Immotthia atrograna (Dacampiaceae, Ascomycota), a new for Ukraine fungicolous fungus from the Carpathians

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84 ISSN 0372-4123. Ukr. Bot. J., 2016, 73(1)
Immotthia M.E. Barr is a small genus currently
classified in the family Dacampiaceae (Pleosporales,
Dothideomycetes, Ascomycota). The Dacampiaceae,
originally named as 'Dacampieae', at present comprises
20 genera. Of them, twelve are lichenicolous, seven
contain saprobes growing on plant litter or non-
specialized plant pathogens, and a single one,
Immotthia, is exclusively fungicolous genus (Barr, 1987b;
Dictionary of the Fungi, 2011; MycoBank, 2015). In
2002, M. Barr transferred the non-lichenicolous genera
to the Teichosporaceae M.E. Barr; however, this familial
placement has not been largely accepted.
Originally, M. Barr (1987a) established a new
genus Immotthia to accommodate a single species,
Immotthia hypoxylon (Ellis & Everh.) M.E. Barr
(basyonym Amphisphaeria hypoxylon Ellis et Everh.,
≡ Otthia hypoxylon (Ellis & Everh.) Ellis & Everh.). This
monotypic genus was characterized by rather small
globose to obpyriform ascomata, usually gregarious
on hypostroma, long-stalked bitunicate asci, and
fusoid pigmented one-septate ascospores, uniseriately
arranged in the asci. The genus name means «not at
all Otthia» reflecting morphological distinctness from
the genus Otthia Nitschke ex Fuckel, where the type
species has been previously placed. The species epithet
derives from association of the fungus with stromata of
Hypoxylon spp. hosts.
Later, after examination of the holotype specimen
of Sphaeria atrograna Cooke & Ellis, M. Barr (1993)
proposed a new combination in the genus Immotthia,
I. atrograna (Cooke & Ellis) M.E. Barr. At the same
time, she pointed out that the studied collection
morphologically was not different from Immotthia
hypoxylon, thus the species were considered as
synonyms. The synonymy was confirmed in further
studies of type material by W. Jaklitsch et al. (2002). Of
the two basyonyms, Sphaeria atrograna was described
seven years earlier than Amphisphaeria hypoxylon,
therefore the priority name of the fungus is Immotthia
atrograna.
In 2002, M. Barr published a new combination,
Immotthia atroseptata (Piroz.) M.E. Barr, based
on Didymosphaeria atroseptata Piroz. parasitizing
apothecia of Pestalopezia rhododendri Seaver on fallen
leaves of Rhododendron maximum L. Described from
USA, this fungus is known exclusively from the holotype
specimen (Grand, 1973; Pirozynski, 1973; Barr, 2002).
One more morphologically similar yet undescribed
species, repeatedly collected in several countries of
Western Europe (Denmark, France, Luxembourg),
is believed to belong to the genus Immotthia. This
parasitic fungus was found in all localities on apothecia
of another discomycete, Velutarina rufo-olivacea (Alb.
& Schwein.) Korf (AscoFrance, 2015).
doi: 10.15407/ukrbotj73.01.084
O.Yu. AKULOV1, V.P. HAYOVA2
1 V.N. Karazin National University of Kharkiv
4, Svobody Sq., Kharkiv, 61077, Ukraine
alex_fungi@yahoo.com
2 M.G. Kholodny Institute of Botany, National Academy of Sciences of Ukraine
2, Tereshchenkivska Str., Kyiv, 01004, Ukraine
v.hayova@gmail.com
IMMOTTHIA ATROGRANA (DACAMPIACEAE, ASCOMYCOTA), A NEW FOR UKRAINE
FUNGICOLOUS FUNGUS FROM THE CARPATHIANS
Akulov O.Yu., Hayova V.P. Immotthia atrograna (Dacampiaceae, Ascomycota), a new for Ukraine fungicolous fungus from
the Carpathians. Ukr. Bot. J., 2016, 73(1): 84–89.
