceae Piceomphale bulgarioides 1589 P Incertae sedis 1 JN673040 Chlorencoelia torta Helotiaceae 0.84 Sarcotrochila longispora CBS 273.74 Hemiphacidiaceae 0.53 KC407777 Vestigium trifidum Incertae sedis 0.98 AY789296 Heyderia abietis Helotiaceae 0.99 AF356694 Fabrella tsugae Hemiphacidiaceae 1 FR774268 Neurospora perkinsii Sordariaceae AY780079 Sordaria fimicola 0.73 KC425274 Chaetomium bostrychodes KC425311 Myceliophthora similis Sorda- 1 Chaetomiaceae riales KJ081755 Thielavia terrestris 1 KC425275 Chaetomium megalocarpum KF833362 Sporoschisma saccardoi Chaetosphaeriaceae 0.99 JQ685872 Hypocrea foliicola 0.58 GU205261 Sphaerodes retispora var. retispora 0.58 0.59 Hypo- 1 KF723005 Trichoderma asperellum Hypocreaceae creales JN941457 Hypocrea lutea 1 0.91 AY283547 Gliocladium viride AB593705 Discosia artocreas 0.97 AB593730 Seimatosporium azaleae 0.77 AF382356 Pestalotiopsis bicilia Amphisphaeriaceae Xylariales 1 0.63 0.85 AF382366 Bartalinia robillardoides AB593724 Immersidiscosia eucalypti 0.72 AY789397 Microglossum olivaceum 1 0.79 AY789359 Leotia lubrica DQ257359 Microglossum rufum DQ257362 Microglossum sp. Leotiaceae 0.98 1 KC834019 Alatospora pulchella 0.70 KC834025 Flagellospora leucorhynchos Helotiales 0.60 KC834023 Flagellospora curvula 1 FJ176884 Leuconeurospora pulcherrima Pseudeurotiaceae AF096198 Pseudeurotium zonatum 1 AY544680 Crinula caliciiformis Bulgariaceae DQ257356 Holwaya mucida Helotiaceae EU754161 Eleutheromyces subulatus Incertae sedis DQ470960 Bulgaria inquinans Bulgaria inquinans CBS 118.31 1 Bulgaria inquinans CBS 129.58 Phacidiaceae Phacidiales Bulgaria inquinans CBS 145.55 (continued) (continued) Bulgaria inquinans CBS 315.71 EU107268 Bulgaria inquinans

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s s p p C l o F S S S S r o e e e C C num C r L h c y p. C num C C C o U C A e e p. C o a i p i p p p p p p p p i s pseudotsugaeC pseudotsugaeC pseudotsugaeC pseudotsugaeC pseudotsugaeC pseudotsugaeC p i i h h o r C a C 286759 g o t C e o 198.68 589.67 C C o 308.68 443.71 b a B p o d Y ccu i 704083 l i t o o o o o o o o o o o R s B B B a a a

p l B B B c i m g g c m e t o h c C S i 584657 r r r r r r r r r B p B i r O C B B B C a c c d y l a l l S S S a o S S S t a a a a ssu a a a a a a s o a o e C y 444.71

B C P a b o i i B c S S lic o S J S n S c cce g a d d i m 338.70 400.81 130.30 ssu c

ssu

761.73 557.70 540.70 c

a a B n p r S u s s s s s s s s s C s n h 01355 457.83 o S 322.63 282.55 B i p i i 287.72 372.81 e B e i e l d o o i u p. C p. C p. C p. C p. C p. C p. C p. C p. C c S p. C u o 322.36 i r m s r 22952 109.22 a r s h a S S i i o a i i m b d d 377.59 o u lla m m m p ilo c s u C l 574.66 365.72 i e n o m e e P p p A n h d g a f i s y m c p s s L L L L L L L L L a L B e . l t u P r o r a r h u i C C o l r X X X X X X X X X X e r m t a i b o l S c n i h t i m s S e l n a i y d b 2710 2357 3538 2406 2362 2379 1009 2095 3539 lin i r 1010 g i 262.85 B o B u f c a l a a p f unk o t e i r e n t i o S S i u g r cch u t o r a s r i ile i l m a m e 590.69 497.72 g a m B B B B B B a p r c n a d i ii r e i d a S S S S S S s a a e n a 841.91 288.37 321.53 562.63 320.53 437.71 i d r u d s Crous etal.

o m h y e m t m h g y o a e c s n e a ii s c e r e v i s i a e

Herpotrichiellaceae Botryosphaericeae Phyllostictaceae Mycocaliciaceae Pertusariaceae Geoglossaceae Incertae sedis Incertae Dothideaceae Lecanoraceae Cladoniaceae Peltigeraceae Arthoniaceae Phacidiaceae Roccellaceae Chaetothyriales Mycocaliciales Geoglossales Pertusariales Lecanorales Dothideales Peltigerales sphaeriales Phacidiales Arthoniales Botryo- ima fUNGUS Phacidium and Ceuthospora

CBS 589.67 ARTICLE CBS 198.68 1 Pha. lauri CBS 443.71 CBS 308.68 CBS 590.69 1 Pha. pseudophacidioides 0.99 CBS 497.72 0.56 Pha. trichophori CPC 22952 CBS 574.66 1 Pha. mollerianum CBS 365.72 Phacidiales, Phacidiaceae CBS 372.81 1 Pha. calderae Phacidium CBS 287.72 0.90 CBS 557.70 0.98 0.74 CBS 761.73 CBS 540.70 Pha. lacerum 0.90 1 CBS 130.30 CBS 400.81 1 CBS 338.70 Pha. vaccinii CBS 444 71 0.98 Pha. fennicum CBS 457 83 0.96 CBS 315.71 0.84 0.84 CBS 145.55 B. inquinans 1 CBS 118.31 Bulgaria CBS 129.58 Pse. ledi CBS 377.59 Pseudophacidium CBS 437.71 CBS 320.53 CBS 841.91 Allanto- 1 A. pseudotsugae CBS 562.63 phomopsiella 0.67 CBS 321.53 CBS 288.37 CBS 322.63 1 Pot. pyri 1 CBS 282.55 Potebniamyces Allantophomopsis sp. CBS 322.36 All. cytisporea CBS 262.85 Allantophomopsis 0.90 “Allantophomopsis” sp. CBS 109.22 Helo- G. infestans CBS 396.48 Gremmenia tiaceae Hyaloscypha sp. CBS 109453 Incertae sedis teaceae Derma- 0.55 CBS 123727 1 CBS 123726 Phl. vincetoxici Phlyctema 1 CBS 123743 1 Neofabraea sp. CPC 22154 Coleophoma sp. CBS 449.70 Incertae sedis Sarcotrochila longispora CBS 273.74

0.01

Fig. 2. A Bayesian 50 % majority rule combined ITS, LSU and RPB2 consensus tree containing representative isolates belonging to the Phacidiales and related orders. Bayesian posterior probabilities support values for the respective nodes are displayed in the tree. A stop rule (set to 0.01) for the critical value for the topological convergence diagnostic was used for the Bayesian analysis. The tree was rooted to Sarcotrochila longispora (CBS 273.74). The scale bar indicates 0.01 expected changes per site.

