Polish Botanical Journal 55(2): 359–371, 2010 CLADOSPORIUM EPICHLOËS, A RARE EUROPEAN FUNGUS, WITH NOTES ON OTHER FUNGICOLOUS SPECIES* MAŁGORZATA RUSZKIEWICZ-MICHALSKA Abstract. Cladosporium epichloës Lobik, associated with Epichloë typhina (Pers.) Tul. & C. Tul. and known from three records worldwide, is reported from Poland for the first time. The morphology and distribution data of this species as well as the first records of Cladosporium uredinicola Speg. and Phoma glomerata (Corda) Wollenw. & Hochapfel as parasites of powdery mildews in Poland are presented. Information concerning specimens of five other hyperparasitic species deposited in Herbarium Universitatis Lodziensis (LOD) is provided. Key words: Cladosporium, Phoma glomerata, Sphaerellopsis filum, Ampelomyces, Tuberculina, hyperparasite, fungicolous fungi, chorology, Poland Małgorzata Ruszkiewicz-Michalska, Department of Mycology, University of Łódź, Banacha 12/16, 90-237 Łódź, Poland; e-mail: mrusz@biol.uni.lodz.pl INTRODUCTION The term ‘fungicolous fungi’ covers species that occur on other fungi as parasites, commensals or saprobionts (Kirk et al. 2008). The nature of the interfungal relationships is not always clear, especially for intimate mycoparasitic interactions (Jeffries & Young 1994). Opinions on the type of relations of particular species with their mycohosts vary considerably. The status of the mycoparasite Ampelomyces quisqualis Ces., considered to be an intracellular necrotroph vs. biotroph (Jeffries & Young 1994), is the best example. It cannot be ruled out that during infection the nature of the contact can be gradually transformed from nearly biotrophic to necrotrophic without the production of any toxins (Kiss 2003 and literature cited therein). Similarly, some recent studies indicate that species of the genus Tuberculina Tode ex Sacc., parasitizing rust fungi, are fusion biotrophs (Lutz et al. 2004), but this has not been confirmed by other researchers (e.g., Bartkowska 2007b). Sphaerellopsis filum (Biv.) B. Sutton is a biotrophic mycoparasite which remains in close trophic contact with rust * This paper is dedicated to Professor Tomasz Majewski on the occasion of his 70th birthday. spores via appresorium-like structures (Płachecka 2005 and literature cited therein). A reliable estimate of the number of fungicolous species is not available (Kirk et al. 2008). Hawksworth (1981), however, traced 1100 anamorphic species recorded on other species of fungi. Fungicolous fungi include members of all higher taxa of true fungi (Chytridiomycota, Zygomycota, Ascomycota including anamorphic taxa, Basidiomycota) and chromistan fungal analogues (Hyphochytriomycota, Oomycota). This taxonomic diversity is also reflected in Polish records of fungi belonging to this ecological group (Mułenko et al. 2008), with most representatives belonging to the ascomycetes [e.g., the genera Hypocrea Fr., Hypomyces (Fr.) Tul. & C. Tul., Nectria (Fr.) Fr.], zygomycetes (mostly belonging to Mortierellales, Mucorales and Zoopagales) and anamorphic fungi. Lichenicolous fungi, recently examined in a study by Czyżewska and Kukwa (2009), are a separate ecological group, relatively numerous in species. Fungi occupied by fungicolous species represent a broad taxonomic spectrum comprising representatives of all taxa of higher ranks belonging to the kingdom Fungi, as well as chromistan and 360 POLISH BOTANICAL JOURNAL 55(2). 2010 protozoan fungal analogues (e.g., myxomycetes) (Hansford 1946; Ellis & Ellis 1998; Samuels et al. 2006; Kirk et al. 2008). Many species parasitize fungi closely related to them; for example, Tuberculina species infect members of the Pucciniales (cf. Lutz et al. 2004). This is also frequently observed among representatives of Oomycota, Chytridiomycota and Zygomycota (Jeffries & Young 1994). For instance, Absidia parricida Renner & Muscat ex Hesselt. & J. J. Ellis, Chaetocladium jonesii (Berk. & Broome) Fresen. and Parasitella parasitica (Bainier) Syd. were recorded on other Mucorales species in Poland (Mułenko 2008). Rust fungi infected by Eudarluca caricis (Biv.) O. E. Erikss. and species belonging to Cladosporium Link, Tuberculina Tode ex Sacc. and Ramularia Unger are the most frequently observed fungi hosting other fungi in Poland (Mułenko et al. 2008). Specimens of the latter taxon [Ramularia coleosporii Sacc., R. uredinis (Voss) Sacc., R. uredinearum Hulea] were recently taxonomi- cally verified and are discussed in a monograph by Wołczańska (2005). The parasitism of Ramularia species on rust spores was recently confirmed by Bartkowska (2007a). Hosts of fungicolous fungi