Abstract. The paper reports first records of new for Ukraine genus and species of fungicolous fungi, Immotthia atrograna
(Cooke & Ellis) M.E. Barr. All collections were made in old-growth forests within protected areas in the Ukrainian
Carpathians during recent mycological surveys. The fungus was found to be associated with two xylariaceous species,
Annulohypoxylon cohaerens (Pers.) Y.M. Ju, J.D. Rogers & H.M. Hsieh and A. multiforme (Fr.) Y.M. Ju, J.D. Rogers et
H.M. Hsieh. Diagnostic description, nomenclature data, geographical distribution and original illustrations for both
sexual and asexual morphs of the reported species are provided. Morphologically similar fungicolous fungi are briefly
reviewed and compared.
Key words: Pleosporales, Immotthia, Hypoxylon, Coniothyrium parasitans, mycoparasite, Ukrainian Carpathians
© O.Yu. AKULOV, V.P. HAYOVA, 2016

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Thus, all known up to now species of the Immotthia
genus are apparently obligate fungicolous fungi. The
genus is not different in ecological preferences from
other fungicolous genera, such as Hypomyces (Fr.)
Tul. & C. Tul. or Cosmospora Rabenh., although is
represented by much lower number of species.
In Ukraine, none of the Immotthia species has been
registered so far. Several specimens recently collected by
the authors, independently in different localities in the
Ukrainian Carpathians, were subsequently identified
as Immotthia atrograna. Since both genus and species
are newly recorded in Ukraine, we provide below
nomenclature data, diagnostic description, original
illustrations, geographical distribution and other related
information for the reported fungus. The specimens are
deposited in Herbarium of the Department of Mycology
and Immunology, V.N. Karazin National University of
Kharkiv (CWU Myc) and in Mycological Herbarium of
M.G. Kholodny Institute of Botany, National Academy
of Sciences of Ukraine (KW).
Immotthia atrograna (Cooke & Ellis) M.E. Barr,
Mycotaxon 46: 71, 1993 (Fig, 1–3)
Syn.: Sphaeria atrograna Cooke & Ellis, Grevillea
8(45): 15, 1879; ≡ Amphisphaeria atrograna (Cooke
& Ellis) Sacc., Syll. Fung. 1: 722, 1882; ≡ Gibbera
atrograna Trans. Brit. Mycol. Soc. 65(3): 396, 1975;
≡ Melamomma atrogranum (Cooke & Ellis) Cooke,
Grevillea, 16: 52, 1887; = Amphisphaeria deformis
[deformata] Ellis & Langl., J. Mycol. 4(12): 123, 1888;
= Amphisphaeria pilosella Ellis & Everh., Proc. Acad.
nat. Sci. Philad. 47: 418, 1895; ≡ Gibbera pilosella (Ellis
& Everh.) Sivan., Trans. Brit. Mycol. Soc. 65(3): 395,
1975; = Amphisphaeria confertissima Ellis & Everh.,
Proc. Acad. nat. Sci. Philad. 47: 418, 1895; ≡ Gibbera
confertissima (Ellis & Everh.) Sivan., Trans. Brit.
Mycol. Soc. 65(3): 395, 1975; = Immotthia hypoxylon
(Ellis & Everh.) M.E. Barr, Mycotaxon 29: 504, 1987;
≡ Amphisphaeria hypoxylon Ellis & Everh., J. Mycol.
2(4): 41, 1886; ≡ Otthia hypoxylon (Ellis & Everh.)
Ellis et Everh., 1892; ≡ Dimerium hypoxylon (Ellis &
Everh.) Petr., Sydowia 11: 338, 1958; ≡ Melamomma
hypoxylon (Cooke & Ellis) Cooke, Grevillea, 16: 53,
1887; ?= Neopeckia episphaeria Höhn., Ann. Mycol.
17(2–6): 120, 1920; ?= Sphaeria insidens Schwein.,
Schr. naturf. Ges. Leipzig 1: 39, 1822; ≡ Myrmaecium
insidens (Schwein.) Sacc., Syll. Fung. 1: 601, 1882;
≡ Anthostoma insidens (Schwein.) Berl. & Voglino, Syll.
Fung. Add. Vol . I–IV: 47, 1886; ≡ Hypoxylon insidens
(Schwein.) Ellis & Everh., N. Amer. Pyren. (Newfield):
653, 1892; ?= Jahnula parasitica Kirschst. ined. Petrak,
1958.
Anamorph: Coniothyrium parasitans (Berk. &
Ravenel) Tassi, Bulletin Labor. Orto Bot. de R. Univ.