RESULTS tree topology for the 70 % reciprocal bootstrap trees, allowing us to combine them in the multigene analyses. For Fig. 1, Phylogeny the LSU overview dataset of 160 sequences (including the The ITS, RPB2 and LSU sequence datasets of 43 sequences outgroup) contained 830 characters, of which 461 contained (including the outgroup) did not show any conflicts in their unique site patterns. For Fig. 2, the LSU dataset contained volume 5 · no. 2 179 180 ARTICLE

Synonym: Phacidiaceae Fr., Phacidiales Leotiomycetes, Phacidiales,Phacidiaceae Type family:Phacidiaceae subreniform, aseptate,withorwithoutappendages. invested inmucilage.Conidiasubcylindrical,ellipsoid-oblongor Conidiogenous cellsphialidic,attimesproliferatingpercurrently, hyaline, smooth,branched,orreducedtoconidiogenouscells. uni- tomultilocular, singletoaggregated.Conidiophores hyaline, anastomosing,investedinmucilage.Conidiomata brown, withoutsheath.Paraphysesbranchedorsimple,septate, subcylindrical orirregularlyso,straighttocurved,hyaline apical dehiscencering.Ascosporesaseptate,ellipsoidto tears in upper layer. discoid, orimmersed,becomingerumpent,openingbyirregular Saprobic or plant pathogenic. conidia withfunnel-shaped,mucoidapicalappendages. hyaline and subcylindrical, conidiogenesis, have phialidic from the subhymenium. The and arise in mucilage, are embedded anastomose, frequently the asci, extendabove Paraphyses or appendages. sheaths lacking ascospores hyaline rings, andaseptate,ellipsoid, dehiscence with amyloid asci thatareclavate,(4–)8-spored, cells, stroma withverticallyarrangedpseudoparenchymatous black layer splittingintoteethorlobes,anexternally covering with a apothecia by erumpent,cleistohymenial characterised The genusPhacidium T 004 (75%)weresampledforthefinaltree. the final tree; for Fig. 2, 2 672 trees were generated of which2 trees were generated of which 54 198 (75 %) were sampled for combined dataset. During the generation of Fig. 1, 72 262 same BayesianparametersasfortheLSUpartitionin sites. FortheLSUoverviewtree,MrModeltestsuggested frequency state (equal) distribution andinversegamma-shapedratevariationacross fixed using analysed was partition and gamma-shapedratevariationacrosssites,theRPB2 was analysed using a fixed (equal) state frequency distribution gamma-shaped rate variation across sites, the ITS partition using dirichlet(1,1,1,1)statefrequencydistributionandinverse GTR modelbuttheLSUpartitionwasanalysedwithMrBayes 537 characters,ofwhich203containeduniquesitepatterns. the ITSdatasetcontained site patterns,and unique contained 355 characters,ofwhich155 the RPB2datasetcontained site patterns, 819 characters,ofwhich101containedunique (endophytic). Foliicolous, caulicolous,corticicolous, parasiticorsaprobic axonomy as “Phacidicei (1917). (1849); as“Bulgariacei For the combined dataset, all data partitions used the For the combined dataset, all data partitions used the Bulgariaceae Ascomata apothecial, discoid, orcircular, Höhn., ”. clavate, unitunicate with or without Asci clavate, unitunicate with or without Summa veg. Scand . represents inoperculate discomycetes inoperculate represents Ber. Deutsch.Bot.Ges. Fr., ”. Fr. 1849. Ceuthospora Summa veg.Scand.2: 357 circular, superficial, superficial, circular, Ascomata asexual morphs 2: 367 (1849); 34: 416 Crous etal.

Allantophomopsiella, Allantophomopsis, Bulgaria, Allantophomopsis, Allantophomopsiella, in Phacidiaceae accepted Type genus: is betterinthe established literature, and hasmoremembers. chosen hereover the latter, as thefamilynamePhacidiaceae asBulgariaceae in thesamepublication was published clusters inthesameclade. The familynamePhacidiaceae myces, andPseudophacidium. Diagnosis genus Allantophomopsis to the similarity Etymology: Namedafteritsmorphological MycoBank MB809673 Allantophomopsiella of 3–4 layers of darkbrown layers of 3–4 multilocular, irregularly erumpent, wall ostiolate; darkbrown, Description fusiform ornaviculateconidia. inequilaterally and inhaving proliferation percurrent in lacking Phacidiopycnis pyri (syn. germ-slit. Potebniamyces with alongitudinal brown, dark being ascospores four oftheeight and exciple, with a brownish outerexciple,a thick gelatinous medullary ascomata by having butisdistinguished 1968), (Bellemère the related (Fig.1).Morphologically Holwaya it isgeneticallyveryclosetothesetaxa(Figs1–2). because wall of layer,upper tears in hymenium; to expose teethopening by irregular opening erumpent, immersed, becoming initially outside based ontheresultsofWang(2010) Huhndorf groups (2006b), group isunresolved, Phacidiaceae groupwithmembersof the strongly supportedmonophyletic Notes: appendage. subreniform, withorwithoutapical,mucoidfunnel-shaped Conidia percurrently, at timesproliferating in mucilage. invested from innerlayerofconidioma. cells, arising to conidiogenous or reduced smooth, branched, angularis to aggregated,withoneseveralostioles.Walls of or not.Conidiomata mucilage in invested or simple,septatehyaline,anastomosing, branched appendages. gelatinous straight tocurved,lacking so, or irregularlyhyalinebrown,withwithoutgermslit, reagent. ring stainingblueinMelzer’s or withoutamyloiddehiscence with unitunicate, (4–)8-spored, from croziers,clavate, arising basal stromaoftexturaangularis in mucilage; invested periphysoids hyaline smooth-walled, in its cleistohymenial development initscleistohymenial Phacidiaceae textura globulosa Bulgaria hyaline, smooth, subcylindrical, ellipsoid-oblong or ellipsoid-oblong smooth,subcylindrical, hyaline, Phacidiaceae

Ascospores to : DistinctfromApostrasseriaandAllantophomopsis mucida : Conidiomata , itsexactplacementwithinthis and although textura globulosa

), which was also placed in Bulgariaceae was alsoplaced pyri), which

inquinans , placed in Bulgariaceae Phacidium aseptate,ellipsoid,fusoid,subcylindrical Bulgariaceae and does not appear to beclosely anddoesnotappear . Crous,gen.nov. to pycnidial, immersed,becoming (formerlyBulgariaceae based on DNA based datainclude textura angularis Fr. 1815.Othergenera textura angularis Conidiogenous cells phialidic, Conidiogenous uni- to multilocular, uni- single . to is reduced to synonymy isreduced Bulgariaceae Conidiophores textura globulosa . by Lumbsch& ; inner layerof Conidiophores ima fUNGUS Paraphyses resembles

hyaline, ) formsa Potebnia- textura , butis . Asci ,

Phacidium and Ceuthospora ARTICLE

Fig. 3. Allantophomopsiella pseudotsugae (CBS 841.91). A. Conidiomata forming on autoclaved barley leaves. B–E. Conidiogenous cells giving rise to conidia. F. Conidia. Bars: A = 300 µm, all others = 10 µm.