often recorded in Poland also include Erysiphales gen. div. and other species of ascomycetes such as Diatrypella favacea (Fr.) Ces. & De Not., Diatrype spp. and Hypoxylon spp., as well as agarics, boletes and polypores (Mułenko et al. 2008). Other species of fungi occurring on parasitic micromycetes, most probably as saprobes, have been observed sporadically (see Table 1). The majority of these records cannot be verified as they are reported in older studies, mostly phytopathological, which are rarely documented by herbarium specimens. The status of a fungus recorded by Domański et al. (1970) on Plasmopara densa (Rabenh.) J. Schroet. (a parasite of Euphrasia sp.), which most probably belonged to Trichothecium plasmoparae Viala, is also unexplained. Table 1. Survey of micromycetes reported in association with basidiomycetous plant parasites in Poland. Mycohost species Fungicolous species Source of data Gymnosporangium sp. Gloeosporium roesteliicola Bubák & Serebrian. Jankowska-Barbacka 1931 Puccinia punctata Link Fusarium uredinicola Jul. Müll. Dominik 1936 Phragmidium rubi-idaei (DC.) P. Karst. Fusarium avenaceum (Fr.) Sacc. Mułenko et al. 2008 Puccinia caricina DC. Hymenula spermogoniopsis (Jul. Müll.) Wollenw. Wollenweber 1916; Mułenko 1996 Puccinia coronata Corda Fusarium avenaceum (Fr.) Sacc. Mułenko et al. 2008 Puccinia sessilis W. G. Schneid. Fusarium avenaceum (Fr.) Sacc., Volutella fusariispora Dominik 1934; Mułenko et al. Dominik (on aecia) 2008 Uromyces fabae (Pers.) de Bary Fusarium graminearum Schwabe, Trichothecium roseum Pruszyńska-Gondek 1974, 1976 (Pers.) Link Ustilago avenae (Pers.) Rostr. Verticillium albo-atrum Reinke & Berthold Ustilago nuda (C. N. Jensen) Kellerm. & Swingle Alternaria tenuis Nees, Botrytis cinerea Pers., Fusarium heterosporum Nees & T. Nees, Trichothecium roPielka 1963 seum (Pers.) Link, Verticillium albo-atrum Reinke & Berthold Ustilago tritici (Pers.) Rostr. Botrytis cinerea Pers., Fusarium culmorum (W. G. Sm.) Sacc., F. javanicum Koord. var. radicicola Wollenw., F. heterosporum Nees & T. Nees, Nigrospora oryzae Pielka 1963 (Berk. & Broome) Petch, Trichothecium roseum (Pers.) Link, Verticillium albo-atrum Reinke & Berthold Exobasidium vaccinii (Fuckel) Woronin Fusarium graminum Corda Pielka 1963 Moesz 1926 (fungicolous species name with a question mark) M. RUSZKIEWICZ-MICHALSKA: CLADOSPORIUM EPICHLOËS, A RARE EUROPEAN FUNGUS CLADOSPORIUM SPECIES: HOST RANGE AND SPECIALIZATION Twenty-six species of the genus Cladosporium Link s.l. have been described as confined to fungi (Heuchert et al. 2005). As well as a few polyphagous saprobionts occasionally colonizing thalli of different species [e.g., C. herbarum (Pers.) Link, C. tenuissimum Cooke, C. cladosporioides (Fresen.) G. A. de Vries], they also include Cladosporium s.str. associated with specific taxa of fungi. Some, such as C. lycoperdinum Cooke, C. epimyces Cooke and C. episclerotiale Bubák, colonize fruitbodies of macromycetes (Agaricales, Boletales, Polyporales, Russulales). Cladosporium episclerotiale is also known from sclerotia of Monilinia laxa (Aderh. & Ruhland) Honey. Cladosporium phyllophilum McAlpine, a parasite of Taphrinales, and C. exobasidii Jaap, limited to Exobasidium Woronin species, exhibit a narrower host spectrum. Some species associated with rust fungi are confined to aecia (C. aecidiicola Thüm., C. gallicola B. Sutton), while others occur on different stages of a variety of genera belonging to Pucciniales (e.g., C. uredinicola Speg.). The latter species also includes specimens morphologically indistinguishable from C. uredinicola known to occur on two Peronospora Corda species from India, Erysiphe euonymi-japonici (Vienn.-Bourg.) U. Braun & S. Takam. from Iran, and Phyllactinia angulata (E. S. Salmon) S. Blumer and P. guttata (Wallr.) Lév. from the U.S.A. (Heuchert et al. 2005; Dugan & Glawe 2006). Finally, C. taphrinae Bubák, C. gerwasiae Heuchert, U. Braun & K. Schub. and C. epichloës Lobik are treated as stenotopic species in the current taxonomic approach of Heuchert et al. (2005). Five species of the genus Cladosporium colonizing other fungi have been recorded in Poland (Heuchert et al. 2005; Ruszkiewicz-Michalska & Mułenko 2008). These are Cladosporium aecidicola, C. uredinicola, C. phyllophilum on Taphrina pruni (Fuckel) Tul., C. fuligineum Bonord. and C. epimyces. Although the name is at present a synonym of Cladosporium herbarum, the taxonomic position of specimens reported in Poland as C. fuligineum is uncertain because the 361 specimens most probably represented other species of Cladosporium, especially the records