Siena 5: 25, 1902; ≡ Sphaeropsis parasitans Berk. &
Ravenel, Grevillea 2(24): 180, 1859; ≡ Macroplodia
parasitans (Berk. & Ravenel) Kuntze, Revis. gen.
pl. (Leipzig) 3: 492, 1898; ≡ Cicinnobella parasitans
(Berk. & Ravenel) Petr., Sydowia 11: 338, 1958.
Icon.: Chlebicki, Acta Mycologica, 2005, 40(1):
76, fig. 1D (as Immotthia hypoxylon); Jaklitsch et al.,
Österreichische Zeitschrift für Pilzkunde, 2002, 11(14):
103, figs. 9–12; Barr, Prodromus…, 1987, p. 101, Pl.
23 Q; Barr, Mycotaxon, 2002, 82(3): 379: figs. q-s; Barr,
Mycotaxon, 1993, 46(1): 69, figs. p-r; Sivanesan, Trans.
Brit. Mycol. Soc., 1975, 65(3): 396, fig. 1A, Pl. 50(8).
Hypostroma appearing as a dark brown to black crust
under pseudothecia on the surface of host stromata, 50–
300 µm wide, in section composed of dark brown textura
angularis made up of thick-walled (0.5–1.5 µm) cells,
6–11 µm in diameter, similar to cells of ascomatal wall
in surface view. Ascomata (pseudothecia) superficial on
hypostroma, numerous, usually gregarious to densely
aggregated, globose, obpyriform, often becoming
laterally compressed by mutual pressure, 120–270 (up
to 385) µm in diameter, black, carbonaceous, surface
glabrous to roughened by protruding cells. Ostioles
pallid to reddish-brown, inconspicuous, 30–50 µm in
diameter, appearing as a circular pore in apical part of
ascomata, occasionally slightly papillate. Interior of the
ostiolar canal is lined with short hyaline to pale brown
periphyses, broadly rounded at the apex, 8–18 × 2–3 µm.
Peridium three-layered, 30–50 µm wide, equally thick
or slightly thicker toward the ascomatal apex. External
layer 10–15 µm wide, composed of dark angular cells
resembling those of hypostroma. Middle layer 6–15 µm
thick, consists of 2–4 layers of more loosely arranged
and lighter coloured cells. Internal layer 10–20 µm in
width, of pale to subhyaline cells 3–8 µm in diameter. In
addition, at the base of ascomata, between the internal
layer and hymenium, a sterile tissue up to 80 µm high
is frequently formed. The outer surface of peridium
smooth to verruculose, covered by protruding cells
of the external layer of pseudothecia. Hamathecium
composed of numerous pseudoparaphyses, hyaline,
filiform, cellular, branching, easily detached from
peridium, exceeding asci in length, 1.5–3.0 µm wide.
Asci bitunicate, oblong cylindrical, thick-walled (up
to 3 µm thick at the apex), formed in a broad basal
fascicle, (4–6–)8-spored, (50–)60–90 × (5–)6–10

86 ISSN 0372-4123. Ukr. Bot. J., 2016, 73(1)
µm; ascospores arranged within the asci in obliquely
uniseriate manner. Ascospores yellow-brown to
reddish-brown, ellipsoid, obovoid to biconical, slightly
asymmetric, inaequilateral, one-septate, constricted at
septum, (8–)9–14(–18) × (4.5–)5–6(–7) µm; septum
central to eccentric, about 1 µm in width, slightly darker
than spore walls; upper cell longer and wider than lower
one, ends subacutely rounded, 1–3 guttules per cell;
walls smooth to verruculose, surrounded by a hyaline
perispore, non-dehiscent in KOH.
Anamorph pycnidial; in external appearance
and general shape conidiomata remarkably similar
to ascomata, sometimes differ in slightly smaller
size. Conidiophores absent. Conidiogenous cells
enteroblastic, hyaline, smooth, discrete, determinate,
irregularly ampulliform or doliiform, (8–)10(–14)
× 5(–7) µm, proliferating as typical phialides, with
minute colarette and conspicuous periclinal thickening.
Conidia ellipsoid, rounded at both ends or slightly
tapering toward one end, one-celled, at first hyaline,
later light- to medium-brown, 6–8(–9) × 3–5 µm,
smooth or finely verruculous, usually bi-guttulate*.