arising from inner layer of conidioma, at times reduced to 321.53). – UK: near Dumfries, dieback of 30-yr-old Picea abies, Oct. conidiogenous cells, branched, septate. Conidiogenous 1937, Peace (CBS 288.37). cells integrated or discrete, ampulliform to subcylindrical or lageniform, hyaline, smooth with minute periclinal thickening Sporulating on PNA: Conidiomata pycnidial, immersed, at apex. Conidia ellipsoid to fusiform, hyaline, smooth, becoming erumpent, irregularly multilocular, to 600 µm aseptate, guttulate, bearing mucoid apical appendages (Type diam, dark brown, ostiolate; wall of 3–4 layers of dark brown C sensu Nag Raj 1993), flabelliform to irregular in shape. textura angularis. Conidiophores arising from inner layer of conidioma, at times reduced to conidiogenous cells, Type species: Allantophomopsiella pseudotsugae (M. branched, septate, 5–15 × 2.5–3.5 µm. Conidiogenous Wilson) Crous 2014. cells integrated or discrete, ampulliform to subcylindrical or lageniform, hyaline, smooth with minute periclinal thickening at apex, 5–8 × 2.5–3 µm. Conidia (4–)5–6(–7) × (2–)3 µm, Allantophomopsiella pseudotsugae (M. Wilson) ellipsoid to fusiform, hyaline, smooth, aseptate, guttulate, Crous, comb. nov. bearing mucoid apical appendages (Type C sensu Nag Raj MycoBank MB809674 1993, only visible in water), flabelliform to irregular in shape. (Fig. 3) Basionym: Phomopsis pseudotsugae M. Wilson, Trans. R. Culture characteristics: Colonies spreading, flat with sparse Scottish Arboricult. Soc. 34(2): 147 (1920). aerial mycelium and feathery margins. On PDA surface Synonyms: Allantophomopsis pseudotsugae (M. Wilson) olivaceous grey, reverse iron-grey. On OA surface olivaceous Nag Raj, Coelom. Anam. App. Conidia: 116 (1993). grey with patches of iron-grey. Phacidiella coniferarum G.G. Hahn, Mycologia 49: 227 (1957). Notes: Distinct from Apostrasseria and Allantophomopsis Phacidium coniferarum (G.G. Hahn) DiCosmo, et al., Canad. in that it lacks percurrent proliferation on its conidiogenous J. Bot. 61: 37 (1983). cells, and has inequilaterally fusiform or naviculate conidia.

Additional synonyms are provided in Nag Raj (1993). Gremmenia: snow-blight pathogens of conifers Specimens examined: Germany: on Pinus wood, Dec. 1991, P. Gremmen (1953) described Phragmonaevia gigaspora Schumacher (CBS 841.91). – The Netherlands: Groesbeek, on as a new pathogen associated with needle blight (snow needles of Pinus sylvestris, Nov. 1970, J. Gremmen (CBS 437.71). mould disease) of Pinus cembra in Europe. In his revision – Norway: on needles of Pinus sylvestris, July 1963, F. Roll-Hansen of Helotiales occurring on conifers, Korf (1962) established (CBS H-15946, culture CBS 562.63); Førde in Sunnfjord, shoot of the genus Gremmenia to accommodate this pathogen, as it Pseudotsuga menziesii, Apr. 1948, H. Robak (CBS 320.53); Gyl in was clearly distinct from Phacidium s. str., lacking a distinct Nordmøre, dead bark of Picea abies, June 1948, H. Robak (CBS upper stroma in its apothecia. He also disagreed with Petrak volume 5 · no. 2 181 182 ARTICLE

Synonym: Basionym MycoBank MB809676 Gremmenia abietis(Dearn.)Crous, (i.e. G.pini-cembrae Type species: curved orreniform,aseptate,finelyguttulate,hyaline,smooth. Melzer’s in reagent. blue reaction giving apex, at flattened stipitate, tip. walled, investedinmucilage,sometimesslightlyswollenatthe Paraphyses simpleorbranched,septate,hyaline,smooth- hyaline smooth-walled hyphae, forming a rise toperiphysoids,investedinmucilage;subhymeniumof becoming hyaline towardsinterior, with inner layer giving clypeus when viewed in section when ascomata are immature, apothecial roofoftexturaglobulosa,brown,appearinglikea backwards, exposingthecreamyhymeniumatmaturity; raising hosttissue,burstingopenvia4–8teachthatcurve or separate,circulartoellipticalinoutline,subepidermal, Plant pathogenic,foliicolous.Ascomatascattered,gregarious Gremmenia Korf, reduced to synonymy with two speciesmadebyDiCosmoetal.(1984),G.gigasporawas not observedinP. infestans . Inadetailedcomparisonofthe frequently haslessthaneightascosporesinitsasci,afeature two speciesbasedonthefactthatPhacidiumpini-cembrae which occurs on Pinus sylvestris,anditssynonym,Phacidiumpini-cembrae abnormal material of (1957), whoconcludedthatP. gigaspora representedoldand accommodate them. 2), and thus the genus species, theyareclearlynotmembersofPhacidiums.str.(Fig. these of range host the define to required is research further which causesdiseaseonAbiesandPseudotsuga.Although pini-cembrae. A thirdspeciesofthiscomplexisP. abietis, thus is more correctly placed in synonymy with than 8-ascospores(clearlyillustratedbyGremmen1953),and gigaspora occursonPinuscembra,andfrequentlyhasless Phacidium infestans,thiswasincorrect,asaPhragmonaevia et al. (1984) listed DiCosmo Although pines. five-needled of needles on occur ellipsoid, frequentlylessthaneightperascus,andapothecia ascospores of formed onneedlesoftwo-needledpines.Incontrast, curved, orcurved-fusiform,eightperascus,withapothecia Ascospores ofP. infestans areelongate-ellipsoid,straight, ascus and ascospore morphologies (DiCosmo but can be distinguished based on different hosts, as well as infestans andP. pini-cembrae aremorphologicallysimilar, non P. abietisRabenh.1844). Mycologia 18: 237(1926). club-shaped, (1–)8-spored, tapering towards base, Asci club-shaped,(1–)8-spored,taperingtowardsbase, biseriate, ellipsoid-elongate, straight to Ascospores biseriate,ellipsoid-elongate,straightto : Phacidium infestans var. (Dearn.)J.Reid&Cain, Phacidium abietis

: 482 (1963) [“1962”] (nom. illegit., [“1962”] (nom.illegit., 54: 482(1963) Art. 53.1; P. pini-cembrae are slightly narrower, elongate- (Gremmen)Korf1962 gigaspora Gremmenia . Korf (1962) distinguished these Pinus cembra. Korf (1962) distinguished these as synonym of Phragmonaevia gigaspora as synonym of (Rehm)Crous2014). Mycologia , which occurs on Phacidium infestans, which occurs on is herewith resurrected to Gremmenia isherewithresurrectedto Phacidium infestans. 54:27(1962). abietis comb.nov. Dearn., Mycologia textura angularis. 1984). et al. 1984). Phacidium Phacidium Phacidium Phacidium Crous etal.