on Exobasidium vaccinii (Fuckel) Woronin. These collections cannot be verified as the exsiccata documenting them are missing. Finally, the specimen of C. epimyces (on Lactarius sp., HBG herbarium) from Prószków near Opole was verified by Heuchert et al. (2005). MATERIALS AND METHODS The present list is based on my unpublished and published data (Ruszkiewicz-Michalska & Michalski 2005; Ruszkiewicz-Michalska 2006) and specimens deposited in the fungal collection of Herbarium Universitatis Lodziensis (LOD) in the subcollection of Parasitic micromycetes (labelled PF). The latter were collected mostly by graduate students conducting research for theses under my supervision. The majority of exsiccata were collected in central Poland, covering Łódź Province and a few sites in Silesia Province (Wyżyna Częstochowska upland, vicinity of Pszczyna). A single specimen was collected in the Bory Tucholskie forest (Fig. 1). Papers by Heuchert et al. (2005) and Ellis and Ellis (1998) were used for determination of fungicolous fungi. Identification of their host species was based on monographs by Majewski (1977, 1979) and Braun (1995). Fig. 1. Sampling sites of fungicolous micromycetes in Poland. 1 – Wierzchlas Reserve, 2 – Łuszczanów, 3 – Łódź, 4 – Przedbórz, 5 – Złoty Potok, 6 – Ligota Dolna Reserve, 7 – Wola. 362 POLISH BOTANICAL JOURNAL 55(2). 2010 Associated plant species are identified according to Rutkowski (1998). Nomenclature follows Mirek et al. (2002) for plants and Mułenko et al. (2008) for fungi. The specimens were studied with a stereomicroscope (Nikon SMZ-10A) and a light microscope (Nikon E400) with phase contrast, using a 1000× oil immersion objective. To have a representative size range, 20 pycnidia, conidiophores and conidia mounted in water were measured in each collection. Photographs were taken with a Nikon DS-L1 digital camera. RESULTS AND DISCUSSION Seven species of fungicolous fungi associated with parasitic micromycetes are listed. Cladosporium epichloës, reported from Poland for the first time, is described and illustrated. Host species, the distribution in Poland and general notes are given for each species. The specimens from LOD given in previous papers are annotated with the reference data. constricted at septa, pale brown, almost smooth, walls conspicuously thickened, apex rounded, with up to 3 hila, base truncate to convex, hila protuberant, thickened, darkened, 1–2 μm in diameter; microcyclic conidiogenesis observed sporadically. This description is in accordance with morphological characteristics given by Heuchert et al. (2005). SPECIMEN EXAMINED. POLAND. On Epichloë typhina (Pers.) Tul. & C. Tul. (on Poa compressa L.): WZNIESIENIA POŁUDNIOWOMAZOWIECKIE HILLS: Łódź, Las Łagiewnicki forest complex, forest district 11, grassy edge of deciduous forest (Tilio-Carpinetum typicum), 8 June 2007, leg. M. Ruszkiewicz-Michalska (LOD 3088 PF). Fig. 2A–C HOST RANGE AND DISTRIBUTION. According to Heuchert et al. (2005) the species is known on Epichloë typhina from three European countries: southwestern Russia (holotype, E. typhina hosted by Bromus inermis Leyss.), central Germany (Dactylis polygama Horv.) and eastern Austria (host of E. typhina not specified). Colonies olivaceous to brown, diffuse to dense, velvety. Mycelium immersed and external, superficial hyphae sparse, creeping, branched, 2.5–5.0 μm wide, septate, cells irregularly swollen (up to 7 μm wide), pale brown, smooth. True stromata lacking. Conidiophores solitary or in loose groups, arising from swollen cells (up to 7–8 μm in diameter), erect, straight or sometimes curved, subcylindrical, geniculate-sinuous, rarely branched, (20–)30–45 × 3–5 μm, slightly swollen at base, 0–3 septate, not constricted at septa, pale to medium brown, paler towards apex, smooth, rarely with subnodulose intercalary swellings (up to 5 μm wide) often with conidiogenous loci. Conidiogenous cells integrated, terminal and rarely intercalary, 8–30 μm long, proliferation sympodial, with (1–)2–7 conspicuous, protuberant, thickened, darkened-refractive conidiogenous loci (1–2 μm in diameter) sometimes situated on small unilateral swellings resembling shoulders. Conidia mainly solitary, rarely in branched chains, straight, polymorphous, ellipsoid, obovoid, fusiform, subcylindrical, (5–)9–22 × 4–7 μm, 0–2(–5)-septate, not NOTES. Cladosporium epichloës is barely distinguishable from C. aecidiicola morphologically. They are tentatively treated as separate taxa (Heuchert et al. 2005), as distinct ecology and host specialization are evident in these two species. Additional studies, including molecular and