* A few collections from USA, quite closely resembling
Immotthia atrograna, differ in having smaller ascospores and
conidia. For example, a specimen from North Carolina at BPI
with ascospores 6–8(–9) × 2–3 µm and conidia 4–5 × 1 µm
has unclear taxonomic position (Barr, 1993).
Specimens examined. On stromata of Annulohypoxylon
cohaerens (Pers.) Y.M. Ju, J.D. Rogers & H.M. Hsieh
on wood of Fagus syvlatica L. — Chernivtsi Region,
Vyzhnytsia District, Vyzhnytsia National Nature
Park, fir-beech forest, 48º 12´N 25º 12´E, 22.08.2015,
V.P. Hayova [KW 60674]. — Ivano-Frankivsk Region,
Nadvirna District, Gorgany Nature Reserve, Gorgany
forestry, spruce-beech forest, quarter 14, 48º 29´N 24º
17´E, 29.08.2010, V.P. Hayova [CWU (Myc) AS 5803,
= KW 60665]. — Lviv Region, Skole District, Skolivski
Beskidy National Nature Park, old-growth beech forest
on the slope of Mount Parashka (protected area), 49º
04´ 04´´N 23º 25´ 32´´E, 22.09.2011, O.Yu. Akulov
[CWU (Myc) AS 4704].
On stromata of Annulohypoxylon multiforme (Fr.)
Y.M. Ju, J.D. Rogers & H.M. Hsieh on bark of Alnus
incana (L.) Moench. — Ivano-Frankivsk Region,
Nadvirna District, Gorgany Nature Reserve, Gorgany
forestry, along Dzhurdzhynets stream, 48º 28´N 24º
17´E, 28.08.2010, V.P. Hayova [KW 60669].
On stromata of A. multiforme on bark of Betula
pendula Roth. — Ivano-Frankivsk Region, Nadvirna
District, Gorgany Nature Reserve, beech-spruce forest
with admixture of birch, quarter 14, 48º 29´N 24º 17´E,
30.08.2010, V.P. Hayova [KW 60671]
General distribution. Europe: Austria, Belgium,
France, Lithuania, Norway, Poland, Russia (European
part), Sweden, Switzerland, Ukraine. Asia: China.
Fig. 1. Immotthia atrograna [CWU Myc AS 4704]: а — habit on stromata of Annulohypoxylon cohaerens, bar: 2 cm; b — general
view of ascomata and conidiomata, bar: 500 µm

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Fig. 2. Immotthia atrograna [CWU
Myc AS 4704]: а — vertical section
through ascomata, bar: 250 µm; b —
surface view of ascoma, bar: 250
µm; c — ascomatal peridium; d —
ascospores, bar: 13 µm, e — ascus
and part of hamathecium, bar: 40
µm
Fig. 3. Anamorph of Immotthia
atrograna, Coniothyrium parasitans
[KW 60665]: а — conidiogenous
cells, bar: 10 µm; b — conidia, bar:
8 µm

88 ISSN 0372-4123. Ukr. Bot. J., 2016, 73(1)
North America: USA. South America: Venezuela,
Puerto-Rico.
Host specialization. Stromata of Annulohypoxylon
cohaerens **, A. multiforme, Hypoxylon rubiginosum
(Pers.) Fr., and H. perforatum (Schwein.) Fr.,
occasionally also a decayed wood of various hardwoods,
particularly in humid forest habitats.
Polish mycologists A. Chlebicki and A. Skirgiełło
(1995) consider Immotthia atrograna «an obligatory
fungicolous saprophyte». On the contrary, W. Jaklitsch
et al. (2002) are inclined to regard I. atrograna as an
obligate parasite attacking host stromata in all stages of
their development. In the latter case, parasitic nature
of the fungus is supported by inability of ascospores to
germinate on artificial media.
The anamorph of this fungus indicated here as
Coniothyrium parasitans belongs to morphologically
extremely variable group of coelomycetous fungi.
Recent studies of cultural characteristics and DNA
sequence data of the Coniothyrium-like fungi have
demonstrated that their morphological features are often
not suitable to delimit formerly recognisable species
and genera. Based on multi-locus DNA phylogeny
combined with detailed morphological analyses, several
new genera were proposed; moreover, some families
within the Pleosporales were redefined (Verkeley et al.,
2004, 2014). In the present study, we did not observe
annellidic conidiogenous cells, the most distinctive
Coniothyrium character. Thus asexual morph of the
reported species, due to phialidic conidiogenesis and
described above conidiomatal and conidial characters,
may be transferred to one of the novel genera; however,
molecular phylogenetic evidence for the redisposal is
required.