Synonyms: collections and epitypification is required to settle the tosettle required is application ofthename. epitypification and collections than onetaxon. As theculturesprovedtobesterile,fresh sequence data, suggesting that they could representmore Notes: in theDNA listedhere showvariation The isolates young tree,31 Aug. 2011, K. of Pinus Kassavare, ondeadneedles Picea, 22 Nov. 1928, J.H. Faull, CBS 263.31.Lappland,Jokkmokk, balsamea balsamea Björkman Specimens examined: Basionym (Fig. 4) MycoBank MB809677 Gremmenia infestans(P. Karst.)Crous, Ascospores aseptate,ellipsoid toellipsoid-fusoid,uni- to reagent) apical discharge mechanism. blue in Melzer’s or absent.Asciclavate,(4–)8-spored,withamyloid (staining Hymenium ofasciandparaphyses;basal stromapresent inner layerwithperiphysoids, investedinmucilage. dark brownpseudoparenchymatal cellsoftexturaglobulosa, erumpent, rupturing host tissue by irregular stellate splits, of scattered or gregarious, circular, immersed, becoming Foliicolous orcaulicolous.Ascomataamphigenous, Ceuthospora Synonyms: Phacidium Fr., Description andillustrations (Gremmen)Korf,Mycologia gigaspora Gremmenia gigaspora Phragmonaevia Basionym MycoBank MB809678 Gremmenia pini-cembrae(Rehm)Crous,comb. nov. clarify itsphylogeneticposition. of available this pathogen, andit will haveto be recollectedto proportions inIdahoandOregon.Noculturesarepresently Bega (1978)reportedthatthediseasecouldreachepidemic and Notes: Description andillustrations Kryptog-fl. Schweiz Bayer. Bot.Ges cons. (1917). (1962). cons. (1886). Pseudotsuga menziesii Gremmenia abietis , CBS 396.48; location unknown, on needles of Abies needles on unknown, location , CBS396.48; , 30 Nov., 30 1928, , Oct 1931, J.H. Faull, ofA. CBS 265.31;onneedles : : Phacidium infestans P.Phacidium Karst., Hedwigia Phacidium lacerumf. Phacidiostroma Phacidium pini-cembrae Grev., Scot.Crypt.Flora Observ. mycol. 1: 167 (1815);nom. . 13:124(1912). Sweden, Vindeln, hostunknown,1946,E. 9:73(1942). , CBS 264.31; on needles of needles on J.H. Faull,CBS264.31; Hansen &I.Olariaga Gremmen,Sydowia causes snow-blight of Abiesspp. causessnow-blight Höhn.,Ber. dt.bot.Ges. : DiCosmoetal.(1984). : DiCosmoetal.(1984). (Dearness 1926,Faull1930). pini-cembrae sylvestris, stillattachedtoa (Rehm) Terrier, 5: 253(1827);nom. (S). comb. nov.

: 141 (1953). 7: 141 Rehm,Ber. ima fUNGUS

25: 232 35: 420 54: 27 Beitr. Phacidium and Ceuthospora ARTICLE

Fig. 4. Gremmenia infestans (S-F207441). A, B. Ascomata on needles. C, D. Vertical section through ascomata. E–G. Asci and ascospores (F in Meltzer’s solution). H. Ascospores. Bars: A = 1 mm, B = 0.5 mm, all others = 10 µm. biseriate, hyaline, smooth, lacking mucoid appendages. Conidia subcylindrical, smooth, granular, hyaline, (17–)18– Paraphyses septate, hyaline, smooth, branched, 20(–22) × 3.5(–4) µm, with a flared, funnel-shaped apical anastomosing, invested in mucilage. Conidiomata pycnidial, mucoid appendage. immersed, becoming erumpent, uni- to multilocular, brown, with ostiole; wall of textura angularis to textura Culture characteristics: Colonies flat, spreading, with sparse globulosa. Conidiophores branched or simple, septate, aerial mycelium and feathery margins. On PDA and OA hyaline, smooth, invested in mucilage. Conidiogenous cells surface and reverse olivaceous grey. phialidic, at times proliferating percurrently, subcylindrical to ampulliform, smooth, hyaline, invested in mucilage. Conidia Specimens examined: Italy: Sardinia: Tacco di Sta Barbara, fallen subcylindrical, aseptate, hyaline, smooth, with irregular leaf of Arbutus unedo, 10 May 1971, W. Gams & J.A. Stalpers funnel-shaped apical mucilaginous appendage. (CBS 287.72). – Spain: Ibiza: Cala Llonga, fallen leaves of Pistacia terebinthus, 17 Apr. 1981, H.A. van der Aa (CBS H-10269, culture Type species: Phacidium lacerum Fr. 1818, typ. cons. CBS 372.81).

Notes: Ceuthospora calderae was originally described from Phacidium calderae (Urries) Crous, comb. nov. leaves of Pistacia lentiscus, collected from the Canary MycoBank MB809679 Islands. Culture CBS 372.81 closely matches the morphology (Fig. 5) of this species; CBS 287.72 was now sterile. Basionym: Ceuthospora calderae Urries, An. Inst. bot. A.J. Cavanilles 14: 165 (1956) [“1955”]. Phacidium fennicum Butin, Sydowia 37: 21 (1984). Description: Conidiomata multilocular, with large central ostiole, papillate. Conidiophores frequently reduced to Description and illustration: Butin (1984). conidiogenous cells, branched, 1–3-septate, up to 30 µm long, 3–4 µm diam. Conidiogenous cells hyaline, smooth, Specimen examined: Finland: Lempäälä Kulju, on needles of Pinus terminal and subterminal, 5–15 × 2.5–3.5 µm, proliferating sylvestris, 18 Apr. 1982, U. Söderholm (CBS 457.83 – ex-type culture). with periclinal thickening (characteristic of this species). volume 5 · no. 2 183 184 ARTICLE

grey. OnPDA surface olivaceous grey, reverseiron-grey. On OAmycelium. aerial moderate olivaceous surface pale with sparseto spreading characteristics: Colonies Culture appendage, (10–)13–15(–18) × (2.5–)3(–4)µm. mucilaginous funnel-shaped with apex scar, flattened central, Conidia aseptate,subcylindrical,basebluntlyroundedwith proliferating percurrently, or with visible periclinal thickening. walled, (5–)7–15 × 2–3(–4) µm, invested in mucilage; cells, liningtheinnerlayerofconidioma,hyaline,smooth- branched, septate,oftenreducedtophialidicconidiogenous individual ostioles up to 25 µm diam. surface view, withindividualloculesupto170µmdiam, the epidermis,multilocular, 500–1500µmdiam,brownfrom scattered, gregarious,toaggregated,subepidermal,splitting pycnidial, flask-shaped to irregularly subglobose, amphigenous, smooth, anastomosing,investedinmucilage.Conidiomata 9.5–12.5 ×3–3.5µm.Paraphysessimple,septate,hyaline, Ascospores aseptate, hyaline, smooth, guttulate, biseriate, reagent)dischargemechanism,70–110Melzer’s ×7–10µm. Asci clavate,8-spored,withamyloid(strainingbluein hymenium; innerlayerofperiphysoids,coveredinmucilage. opening by3–5teethofbrowntexturaglobulosa,toexpose solitary, initially immersed, subepidermal, becoming erumpent, amphigenous, circular, 300–2000µmdiam,aggregatedto Description: Foliicolousoronconescales.Ascomata Additional synonymsareincludedinNagRaj(1993). Fig. 5. Ceuthospora pinastri Ceuthospora Synonyms: Phacidium lacerum Fr., (Figs 6–7) Hochsch. Wien 2(4):104(1925). : Fr., Phacidium Phacidium calderae (CBS 372.81). Syst. mycol.2:575(1823). Dothidea pinastri (Fr.) Höhn., Observ. mycol.2:312(1818) Fr., Elench.fung.2: 123 (1828). simple or Condiophores simple or A, B. Conidiomata. Mitt. bot. Inst. tech. C, Crous etal. Conidiogenous cells. E, D. F.Conidiogenous Conidia. Bars: A, B = 200 µm, all others= 10 µm.