inoculation experiments, are needed to resolve their taxonomy. C. epichloës also resembles the saprobic C. herbarum. The main character distinguishing them is the presence of multilateral nodes of conidiophores in the latter (swellings 7–9 μm in diam.; compare illustration in Schubert et al. 2007: figs 17, 18b) which are absent in the former. In addition, C. epichloës forms characteristic geniculate conidiophores with small unilateral shoulders (Heuchert et al. 2005). As Epichloë typhina is a common pathogen of many of the world’s graminicolous species (including cereal crops), its parasites are of great importance as prospective biocontrol agents. Most of the species observed on E. typhina are known from a limited number of localities such as Radulidium epichloës (Ellis & Dearn.) Arzanlou, W. Gams & Crous (Arzanlou et al. 2007), Phyllosticta epi- Cladosporium epichloës Lobik Bolezni Rast. 17(3–4): 189. 1928. 363 M. RUSZKIEWICZ-MICHALSKA: CLADOSPORIUM EPICHLOËS, A RARE EUROPEAN FUNGUS B A t D C Fig. 2. Fungicolous Cladosporium species. A–C – Cladosporium epichloës (A – developing ascoma of Epichloë typhina covered by conidiophores of the parasite, B & C – conidiophores and conidia); D – C. uredinicola [conidiophores and conidia on telial column (t) of Cronartium flaccidum]. Scale bars: A, C, D = 25 μm, B = 10 μm. chloës Thirum. (Aa & Vanev 2002), Cladosporium epichloës (Heuchert et al. 2005), and Dicyma pulvinata (Berk. & M. A. Curtis) Arx recorded on E. typhina for the first time only recently (Alderman et al. 2010). Bionectria epichloë (Speg.) Schroers, whose presence in the anamorph stage (Clonostachys epichloë Schroers) was recently confirmed by Kirschner (2006) in Europe, occurs worldwide. Only Alternaria sp. was observed on the stroma of Epichloë typhina in Poland (Mułenko et al. 2008). Cladosporium aecidiicola Thümen Mycoth. univ., Cent. IV, No. 373. 1876. SPECIMEN EXAMINED. POLAND. On aecia of Puccinia coronata Corda (on Rhamnus cathartica L.): WYŻYNA PRZEDBORSKA UPLAND: Murawy Dobromierskie Reserve near Przedbórz, edge of xerothermic grassland, 14 Sept. 2009, leg. M. Ruszkiewicz-Michalska (LOD 3090 PF). HOST RANGE AND DISTRIBUTION. This species is known to parasitize aecial sori of numerous rust 364 POLISH BOTANICAL JOURNAL 55(2). 2010 species worldwide (Heuchert et al. 2005; Farr & Rossman 2010). Recently it was reported for the first time also on spermogonia (Blanz & Braun 2008). The species is very likely to be common in Poland although the published records comprise only the northern and southern parts of the country (Ruszkiewicz-Michalska & Mułenko 2008). It infected seven host species of the genera Coleosporium Lév., Puccinia Pers. and Uromyces (Link) Unger. See also notes on C. uredinicola. NOTES. Like C. epichloës, C. aecidiicola is morphologically similar to the common saprobic C. herbarum, which differs in having ovoid-ellipsoid conidia that are not attenuated toward the base as observed in the former species. In addition, C. herbarum forms characteristic intercalary swellings round about the conidiophore, which are not present in C. aecidiicola (Heuchert et al. 2005). Cladosporium uredinicola Spegazzini Fig. 2D Anales Mus. Nac. Hist. Nat. Buenos Aires 23: 122–123. 1912. of numerous genera of the rust families Phragmidiaceae, Pucciniaceae and Uropyxidiaceae; it has not been reported on Ochropsora species (Heuchert et al. 2005; Farr & Rossman 2010). The specimen of C. uredinicola on Ochropsora ariae (LOD 3113) concerns a rust species very rare on this host plant in Poland (Majewski 1977; Majewski & Ruszkiewicz-Michalska 2008) and in the world (Farr & Rossman 2010). Cladosporium uredinicola on Phyllactinia guttata is the first Polish report of this species on a powdery mildew. NOTES. A parasite of Cronartium ribicola J. C. Fisher assigned by Madej (1965) to C. aecidicola most probably belongs to C. uredinicola. This record is provisionally included (exsiccatum not available) in C. uredinicola due to its occurrence on telial sori, as C. aecidicola is confined to the aecial stage of rusts, similarly to C. gallicola associated with members of the Cronartiaceae. Although C. uredinicola was already reported in association with telial columns of Cronartium Fr. species (Morgan-Jones & McKemy 1990), it cannot be ruled out that Madej’s record (1965) represents the recently described C. gerwasiae, which is at present known only from Gerwasia sp. from the Phragmidiaceae (Heuchert et al. 2005). SPECIMENS EXAMINED. POLAND. On uredinia of Ochropsora ariae (Fuckel) Ramsb. (on Pyrus communis L.): WZNIESIENIA POŁUDNIOWOMAZOWIECKIE HILLS: Łódź, Las Łagiewnicki Reserve, deciduous forest (Potentillo albae-Quercetum), roadside, 30 Sept. 2007, leg. K. Brózio (LOD 3113 PF); on telia of Cronartium flaccidum (Alb. & Schwein.) G. Winter (on Vincetoxicum hirundinaria Medik.): WZNIESIENIA POŁUDNIOWOMAZOWIECKIE HILLS: Łódź, Botanical Garden, section of healing plants and plants grown for industrial purposes, 4 Sept. 2004, leg. E. Połeć (LOD 3089 PF); WYŻYNA ŚLĄSKA UPLAND: Ligota Dolna Nature Reserve near Opole, Kamienna Góra hill, xerothermic meadow (Adonido-Brachypodietum), top of hill, 16 Aug. 2000, leg. E. Moliszewska [as Cladosporium aecidiicola Thüm. (in Moliszewska 2008); voucher stored in herbarium of the Department of Biotechnology and Molecular Biology, University of Opole, and LOD 3130 PF]; on chasmothecia of Phyllactinia fraxini (DC.) Fuss (on Fraxinus excelsior L.): WZNIESIENIA POŁUDNIOWOMAZOWIECKIE HILLS: Łódź, by Jaracza Street, 10 Sept. 2010, leg. M. Ruszkiewicz-Michalska (LOD 3129 PF). SPECIMENS EXAMINED. POLAND. On aecia of Puccinia coronata Corda (on Frangula alnus Mill.): KOTLINA OŚWIĘCIMSKA BASIN: Wola village near Pszczyna, edge of coniferous forest (Molinio-Pinetum), 11 June 2007, leg. A. Myszka (LOD 2828 PF); on Puccinia poarum E. Nielsen (on Tussilago farfara L.): KOTLINA OŚWIĘCIMSKA BASIN: Wola village near Pszczyna, edge of coniferous forest (Molinio-Pinetum), 11 June 2007, leg. A. Myszka (LOD 2929 PF); on Puccinia sessilis W. G. Schneid. [on Polygonatum odoratum (L.) Mill.]: Bory Tucholskie forest: Cisy Staropolskie Reserve in Wierzchlas near Tuchola, deciduous forest (TilioCarpinetum), 26 June 2006, leg. M. Ławrynowicz (LOD 3094 PF). HOST RANGE AND DISTRIBUTION. This species is known worldwide on aecia, uredinia and telia HOST RANGE AND DISTRIBUTION. Three commonly recognized Tuberculina species [T. maxima Tuberculina persicina (Ditmar) Saccardo Fungi italica autogr. del. 17–28: tab. 964. 1881. M. RUSZKIEWICZ-MICHALSKA: CLADOSPORIUM EPICHLOËS, A RARE EUROPEAN FUNGUS Rostr., T. persicina (Ditm.) Sacc. and T. sbrozzii Cavara & Sacc.] parasitize more than 150 species of 15 genera of Pucciniales worldwide (Lutz et al. 2004). In Poland, two of the three species were reported on aecial sori of rusts: T. maxima on Cronartium flaccidum and T. persicina on 12 species belonging to six genera (Mułenko et al. 2008). The latter species was wrongly reported on Coleosporium tussilaginis (Pers.) Berk. also present in the specimen from the Wyżyna Częstochowska upland (Ruszkiewicz-Michalska 2006). My verification of the specimen (LOD 1002 PF) revealed the parasite associated with aecia of Puccinia poarum. The taxonomy of T. cracoviae Pielka and T. ustilaginum Pielka, two Tuberculina species described by Pielka (1963) from Poland parasitizing smut fungi [Ustilago avenae (Pers.) Rostr., U. nuda (Jens.) Kellerm. & Swingle and U. tritici (Pers.) Rostr.], is unclear. The species probably do not belong to Tuberculina s.str., as in the opinion of Lutz et al. (2004) the genus should be restricted to rust species. Parasites of smut fungi are barely noted in the available professional literature. Among the species reported as parasites of smuts, Sphaerellopsis filum and Pythium vexans de Bary are listed (Jeffries & Young 1994). SEM observations of an unidentified ascomycete parasitizing spores of Vankya vaillantii (Tul. & C. Tul.) Ershad were given by Protzenko (1989). The ability of Stephanoma phaeosporum Butler & McCain to parasitize Ustilago avenae (Pers.) Rostr. and 365 U. maydis (DC.) Corda in culture was also reported (Rakvidhyasastra & Butler 1973). NOTES. Based on molecular and ultrastructural studies, Lutz et al. (2004) recently showed that members of the genus are closely related to their hosts (rust species), sharing with them features including the uredinalean architecture of septal pores. Lutz et al. (2004) and Bauer et al. (2004) also reported the lack of haustoria and other intracellular structures of T. persicina, and its interaction with the host via large fusion pores in the frame of which Tuberculina nuclei and other organelles are horizontally transferred to the rust cells. In contrast, observations by Bartkowska (2007b) do not confirm such fusion of host and parasite via large pores. She observed direct penetration of aeciospores by Tuberculina hyphae (presumably vegetative) and invasion of plant tissue by the hyperparasite causing rapid destruction of plant cells infected with the rust. Ampelomyces quisqualis Cesati s.l. Fig. 3A Bot. Ztg. 10: 301. 