Morphologically very close to Immotthia atrograna
is a non-fungicolous species, Coleroa pusiola (P. Karst.)
Sivan., described from wood of Salix myrsinifolia Salisb.
(as S. nigricans Smith) from Finland. This fungus was
previously known as Amphisphaeria pusiola P. Karst., or
Didymosphaeria pusiola (P. Karst.) Rehm. Description
and illustrations of the holotype specimen of Coleroa
pusiola provided by A. Sivanesan (1975) show ascospore
size 9–12 × 3.5–4.5 µm, while the holotype collection
of Immotthia atrograna features slightly longer and wider
ascospores (13–15 × 5–7 µm). M. Barr (1993) who
examined numerous specimens of Immotthia atrograna,
observed very wide ascospore size range in this species.
She suggested that Coleroa pusiola might be another
** On stromata of Annulohypoxylon cohaerens this fungus is
first reported here.
species of Immotthia, with a thinner peridium, up to 25
µm wide, and slightly smaller ascospores.
Quite resembling Immotthia atrograna in general
appearance and micromorphological features are
species of another genus, Didymosphaeria Fuckel. They
also have one-septate brown ascospores; moreover,
some species are characterized by fungicolous habit, for
example, Didymosphaeria eutypae Sureya, D. cocconiae
Arx (= Didymosphaeria cocconiae var. major Bat. &
Peres), and D. conoidea Niessl. D. eutypae, a parasite
of stromatic pyrenomycete Eutypa spp. described from
France, differs in having wider ascospores (8 × 7.5–8
µm). D. cocconiae has significantly larger ascospores
(18–24 × 7–8.5 µ m). This species described as a parasite
of Hysterostomella spurcaria (Berk. & Broome) Höhn.
(as Cocconia spurcaria (Berk. & Broome) Arx) is known
from the Philippines islands and Brazilia (Pirozynski,
1973). D. conoidea has almost identical in size ascospores
(9–)10–12(–14) × 4–5(–6) µm; however, it occurs
mostly on members of the Leptosphaeriaceae M.E. Barr
on plant litter (Shoemaker, Babcock, 1990).
A student of V.N. Karazin National University of
Kharkiv, O.V. Romanchenko, is thanked for preparing
line drawings for this publication.
REFERENCES
ASCOFrance, available at: http://www.ascofrance.com (ac-
cessed 12 March 2015)
Barr M.E. New taxa and combinations in the loculoascomy-
cetes, Mycotaxon, 1987a, 29: 501–505.
Barr M.E. Prodromus to class Loculoascomycetes, Amherst:
University of Massachusetts, 1987b, 168 pp.
Barr M.E. Redisposition of some taxa described by J.B. El-
lis, Mycotaxon, 1993, 46: 45–76.
Barr M.E. Teichosporaceae, another family in the Pleospora-
les, Mycotaxon, 2002, 82: 373–389.
Chlebicki A. Some ascomycete fungi from primeval forests of
north-east Poland, Acta Mycologica, 2005, 40(1): 71–94.
Chlebicki A., Skirgiełło A. Some mycogenous fungi from
Poland, Acta Mycologica, 1995, 30(1): 81–93.
Dictionary of the Fungi, 10th ed. Eds P.M. Kirk and
P.F. Cannon, D.W. Minter, J.A. Stalpers, Egham, UK:
CABI Bioscience; Utrecht, The Netherlands: CBS,
2011, 784 pp.
Eriksson O.E. The non-lichenized Pyrenomycetes of Sweden,
Lund: SBT-forlaget, 1992, 208 pp.
Grand L.F. Pestalopezia rhododendri and an associated hy-
perparasite in North Carolina, J. Elisha Mitchell Sci. Soc.,
1973, 89: 111–114.
Jaklitsch W., Scheuer Ch., Voglmayr H. Notes on the genus
Immotthia (Pleosporales, Ascomycetes), including some
type studies, Österreichische Zeitschrift für Pilzkunde,
2002, 11(14): 93–106.