(UPS). 540.70). –Sweden: on needlesof , 12 sylvestris of Pinus needles Apr. 1970, on CBS 338.70);Hulshorsterzand, culture der Aa(CBSH-10292, Maarschalksbos, on leaves of Ilexaquifolium on leaves Maarschalksbos, verticillata Vliet (CBS130.30);Baarn,Cantonspark,onleavesofSciadopitys The Apr. 1887, of P.on needles & Grove](K(M)189268);Königstein, sylvestris,30 needles, Bastadt, onpine Apr. 1877, MBT178726; CBS761.73–ex-neotypeculture).Germany:Baden, – neotype (CBS H-10302 Bas Rhin,onneedlesofPinussylvestris, 26 Aug. 1968, M. Morelet communis Juniperus Specimens examined Notes: so aneotypeisdesignatedfor Fries’ namehere. and or 1823 Frieshad beforehimin1818 collection” “original (pers. comm.)(Fig.8). That specimenisnot,therefore, the Fries cameto Uppsala in1835 accordingto Stefan Ekman and hasnodate,appearstohavebeencollectedafter & Nannfeldt (1962:38). The UPS specimen is unlocalised by Holm done andthenameislistedas“materialunknown” exsiccate his in material to distribute intended that hehad implying and later“S.S.”(Fries1849) refers to“(Exs.ined.)” (1823) work, Fries In thesanctioning was statedtobedepauperate. the “original collection”by (DiCosmo et al. 1984) inUPS on thishost. commonly in UPSreferredtoas The material distributedthroughout Europe,whereit is widely occurs sylvestris, and from Pinus on needles collections European Netherlands: ofPinussylvestris, on needleFeb.1930, J.Y. van Phacidium lacerum Phacidium , 27 Apr. 1970, W. Krieger [FungiSaxoniciexs.290](K(M)189269).– Scleromycetes Suecici. , May1981,O. Petrini (CBS400.81);Hagenau, : France: Mt. Ventoux, livingneedleof Phacidium lacerum for Phacidium H.A. van der H.A. van Baarn, Aa (CBS557.70); was originally described from described originally was Pinus sylvestris, ex-herb Fries Schroeter [exherbs Thuemen This was not, however, H.A. vander Aa (CBS , July 1969, , July designatedhere, ima fUNGUS H.A. van Phacidium and Ceuthospora ARTICLE

Fig. 6. Phacidium lacerum (CBS 761.73). A, B. Conidiomata on autoclaved barley leaves. C–E. Conidiogenous cells. F. Conidia. Bars: A, B = 300 µm, all others = 10 µm.

Fig. 7. Phacidium lacerum (K(M) 189269). A, B. Ascomata on needles. C–E. Vertical section through ascomata. F, G. Asci and ascospores. H, I. Conidia. Bars: A, B = 1000 µm, all others = 10 µm. volume 5 · no. 2 185 186 ARTICLE

Raj (1993;asexualmorphonly). DiCosmo and illustrations Descriptions Additional synonymsarelistedinNagRaj(1993). Ceuthospora phacidioides Ceuthospora Phacidium multivalve Phacidium Xyloma multivalve Ceuthospora lauri Ceuthospora Synonyms: 1 Fig. 8. – culture ex-epitype);Wageningen, on leavesof of laurocerasus as on : examined Specimens Basionym MycoBank MB810293 Phacidium lauri(Sow.) Crous&D.Hawksw., comb. the diagnosis for this fungus appears on the53 appears forthisfungus the diagnosis the wholevolume wascompletedin1803. before that thetextwas issued dated “1802”,butitseemsunlikely is and name, or diagnosis no has which 9), (Fig. 4 fig. 371 pl. on Ekman. sine loc., on dead leavesof der Aa(CBSH-10271,cultureCBS198.68,asC.feurichii 90, on leavesofVincaEemnesserweg , 3Mar. minor 1968, H.A. van May 1969, The texttheplatesin thisworkarenotnumbered,and to illustrate Sphaeria lauri Ilex aquifolium C. phacidioides (1826); nom.illegit.(Art.52,1). (1817) :Fr., nov. : Fr., (1824). pagin Phacidium lacerum Phacidium Syst. mycol.,Index:167(1832). . [53](1803 : . (1984; sexual and asexual morph) andNag and asexual et al.(1984;sexual L. Marvanová , 2Mar. 1968,H.A.vander – epitype Aa (CBSH-10276 Sphaeria lauriSow., Cryptosphaeria lauri (Sow.)Cryptosphaeria Grev., Syst. mycol.2:576(1823). , 8Oct. 1967, H.A.vander Aa (CBS589.67, (Sow.) Grev.,(1826) Scott.Crypt.Fl.5:254 ); Baarn, Cantonspark, on leaves of Prunus on leaves ); Baarn,Cantonspark, DC.,Fl.franç., designated here, The 1 ). (DC.)J.C.Schmidt,Mykol.Heft. (CBS443.71,asC.phacidioides (UPS – Fries' specimen). Photo: Stefan (UPS–Fries'specimen). Netherlands: Baarn,Drakenburgerweg, Prunus laurocerasus Grev., : Sutton (1972, asexual morph), : Sutton(1972,asexual Col. Fig. Br. Fungi 3 rd Scott. Crypt. Fl. MBT178727; CBS308.68 edn2:303(1805). rd ; the illustration is ; theillustration , G.B.W. Kirby (K Ilex aquifolium Fl. Edin.: 361 ); Baarn, ). – UK: 3: 5: 253 1: 42 sine Crous etal. , Sphaeria lauri Fig. 9. Sowerby’s diagnosis original names Nag Raj1993),andsuspectthattheyhavebeenlost. The see Sutton 1980, these synonyms(also of all material original more details.However,to locatethe wehavebeenunable a lectotypefor investigated inthis study. Di Cosmo past, but appear to be synonymousbasedonthe cultures we aquifolium , Europe),andC.phacidioides Prunus laurocerasus which is the type speciesof the genus Ceuthospora lauri multivalve assexualmorph.Ceuthospora Phacidium (1984) linkC.phacidioides phacidioides including synonyms, several has which 1993), Raj Nag ofC.lauri 3(–3.5) µmfallintothevariation Notes: × Conidia ofthepresentstrains(12–)13–15(–16) is thereforeasuperfluousname. multivalve name multivalve than Phacidium in literature used has priority. Ceuthosporalaurialsoappearsmorecommonly lauri the ICNisSphaeria under is legitimate which epithet species as theearliest by Fries,and both sanctioned also confirmed that Greville’s specimens were on the leaves arestillsometimesused byourcooksassuch.”He nobilis as thetrueBay Laurel (Laurus regarded confusion hasarisen.It has frequently beenerroneously of PrunusLaurocerasus of speaking “Owing tothecustom,inBritain, by Grove(1935:291–292): ex holotype). – holotype of The choiceoftheepithet“lauri Xyloma as isillegitimate Ceuthospora phacidioides Ceuthospora lauri was listed as a synonym when it was introduced; it it wasintroduced; when waslistedasasynonym , commonlyconfusedinthe Europe), havebeen . FromSowerby(1803). Sphaeria lauri[not traced]; K (M) IMI 153020 – slides (NagRaj1993).Furthermore,DiCosmo Xyloma multivalve inG but didnotprovide Phacidium multivalve andPhacidium (conidia10–17×2.5–3µm)to ” bySowerbyisexplained (A) andillustrations et al. (1984) designated as‘theLaurel’,great (7–17 × 2–3µm, (7–17 . Notethatthe ima fUNGUS , thatname ), and its ), and Prunus were (Ilex (B) of et al. B (on A C. , Phacidium and Ceuthospora

Synonyms: Phoma molleriana (Thüm.) Sacc., Syll. fung. : 3 ARTICLE 110 (1884). Macrophoma molleriana (Thüm.) Berl. & Voglino, Syll. fung., Addit. I: 314 (1886). Ceuthospora molleriana (Thüm.) Petr., Annls mycol. 23: 29 (1925).

Description: Conidiomata uni- to multilocular, up to 400 µm diam. Conidiophores branched, up to 45 µm long, 2–3 µm diam. Conidiogenous cells terminal and lateral, 5–10 × 2–3 µm, with prominent periclinal thickening, rarely proliferating percurrently. Conidia hyaline, smooth, granular, subcylindrical, (9–)10–12(–13) × (2–)2.5 µm; apical mucoid appendage only visible when mounted in water.