1852. SPECIMENS EXAMINED. POLAND. On Erysiphe artemisiae Grev. [= Golovinomyces artemisiae (Grev.) V. P. Heluta] (on Artemisia vulgaris L.): WZNIESIENIA POŁUDNIOWOMAZOWIECKIE HILLS: Łódź city, Bolesław Chrobry Municipal Park, children’s playground, 9 Oct. 2006, leg. D. Papierz (LOD 3049 PF); on Erysiphe cichoracearum DC. var. cichoracearum [= Golovinomyces cp A B Fig. 3. Pycnidial fungicolous fungi. A – Ampelomyces quisqualis [pycnidium in conidiophore (cp) of Sphaerotheca spireae]; B – Phoma glomerata (globose pycnidium). Scale bars = 25 μm. 366 cichoracearum (DC.) V. P. Heluta var. cichoracearum] (on Centaurea scabiosa L.): WYŻYNA PRZEDBORSKA UPLAND: Murawy Dobromierskie Reserve near Przedbórz, xerothermic grassland, 14 Sept. 2005, leg. M. Ruszkiewicz-Michalska (LOD 3116 PF); (on Tanacetum vulgare L.): WZNIESIENIA POŁUDNIOWOMAZOWIECKIE HILLS: Łódź, Bolesław Chrobry Municipal Park, children’s playground, 16 Sept. 2006, leg. D. Papierz (LOD 3050 PF); on Erysiphe cruciferarum Opiz ex Junell [on Sisymbrium officinale (L.) Scop.]: WZNIESIENIA POŁUDNIOWOMAZOWIECKIE HILLS: Łódź, Bolesław Chrobry Municipal Park, children’s playground, 12 Oct. 2006, leg. D. Papierz (LOD 3048 PF); on Erysiphe heraclei DC. [on Petroselinum crispum (Mill.) A. W. Hill]: NIZINA ŚRODKOWOMAZOWIECKA LOWLAND: Łuszczanów village near Pacyna, vegetable garden, 17 Sept. 2004, leg. W. Kaczmarek (LOD 3115); on Erysiphe sordida (L.) Junell [= Golovinomyces sordidus (L. Junell) V. P. Heluta] (on Plantago intermedia Gilib.): WZNIESIENIA POŁUDNIOWOMAZOWIECKIE HILLS: Łódź, Las Łagiewnicki forest complex, deciduous forest, 21 Oct. 2007, leg. M. Ruszkiewicz-Michalska (LOD 3117 PF); (on Plantago major L.): NIZINA ŚRODKOWOMAZOWIECKA LOWLAND: Łuszczanów village near Pacyna, vegetable garden, weed, 17 Sept. 2004, leg. W. Kaczmarek, (LOD 3114 PF); KOTLINA OŚWIĘCIMSKA BASIN: Wola village near Pszczyna, boggy meadow (community of Molinion caeruleae alliance), 28 Aug. 2006, leg. A. Myszka (LOD 2780 PF); on Microsphaera trifolii (Grev.) U. Braun [= Erysiphe trifoliorum (Wallr.) U. Braun] (on Melilotus alba Medik.): WZNIESIENIA POŁUDNIOWOMAZOWIECKIE HILLS: Łódź, Las Łagiewnicki forest complex, rest stop/parking lot, lawn, 21 Oct. 2007, leg. M. Ruszkiewicz-Michalska (LOD 3118 PF); on Podosphaera tridactyla (Wallr.) de Bary [on Padus serotina (Ehrh.) Borkh.]: KOTLINA OŚWIĘCIMSKA BASIN: Wola village near Pszczyna, edge of coniferous forest (MolinioPinetum), 11 June 2007, leg. A. Myszka (LOD 2825 PF); WZNIESIENIA POŁUDNIOWOMAZOWIECKIE HILLS: Łódź, Las Łagiewnicki Reserve, deciduous forest (Potentillo albae-Quercetum), 19 Aug. 2007, leg. K. Brózio (LOD 3119 PF); deciduous forest (Tilio-Carpinetum calamagrostietosum), roadside, 8 Sept. 2007, leg. K. Brózio (LOD 3120 PF); same locality and collector, 30 Sept. 2007 (LOD 3121 PF); Las Łagiewnicki forest complex, deciduous forest (Tilio-Carpinetum), roadside, 30 Sept. 2007, leg. A. Kuchnik (LOD 3122 PF); same locality, deciduous forest (Calamagrostio-Quercetum typicum), roadside, 22 July 2007, leg. A. Kuchnik (LOD 3123 PF); on Sphaerotheca balsaminae (Wallr.) Kari (on Impatiens noli-tangere L.): KOTLINA OŚWIĘCIMSKA BASIN: Wola village near Pszczyna, edge of coniferous forest (Molinio- POLISH BOTANICAL JOURNAL 55(2). 2010 Pinetum), 11 July 2006, leg. A. Myszka (LOD 2777-2779 PF); on Sphaerotheca fusca (Fr.) Blumer [= Podosphaera fusca (Fr.) U. Braun & Shishkoff ] (on Solidago canadensis L.): WZNIESIENIA POŁUDNIOWOMAZOWIECKIE HILLS: Łódź, Las Łagiewnicki forest complex, deciduous forest (Calamagrostio-Quercetum typicum), roadside, 28 July 2007, leg. A. Kuchnik (LOD 3124 PF); on Sphaerotheca spiraeae Sawada [= Podosphaera filipendulae (Z. Y. Zhao) T. Z. Liu & U. Braun] [on Filipendula ulmaria (L.) Maxim.]: WYŻYNA CZĘSTOCHOWSKA UPLAND, Parkowe Reserve near Złoty Potok, forest district no. 270, shaded thickets at Wiercica stream, 14 July 1998, leg. M. Ruszkiewicz (LOD 1799 PF). Published reports of this species from Central Poland (Ruszkiewicz-Michalska & Michalski 2005; Ruszkiewicz-Michalska 2006) are based on the following specimens from LOD (PF): nos 784, 1227 (Erysiphe cichoracearum), no. 816 (E. cruciferarum), no. 392 (E. galii Blumer) [= Neoerysiphe galii (S. Blumer) U. Braun], no. 791 (E. heraclei DC.), no. 762 (E. magnicellulata U. Braun var. magnicellulata) [= Golovinomyces magnicellulatus (U. Braun) V. P. Heluta], no. 515 (E. orontii Castagne) [= G. orontii (Castagne) V. P. Heluta], nos 140, 141, 150, 151, 1317, 1785, 1797, 2553 (E. sordida), no. 1900 (Microsphaera alphitoides Griffon & Maubl. var. alphitoides) [= Erysiphe alphitoides (Griffon & Maubl.) U. Braun & S. Takam.], no. 803 [M. berberidis (DC.) Lév. var. berberidis] [= E. berberidis (DC.) U. Braun & S. Takam.], no. 676 [Phyllactinia guttata (Wallr.) Lév.], no. 780 [Sawadaea bicornis (Wallr.) Homma], nos 710, 2104 [Sphaerotheca aphanis (Wallr.) U. Braun var. aphanis] [= Podosphaera aphanis (Wallr.) U. Braun & S. Takam.], nos 896, 916, 1346 [Sphaerotheca ferruginea (Schltdl. ex Fr.) L. Junell] [= P. ferruginea (Schltdl. ex Fr.) U. Braun & S. Takam.], no. 1998 [Sphaerotheca fusca (Fr.) Blumer emend. U. Braun], nos 602, 620 (Sphaerotheca spiraeae Sawada). HOST RANGE AND DISTRIBUTION. The name A. quisqualis is traditionally applied to all pycnidial intracellular hyperparasites of powdery mildews although more than 40 species are validly described in the genus in the older literature. A. quisqualis s.l. infects ca 66 species belonging to nine genera, including the rarely attacked Blumeria graminis (DC.) Speer, worldwide (Kiss 1998; Szentiványi & Kiss 2003). Linnemann (1968) reported also A. quisqualis as a parasite of Mucorales species. Recent studies on the taxonomy of the species revealed that it is an assemblage of several M. RUSZKIEWICZ-MICHALSKA: CLADOSPORIUM EPICHLOËS, A RARE EUROPEAN FUNGUS distinct lineages rather than a single species (Kiss & Nakasone 1998; Szentiványi et al. 2005). Some groups of A. quisqualis isolates show differences in ITS rDNA and actine gene sequences reflecting their association with certain taxa of mycohosts (Szentiványi et al. 2005; Park et al. 2010). According to Mułenko et al. (2008; erysiphacean nomenclature following Braun 1995) A. quisqualis s.l. commonly occurring in Poland was noted on species of Erysiphe (21 taxa), Microsphaera (6), Oidium (1), Podosphaera (2), Sawadaea (1), Sphaerotheca (10) and Uncinula (2). The only record of the parasite on Phyllactinia concerns P. mali (Duby) U. Braun (Czerniawska 2005). The list of Polish hosts of A. quisqualis is supplemented here with four new species: Microsphaera berberidis, Phyllactinia guttata, Sphaerotheca balsaminae and S. spiraeae. Ampelomyces quisqualis often accompanies alien expansive species of powdery mildews such as Erysiphe elevata (Burrill) U. Braun & S. Takam., E. flexuosa (Peck) U. Braun & S. Takam. and E. platani (Howe) U. Braun & S. Takam. (Pastirčáková 2007) and taxa newly spreading on a host [e.g., Phyllactinia guttata on Padus serotina (Ehrh.) Borkh.; Ruszkiewicz-Michalska & Michalski 2005]. NOTES. Some of the Ampelomyces species [A. heraclei (Dejeva) Rudakov, A. humuli (Fautrey) Rudakov and A. quercinus (Syd.) Rudakov] isolated from powdery mildews (Microsphaera and Sphaerotheca) and a downy mildew fungus [Plasmopara viticola (Berk. & M. A. Curtis) Berl. & De Toni] were recognized as closely related to Phoma glomerata (Corda) Wollenw. & Hochapfel (Kiss & Nakasone 1998; Sullivan & White 2000). One of those fast-growing Ampelomyces species (A. quercinus) was recently reallocated to Phoma fungicola Aveskamp, Gruyter & Verkley based on molecular studies (Aveskamp et al. 2010). Phoma glomerata (Corda) Wollenweber & Hochapfel Fig. 3B Z. ParasitKde 3(5): 592. 1936. SPECIMENS EXAMINED. POLAND. On Phyllactinia fraxini (DC.) Fuss (on Fraxinus excelsior L.): WZNIESIENIA POŁUDNIOWOMAZOWIECKIE HILLS: Łódź, 367 by Jaracza Str., 12 Oct. 2008, leg. M. RuszkiewiczMichalska (LOD 3091 PF); same locality, host and collector, 10 Sept. 2010 (LOD 3128 PF). HOST RANGE AND DISTRIBUTION. The species is reported from Poland as a parasite of powdery mildews for the first time although it is known to occur as a saprotroph on different substrates, including plant tissues (Mułenko et al. 2008). It is very likely that either P. glomerata or another fungicolous Phoma species on Phyllactinia mali was also observed by Czerniawska (2005) in Gorzów Wielkopolski (western Poland), as suggested by the notes on the size of spores produced by a parasite of Phyllactinia (Czerniawska 2005). The size range considerably exceeds values reported for A. quisqualis (Sutton 1998) and the range observed on other hosts in the study by Czerniawska (2005), but it is within the range reported for P. glomerata (Boerema et al. 2004). The presence of P. glomerata (probably in a combined infection with Ampelomyces quisqualis) on the mycelium of P. mali also seems to be confirmed by the photographs (Czerniawska 2005: Fig. 1a, d), which show globose pycnidia with a multi-layered wall resembling pycnidia of Phoma. NOTES. Phoma glomerata is a cosmopolitan, ubiquitous species associated with a wide range of substrates, and a secondary invader, probably feeding on fungal saprotrophs and parasites of diseased tissues of plants (White & Morgan-Jones 1987; Sullivan & White 2000). P. glomerata is frequently