89
ISSN 0372-4123. Укр. ботан. журн., 2016, 73(1)
MycoBank, available at: http://www.mycobank.org (ac-
cessed 6 April 2015)
Petrak F. Mykologische Bemerkungen, Sydowia, 1957
[1958], 11(1–6): 37–353.
Pirozynski K.A. Three hyperparasites of Ascomycetes, My-
cologia, 1973, 65(4): 761–767.
Shoemaker R.A., Babcock C.E. Didymosphaeria conoidea
Niessl. In: Fungi Canadenses, 1990, Pl. 328.
Sivanesan A. Redisposition and descriptions of some Am-
phisphaeria species and a note on Macrovalsaria, Trans-
actions of the British Mycological Society, 1975, 65(3):
395–402.
Verkley G.J.M., da Silva M., Wicklow D.T., Crous P.W.
Paraconiothyrium, a new genus to accommodate the my-
coparasite Coniothyrium minitans, anamorphs of Para-
phaeosphaeria, and four new species, Studies in Mycology,
2004, 50: 323–335.
Verkley G.J.M., Dukik K., Renfurm R., Göker M.,
Stielow J.B. Novel genera and species of coniothyrium-
like fungi in Montagnulaceae (Ascomycota), Persoonia,
2014, 32: 25–51.
Wang, Y.Z., Aptroot A., Hyde K.D. Revision of the
ascomycete genus Amphisphaeria, Hong Kong: Fungal
Diversity Press, 2004, 168 pp.
Recommended for publication Submitted 15.10.2015
by V.P. Heluta
Акулов О.Ю.1, Гайова В.П.2 Immotthia atrograna — новий
для території України вид мікофільних грибів із Карпат. —
Укр. ботан. журн. — 2016. — 73(1): 84–89.
1 Харківський національный університет імені
В.Н. Каразіна
пл. Свободи, 4, м. Харків, 61077, Україна
2 Інститут ботаніки імені М.Г. Холодного НАН України
вул. Терещенківська, 2, м. Київ, 01004, Україна
Повідомляється про перші знахідки в Україні мікофіль-
ного гриба Immotthia atrograna (Cooke & Ellis) M.E. Barr.
Усі зразки зібрані в старовікових лісах під час мікологіч-
них обстежень на природоохоронних територіях Україн-
ських Карпат. Гриб знайдений на двох видах грибів —
Annulohypoxylon cohaerens (Pers.) Y.M. Ju, J.D. Rogers
& H.M. Hsieh і A. multiforme (Fr.) Y.M. Ju, J.D. Rogers &
H.M. Hsieh. Наведено діагностичний опис, номенкла-
турні дані, географічне поширення та оригінальні ілю-
страції, а також вміщено стислі відомості для порівнян-
ня з морфологічно подібними мікофільними грибами.
Ключові слова: Pleosporales, Immotthia, Hypoxylon,
Coniothyrium parasitans, мікопаразити, Українські
Карпати.
Акулов А.Ю.1, Гаевая В.П.2 Immotthia atrograna — новый
для территории Украины вид микофильных грибов из
Карпат. — Укр. ботан. журн. — 2016. — 73(1): 84–89.
1 Харьковский национальный университет имени
В.Н. Каразина
пл. Свободы, 4, г. Харьков, 61077, Украина
2 Институт ботаники имени Н.Г. Холодного НАН
Украины
ул. Терещенковская, 2, г. Киев, 01004, Украина
Сообщается о первых находках в Украине микофильно-
го гриба Immotthia atrograna (Cooke & Ellis) M.E. Barr. Все
образцы собраны в старовозрастных лесах во время ми-
кологических обследований на природоохранных тер-
риториях Украинских Карпат. Гриб обнаружен на двух
видах грибов — Annulohypoxylon cohaerens (Pers.) Y.M. Ju,
J.D. Rogers & H.M. Hsieh и A. multiforme (Fr.) Y.M. Ju,
J.D. Rogers & H.M. Hsieh. Приведены диагностическое
описание, номенклатурные данные, географическое
распространение и оригинальные иллюстрации, крат-
кие сведения для сравнения с морфологически сходны-
ми микофильными грибами.
Ключевые слова: Pleosporales, Immotthia, Hypoxylon,
Coniothyrium parasitans, микопаразиты, Украинские
Карпаты.