Culture characteristics: Colonies spreading with sparse aerial Fig. 10. Sphaeria lauri packet with slides made Brian Sutton from mycelium and feathery margins. On PDA and OA surface and Sowerby’s holotype (K(M) IMI 153020). reverse olivaceous grey.

Cultures examined: Italy: Sardinia: Su Cologone, on leaves and not Laurus, contrary to his publications, which led to that of Eucalyptus, 6 May 1971, W. Gams (CBS H-10285, 10286; error being perpetuated although the misidentification was CBS 365.72 – culture). – The Netherlands: Baarn, leaf spot on soon recognized, for example, by Berkeley (1836: 283) who Polygonatum odoratum, 8 Aug. 1966, H.A. van der Aa (CBS 574.66). gave it the English name “Cherry-laurel Ceuthospora” and stated “On dead leaves of Prunus Lauro-cerasus (not Laurus Notes: Ceuthospora molleriana was originally described from nobilis as is stated by Dr. Greville)”. Eucalyptus leaves collected in Portugal (conidia 10–13 × 2–2.5 Sowerby (1803) referred to a single unlocalized, but µm, Petrak 1925), and closely fits the morphology observed presumably English (as his work was devoted to fungi from in the present collections. Although originally described from that country) collection made by the Rev. William Kirby Eucalyptus, this taxon appears to have a wider host range. (1759–1850) when introducing the name Sphaeria lauri (Fig. 9); Kirby was born in and died in Suffolk, England. The original material was located in K by Sutton (1972) who Phacidium pseudophacidioides Crous, sp. nov. prepared sections (now preserved as K(M) IMI 153020; Fig MycoBank MB809681 10) and published drawings made from Kirby’s specimen (Fig. 12) (Sutton 1972: 323 fig. 1). That specimen could not now be re-located in K, despite extensive searches by Begoña Etymology: Named after its morphological similarity to the Aguirre-Hudson, Heidi Döring, and D.L.H. Sutton did, asexual morph, Ceuthospora phacidioides. however, include a sketch of the material he saw which evidently comprised two specimens, the right-hand one Diagnosis: Conidiogenous cells discrete or integrated, of which had Kirby’s name below it and is therefore the subcylindrical with perominent periclinal thickening or holotype of Sowerby’s name. percurrent proliferation, 7–19 × 2–2.5 µm. Conidia hyaline, Sutton (1972, 1980) attributed the epithet “lauri” to smooth, granular, subcylindrical, tapering at ends, bearing a Greville and not Sowerby, in accordance with the Code funnel-shaped mucoid apical appendage, (11–)11.5–12.5(– then in operation which ruled that names of “Fungi caeteri” 13) × (2–)2.5 µm. published before 1 January 1821 were not validly published; that situation changed with the deletion of the later starting Type: The Netherlands: Baarn, Eenmnesserweg 92, on Ilex point dates for fungi at the Sydney IBC in 1981. In using the aquifolium, 8 Aug. 1968, H.A. van der Aa (CBS H-10289 – term “lectotype”, Sutton meant as the type of Ceuthospora holotype; CBS 590.69 – ex-type culture). lauri “Grev.”, and that is clear from his annotation on the packet with the slides (Fig. 10) and not Sowerby’s original Description: Conidiomata stromatic, pycnidioid, scattered, binomial. The use of that term was correct for Greville’s name, black, subepidermal, multiloculate, up to 600 µm diam, with under the pre-1981 Code, as Greville had studied collections papillate ostioles. Conidiophores arising from inner layers of other than Sowerby’s. cavity, subcylindrical, hyaline, smooth, extensively branched, up to 120 µm long, 2.5–3 µm diam, invested in mucus. Conidiogenous cells discrete or integrated, subcylindrical with Phacidium mollerianum (Thüm.) Crous, comb. nov. perominent periclinal thickening or percurrent proliferation, MycoBank MB809680 hyaline, smooth, 7–19 × 2–2.5 µm. Conidia hyaline, smooth, (Fig. 11) granular, subcylindrical, tapering at ends, apex subobtuse, Basionym: Sphaeropsis molleriana Thüm., Inst. Coimbra 27: base with truncate hilum, 1 µm diam, bearing a funnel-shaped 40 (1879). mucoid apical appendage, (11–)11.5–12.5(–13) × (2–)2.5 µm. volume 5 · no. 2 187 188 ARTICLE

497.72). sagittale dead leafofChamaespartium cells andconidia. Fig. 12. 300 µm,allothers=10µm. more extensivelybranchedthan in P.are much andthe conidiophores laurihaslargerconidia, originally identified as Notes: The twoisolatesofC.pseudophacidioides Other specimenexamined Fig. 11. Phacidium molleriana Phacidium (CBS Phacidium pseudophacidioides 590.69). A, B. onautoclavedbarleyleaves.C–E.conidiogenous Conidiomata Conidiophores, F. Conidia.Bars: A =300µm,B400allothers =10µm. C. (i.e. phacidioides : Switzerland: Conidiogenous cells. F,on autoclavedbarleyleaves.B–E.Conidiogenous (CBS574.66).A.Conidiomata , 3 May1952,E. Müller P. pseudophacidioides Zürich: Weiacherberg,on P. lauri ). However, were (CBS Crous etal. . subcylindrical with prominent periclinal thickening, 5–12× with prominentpericlinal subcylindrical Diagnosis collected, Trichophorum Etymology: Named after the host genus on whichit was Phacidium trichophori (Fig. 13) MycoBank MB809682 : Conidiogenous cells discreteorintegrated, Conidiogenous . Crous &Quaedvlieg, G. Conidia.Bars: A = ima fUNGUS sp. nov. Phacidium and Ceuthospora ARTICLE

Fig. 13. Phacidium trichophori (CPC 22952). A, B. Conidiomata in culture. C–E. Conidiophores, conidiogenous cells and conidia. F, G. Conidia. Bars: A, B = 300 µm, all others = 10 µm.