confused with Ampelomyces quisqualis (Sullivan & White 2000), but the two species are distinguishable on the basis of pycnidial morphology (pycnidia sessile and formed mainly on the leaf surface vs. stipitate pycnidia formed mainly within the host’s hyphae and conidiophores). These two species also differ by their growth rates in culture: average growth on PDA for P. glomerata isolates is 8±1 mm/day and for A. quisqualis it is 0.8±0.1 mm/day (Sullivan & White 2000). Occasionally the conidia of P. glomerata are 1-septate (White & Morgan-Jones 1987), as observed in a single pycnidium in one of the specimens mentioned above (LOD 3091 PF). Additionally, in culture P. glomerata produces generally multicellular 368 alternarioid dictyochlamydospores formed in branched or unbranched chains of 2–20 elements (Boerema et al. 2004). Sphaerellopsis filum (de Bivona-Bernardi) B. Sutton Mycol. Pap. 141: 196. 1977. SPECIMENS EXAMINED. POLAND. On telia of Puccinia chaerophylli Purton [on Torilis japonica (Houtt.) DC.]: WZNIESIENIA POŁUDNIOWOMAZOWIECKIE HILLS: Łódź city, Las Łagiewnicki forest complex, deciduous forest (Tilio-Carpinetum calamagrostietosum), roadside, 25 Aug. 2007, leg. A. Kuchnik (LOD 3125 PF); on uredinia of Puccinia menthae Pers. (on Mentha citrata Ehrh. subsp. citrata): WZNIESIENIA POŁUDNIOWOMAZOWIECKIE HILLS: Łódź, Botanical Garden, section of healing plants and plants grown for industrial purposes, 29 Sept. 2002, leg. M. Siennicka (LOD 2286 PF); on telia of Puccinia punctata Link [on Galium odoratum (L.) Scop.]: WZNIESIENIA POŁUDNIOWOMAZOWIECKIE HILLS: Łódź, Las Łagiewnicki forest complex, deciduous forest (Tilio-Carpinetum typicum), roadside, 30 Sept. 2007, leg. A. Kuchnik (LOD 3126 PF); same host and locality, 21 Oct. 2007, leg. M. Ruszkiewicz-Michalska (LOD 3127 PF). Published reports of this species from Central Poland (Ruszkiewicz-Michalska 2006) are based on the following specimens from LOD (PF): no. 960 (uredinia and telia of Puccinia asperulae-cynanchicae Wurth), no. 1174 (telia of P. behenis G. H. Otth), no. 409 (telia of P. calcitrapae DC.), no. 393 [telia of P. maculosa (Str.) Rohl.], nos 982, 1003 (uredinia of P. obscura Schroet.), nos 1007, 1008 (uredinia of P. poae-nemoralis G. H. Otth), nos 395, 999, 1000 (uredinia of P. punctata Link), nos 978, 981, 983, 1004–1006 (telia of P. violae DC.), no. 959 (telia of Uromyces anthyllidis Schroet.), no. 1960 [telia of U. fallens (Arth.) Kern ex Barthol.], nos 422, 432 (telia of U. striatus Schroet.). HOST RANGE AND DISTRIBUTION. This most common parasite of uredinial and occasionally of aecial and telial sori of rust fungi was reported on 369 species belonging to 30 genera worldwide (after Płachecka 2005). In Poland it has been observed in uredinia and telia of members of seven genera: Chrysomyxa Unger (1 species), Coleosporium (1), Cumminsiella Arthur (1), Melampsora Castagne (6), Phragmidium Link (1), Puccinia (36) and Uromyces (9) (Nowak & Majewski 2005; Mułenko et al. 2008). Recently, S. filum POLISH BOTANICAL JOURNAL 55(2). 2010 was also reported to destroy aeciospores of Puccinia caricina and P. dioicae Magnus parasitizing Carex sp. (Adamska et al. 2010). However, these records are doubtful, as on that host genus both Puccinia species form uredinial and telial stages exclusively (Majewski 1979). The present paper lists one new host species for S. filum, namely Puccinia chaerophylli. Specimens reported as Darluca dianthi (Alb. & Schwein.) Migula on Puccinia arenariae (Schumach.) G. Wint. and Darluca genistalis (Fr.) Sacc. var. hypocreoides Fuckel on Melampsora epitea Thüm. and Melampsora sp. probably also belong to S. filum s.l. (Mułenko et al. 2008). NOTES. Although S. filum has been assumed to be a nonspecific rust hyperparasite, molecular studies revealed genetic diversity within the species, suggesting that it either comprises more than one species or shows host specialization (Liesebach & Zaspel 2004; Nischwitz et al. 2005; Bayon et al. 2006). Some studies based on ITS sequence data indicate that S. filum and Ampelomyces quisqualis, both placed among the pleosporalean genera, evolved from a common ancestor (Nischwitz et al. 2005). ACKNOWLEDGEMENTS. I am grateful to Professor Tomasz Majewski for his supervision, support and encouragement. I thank Professor Uwe Braun (Martin Luther University, Germany) for providing essential literature, Professor Maria Ławrynowicz (University of Łódź, Poland), Curator of the Fungal Collection of Herbarium Universitatis Lodziensis, for permission to analyze herbarium materials, Dr. hab. 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