2–3 µm. Conidia hyaline, smooth, granular, subcylindrical, Culture examined: The Netherlands, Loenen, on leaves of Vaccinium tapering towards a truncate basal scar, 0.5–1 µm diam, vitis-idaea, May 1969, L. Marvanová (CBS 444.71). bearing a funnel-shaped mucoid apical appendage, 3–5 µm long, 2–4 µm diam at apex, (9–)10–11(–13) × (2–)2.5(–3) µm. Note: Unfortunately this culture proved to be sterile, and thus its morphology could not be confirmed. Type: The Netherlands: Korenburgerveen, Winterswijk, on Trichophorum cespitosum subsp. germanicum, 29 Apr. 2013, W. Quaedvlieg (CBS H-21816 – holotype; CBS 138246 = Pseudophacidium P. Karst., Acta Soc. Fauna Fl.fenn. CPC 22952 – ex-type culture). 2: 157 (1885) [“1881–1885”]. Synonym: Myxofusicoccum Died., Annls mycol. 19: 68 Description: Foliicolous. Conidiomata pseudostromatic, (1912). pycnidioid, scattered to gregarious, black, subepidermal, uniloculate, up to 300 µm diam; walls of the pseudostroma Caulicolous. Ascomata scattered to gregarious, immersed, 40–90 µm thick. Conidiophores arising from inner layers raising the host epidermis, causing it to rupture; ascomata oblong, of cavity, subcylindrical, hyaline, smooth, thin-walled, hemispherical to irregularly pulvinate, black, carbonaceous; branched, to 30 µm long, invested in mucus. Conidiogenous wall of brown textura globulosa, becoming pale brown towards cells discrete or integrated, subcylindrical with prominent interior; inner layers of subglobose, hyaline cells, invested in periclinal thickening, hyaline, smooth, 5–12 × 2–3 µm. mucilage; subhymenium of hyaline textura intricata; basal stroma Conidia hyaline, smooth, granular, subcylindrical, tapering of pale brown pseudoparenchymatal cells. Asci club-shaped, towards a truncate basal scar, 0.5–1 µm diam, apex 8-spored flattened at apex which does not strain in Melzer’s subobtuse, bearing a funnel-shaped mucoid apical reagent. Ascospores irregularly biseriate, oblong-ellipsoidal, appendage, 3–5 µm long, 2–4 µm diam at apex, (9–)10– straight to curved, hyaline, smooth-walled. Conidiomata brown, 11(–13) × (2–)2.5(–3) µm. pycnidioid, stromatic, gregarious to crowded, immersed, discoid to orbicular, opening by means of irregular ruptures, multilocular; Notes: Phacidium trichophori resembles P. lauri (conidia wall of brown textura globulosa; basal stroma of pale brown (12–)13–15(–16) × 3(–3.5) µm), but is morphologically distinct in textura epidermoidea. Conidiogenous cells phialidic, lining having smaller conidia. Ceuthospora gaeumannii (conidia 8–11 inner cavity, lageniform to ampulliform, hyaline, smooth-walled; × 2–2.5 µm, av. 9 × 2.2 µm; Nag Raj 1993) has slightly smaller proliferating percurrently at apex. Conidia ellipsoid to oblong or conidia, much larger conidiomata (500–1500 µm diam). subreniform, apex rounded, base with truncate scar, guttulate, hyaline, smooth-walled.

Phacidium vaccinii Fr., Syst. Mycol. 2(2): 575 (1823). Type species: Pseudophacidium ledi (Alb. & Schwein.) P. Karst. 1885. Synonyms are listed in DiCosmo et al. (1984). volume 5 · no. 2 189 190 ARTICLE

Fig. 14. A =500µm,allothers10µm. (Smerlis 1969). species The genuscontainssaprobic,andplantpathogenic covering layer, exposing creamto greyish orbrownishdiscs. Ascomata openbymeansofanirregularrupture the discomycetes thatoccuronbarkofhard-orsoftwood. ofPseudophacidium Notes: Species abies : examined Specimen Description Basionym (Fig. 14) Pseudophacidium P.ledi (Alb.&Schwein.) Karst., (4–)5(–6) µm. base truncate,1µmdiam,(8–)9–11(–12)apex subobuse, × third, mostly widestinupper guttulate, thin-walled, hyaline, near apex.Conidia percurrently times 1–2 µm, proliferating × 3–4 smooth, 5–10 hyaline, phialidic, or ampulliform, cells subcylindrical Conidiogenous cells. to condiogenous diam, orreduced 3–4µm long, or not,1–3-septate,to30µm branched subcylindrical, diam. opening by irregular rupture,multilocular, up to 500 µm to pulvinate,brown, scattered, erumpent,irregular 1885”]. Acta Soc.FaunaFlorafenn. (1805)l , Apr.1959, arising from inner cavity,from inner arising Conidiophores hyaline, Pseudophacidium ledi Pseudophacidium : : Sporulating onPNA. Xyloma ledi J. Gremmen(CBS377.59). , Graubünden, Bergün, on Picea Bergün, Switzerland, Graubünden, Alb. &Schwein.,Consp.fung.:60 Conidiogenous cells giving rise toconidia. giving cells on PNA. B–E. Conidiogenous forming (CBS377.59).A.Conidiomata solitary,to oblong, ellipsoid : 157 (1885) [“1881– 2: 157(1885) Conidiomata areimmersed stromatic, Crous etal. Basionym Dennis, pyri (Berk.&Broome) Potebniamyces reagent. in Meltzer’s staining smooth-walled. numerous, subcylindrical,branchedorseptate,hyaline, brown inwater, in2%KOH. turning blue-green red- hymenium investedinmucilage; periphysoids, hyaline septate, branched, consists ofsubcylindrical, which region, of brown textura globulosa opening by irregular ruptureto expose hymenium;apicalroof initially immersed, becoming erumpent,irregularinoutline, and corticolous.Ascomatascatteredtogregarious, Caulicolous Potebniamyces Smerlis,Canad.J.Bot. 1978. Type species: DiCosmo yellow, fromSutton1980,and (adapted smooth-walled obovoid, guttulate,aseptate,base truncate, hyaline to pale to curved, smooth-walled.Conidia hyaline, cavity,to ampulliform, straight lageniform subcylindrical, prismatica. above, palebrownbelow, with walloftextura yellow-brown wall of brown textura to globulosa ostiolate, multilocular,rupture; by irregular brown,opening depressed globose,papillate,non- immersed, subepidermal, yellow, smooth-walled. to subglobose ellipsoid,aseptate, guttulate, hyalineto pale (1962). Mag. nat.Hist. , ser.17: 144(1876). 4 Ascom., 2 et al.1984). : Stictis lecanora phialidic, lining the inner lining cells phialidic, Conidiogenous Potebniamyces pyri (Berk.&Broome)Dennis Potebniamyces nd Asci club-shaped, 8-spored, withapex not edn:231(1978). Conidiomata var. , becoming palertowardsinner Ascospores pyri Berk. & Broome, Ann. scatteredtoaggregated, textura epidermoidea uni- to biseriate, uni- to subglobose F. Conidia. Bars: Conidia. ima fUNGUS Paraphyses 40: 352 Brit.

Phacidium and Ceuthospora

Specimen examined: : Wilhelminadorp, bark such as Leotia, Neobulgaria and Ombrophila (Helotiaceae)

The Netherlands ARTICLE of Pyrus communis, Mar. 1963, G.S. Roosje,(CBS 322.63); are not closely related (Wang 2006a, b), Lantz et al. (2011) Wageningen, on peduncle of P. communis, Mar. 1955, Zweede (CBS placed Bulgaria in a strongly supported clade with Phacidium 282.55). and Potebniamyces. The relationship between Bulgaria and Phacidium was also confirmed by Hustad & Miller (2011). Notes: Although we assume that Phacidiopycnis is the The ultrastructure of the ascus apical apparatus of Bulgaria, asexual morph of Potebniamyces, the type species of however, is unique compared to other members of Helotiales Phacidiopycnis, P. malorum, still needs to be recollected (Verkley 1992, 1994). For the present, we treat Bulgariaceae to confirm this assumption. For this reason the synonymy as a synonym of Phacidiaceae, but it could be that the family between these two generic names remains unconfirmed. If is still heterogeneous, and as more members of Phacidiaceae shown to be synonymous, the older name, Phacidiopycnis, and Helotiaceae get added in future molecular studies, the will have preference for the holomorph (Johnston et al. 2014). Phacidiaceae may yet prove to be paraphyletic. In our results, the genus Allantophomopsis (based on A. cytisporea), clusters sister to the canker pathogen Discussion Potebniamyces pyri (asexual morph: Phacidiopycnis pyri). Potebniamyces produces black, gelatinous discs, erumpent The primary aim of this study was to obtain molecular support from submerged stromata in tree bark. Although DiCosmo for the assumption that Ceuthospora and Phacidium are et al. (1984) considered this genus a possible member of congeneric, and at the same time elucidate the position of Phacidiaceae, Rhytismataceae or Dermateaceae, it appears Phacidiaceae within Leotiomycetes, as the latter family and to be a member of an independent family. As the majority of order were not represented in the phylogenetic study of Wang the families in Helotiales still lack molecular data, we consider et al. (2006b). it premature to introduce any new family name here to The order Helotiales contains members that represent accommodate Potebniamyces. DiCosmo et al. (1984) discussed a broad range of ecologies, including species that are the morphological differences between Potebniamyces and plant pathogenic, saprobic, endophytic, mycorrhizal or Phacidium, and Wang et al. (2006b) suggested a possible sister ectomycorrhizal parasites, aquatic saprobes, or wood rotting relationship between the two genera. fungi (Wang et al. 2006a). Although lacking many genera of In the present study we introduce the generic name Helotiales in their phylogenetic analysis, Wang et al. (2006b) Allantophomopsiella to accommodate A. pseudotsugae, concluded that the concept of Helotiales adopted by Eriksson a pathogen of conifers (Roll Hansen 1992). The genus (2005) included many non-monophyletic taxa. They placed Gremmenia is resurrected to accommodate the snow-blight 13 families in the order. In spite of earlier work on the class pathogens of conifers, namely G. abietis (on Abies spp. and (Gernandt et al. 2001, Lutzoni et al. 2004, Wang et al. 2005), Pseudotsuga menziesii), G. infestans (on Abies balsamea), no members of Phacidium were present in the phylogeny of and G. pini-cembrae (on Pinus cembra). Wang et al. (2006b). By adding Phacidiaceae to Helotiales Finally, the position of the Phacidiaceae is clarified within data present in GenBank, Phacidiales clearly separated from the resurrected Phacidiales, with respectively a neotype and Helotiales, which still appear heterogeneous (Fig. 1). The an epitype designated to fix the phylogenetic application separation of Phacidiales from Helotiales is also supported by of the names Phacidium lacerum (type of Phacidium) and the phylogeny of Wang et al. (2006b), as Bulgaria inquinans Sphaeria lauri (type of Ceuthospora) (i.e. Phacidium lauri). and Phacidiophicnes pyri cluster outside of the Helotiales Phacidium (1815) is accepted as the correct name for this clade in their study. genus over Ceuthospora (1827), based on nomenclatural A surprising result of our study concerns the phylogenetic priority, as both names are conserved; it also has a greater position of Bulgariaceae. The genus Bulgaria, based on number of species. B. inquinans, effectively clusters in Phacidiaceae (based on P. lacerum). A second species has been reported, B. nana (Döring & Triebel 1998, Wang 2006b), but that is Acknowledgements considered a species of Austrocenangium (see Gamundi 1997). Bulgaria inquinans is characterised by large (up to We thank the technical staff, Arien van Iperen (cultures), Marjan 4 cm), dark brown to black turbinate, gelatinous apothecia Vermaas (photographic plates), Mieke Starink-Willemse (DNA with brown ascospores. It occurs on bark of fallen trunks and isolation, amplification and sequencing), and Emma Hultén branches, but has also been observed on living trees. The (macroscopic photographs of Gremmenia infestans) for their species is considered saprobic, and possibly facultatively invaluable assistance. In addition, Stefan Ekman searched for plant pathogenic. The relationship of Bulgaria has been material of Phacidium lacerum in UPS and provided information on debatable. It has been placed close to other species the single Friesian specimen there, and Begoña Aguirre-Hudson producing gelatinous apothecia in Leotiaceae/Helotiaceae, and Heidi Döring kindly gave of their time in quests to re-locate the e.g. Leotia, Neobulgaria, Ombrophila (in Leotieae, Korf Sowerby material of Sphaeria lauri in K. We are also thankful for 1973; in Ombrophiloideae, Dennis 1978), or considered to be the review and subsequent intense discussions with H.O. Baral, who rather distinct, and placed as the sole genus in Bulgariaceae brought many aspects to our attention, and provided a better insight (Helotiales) (e.g. Eriksson 2006). Although some molecular into this exciting group of fungi. This work was carried out while phylogenetic studies have not been able to place Bulgaria D.L.H. was in receipt of a grant from the Ministerio de Economía y in a larger group with support, and instead suggest species Competitividad of Spain (project CGL2011-25003). volume 5 · no. 2 191 192 ARTICLE

Gernandt DS, Platt JL, Stone JK, Spatafora JW,DS, Platt JL, StoneJK,Spatafora Gernandt Holst-Jensen A, Gamundi IJ(1997)Austrocenangium Arx JA von, MüllerE (1954) DieGattungenderamerosporen References Gremmen J (1953) Some noteworthy discomycetous fungion Gremmen J(1953)Somenoteworthy Fries EM(1849) Greville RK (1828) Greville Fries EM(1832)SystemaMycologicum Fries EM (1825) Systema Orbis Vegetabilis Fries EM(1823)SystemaMycologicum Faull JH(1930) The spread andcontrolof Phacidium blightinspruce Eriksson OE (ed.) (2006)OutlineofAscomycota –2006.Myconet Eriksson OE (ed.) (2005) Notesonascomycetesystematics. Nos. Döring H, Triebelof D relationships (1998) Phylogenetic DiCosmo F, Nag Raj TR, Kendrick WB (1984) A revisionof the . [The English Flora bySirJamesEdward Fungi.[TheEnglish MJ (1836) Berkeley de dévelopment du l’étude a Contribution (1968) A Bellemère Disease of Pacific Coast Conifers Coast Pacific Bega RVof (1978) Disease Fries EM (1815) ObservationesMycologicae Crous PW, GamsW, StalpersJA,RobertV,G (2004) Stegehuis Dennis RWG (1978) Dearness J (1926) Newandnoteworthyfungi-IV. Crous PW, Schoch CL,HydeKD,Wood AR, GueidanC, et al. Crous PW, Wingfield MJ, Mansilla JP, Alfenas AC, Groenewald JZ Groenewald JP, AC, Alfenas Mansilla MJ, Wingfield PW, Crous PW,Crous VerkleyRA JZ,Samson GJM,Groenewald (2009b) (eds) DNA sequences.Mycologia Rhytismatales of Helotiales RC, KohnLM (2001).Phylogenetics Hamelin America. Mycotaxon63:261–268. TypographiaAcademica. MacLachlan &Stewart. Pyrenomyceten. coniferous hosts.Sydowia Mauritius. Academica. Mauritius. Bryologie, Lichénologie by 18SrDNAinferred analysis. sequence G. Bonnier. plantations. 12: 1–82. 3912–4284. Phacidiaceae Smith (HookerWJ),Vol. 5(2).]London:Longman I. inoperculés discomycètes trimestriel delaSociétéMycologiqueFrance les chez l’apothécie Office, Washington, DC:USDA. of Printing No. 521.]USGovernment Handbook Agriculture 11 century. StudiesinMycology MycoBank: anonlineinitiativeto launch mycologyintothe21st 236–255. 55: 99–131. in Mycology on Eucalyptus.II.Studies their anamorphsoccurring Mycology in theCapnodiales lineages (2009a) Phylogenetic (2006) Phylogenetic reassessment of (2006) Phylogenetic CBS-KNAW FungalBiodiversityCentre. Fungal Biodiversity (1): 1–434. 64:17–47. Journal of the ArnoldArboretum 11 of Journal Summa VegetabilumScandinaviae Myconet 11 andrelatedanamorphs. based onpartial smallsubunitnuclearribosomal Scottish Cryptogamic Flora. Vol.Scottish Cryptogamic 6.Edinburgh: British Ascomycetes. 2 Beiträge . [CBS Laboratory ManualSeries1.] . [CBSLaboratory Utrecht: 19:123–136. : 1–51.

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