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Genus Skeletocutis (Polyporaceae) in the Ukrainian left bank forest steppe

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Upon review of the reference data and the author's herbarium specimens were recorded five species of the Skeletocutis genus in the Ukrainian Left Bank Forest Steppe. One species, S. brevispora, is a new record for Ukraine, another one, S. carneogrisea, was found for the first time in the Ukrainian Left Bank Forest Steppe. For other two species, S. nivea, S. subincarnata, new localities were found within the studied region.
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~ 34 ~ ВІСНИК Київського національного університету імені Тараса Шевченка ISSN 1728-3817
УДК 582.284 (477.5)
M. Shevchenko, PhD student, I. Dudka, DSc
M.G. Kholodny Institute of Botany, the National Academy of Sciences of Ukraine, Kyiv, Ukraine
GENUS
SKELETOCUTIS
(POLYPORACEAE)
IN THE UKRAINIAN LEFT BANK FOREST STEPPE
Upon review of the reference data and the author's herbarium specimens were recorded five species of the Skeletocutis
genus in the Ukrainian Left Bank Forest Steppe. One species, S. brevispora, is a new record for Ukraine, another one,
S. carneogrisea, was found for the first time in the Ukrainian Left Bank Forest Steppe. For other two species, S. nivea,
S. subincarnata, new localities were found within the studied region.
Key words: genus, Skeletocutis brevispora, distribution, Ichnia National Natural Park, Ukraine.
Introduction. Skeletocutis Kotl. et Pouzar is a genus of
aphyllophoroid fungi with poroid hymenophore belonging to
Polyporaceae Fr. ex Corda family. The genus was
described in 1958 by Czech mycologists using the type
species Skeletocutis amorpha (Fr.) Kotl. & Pouzar [15]. In
1982 A. David referred several species to this genus
(S. azorica (D.A. Reid) Jülich, S. jelicii Tortič & A. David,
S. portcrosensis A. David and S. subsphaerospora
A. David) with incrusted generative hyphae and monomitic
hyphal system [12]. However, reference of such species to
Skeletocutis genus has been deemed arguable. Thus,
L. Ryvarden, R. Gilbertson [20] and A. Bernicchia [7]
treated the species with monomitic hyphal systems as a
part of Ceriporiopsis Domański genus and delimited
Skeletocutis based on its dimitic system and encrustation
of the hyphae at dissepiment edges.
The molecular and phylogenetic studies completed in
the early ХХІ century demonstrated that the hyphal
system's type is not a diagnostic feature for Skeletocutis
genus [18, 23]. Relying on the foregoing data it was found
out that the genus also comprises the species with
monomitic hyphal system [18, 23].
Two species, Skeletocutis lenis (P. Karst.) Niemelä and S.
vulgaris (Fr.) Niemelä & Y.C. Dai, were transferred to a new
genus Sidera Miettinen & K.H. Larss. (Hymenochaetales
Oberw.) based on ITS (internal transcribed spacer) and LSU
(large subunit) nuclear ribosomal DNA (nrDNA) sequence
analyses [18]. The main morphological difference between the
two genera is that Skeletocutis usually has hyphae encrusted
by fine crystals on the tube mouths, whereas in Sidera the
dissepiment edge hyphae are smooth or covered with a few
faceted crystal clusters [18, 19].
Genus Skeletocutis comprises the species with annual
and perennial basidiocarps. Basidiocarps resupinate or
pileate; normally white or occasionally grey, pink or purple
colored [21]. The most of this species have a dense
cartilaginous zone above the tube layer [21]. According to
the contemporary concept, the genus comprises the
species with monomitic, dimitic and trimitic hyphal system
[18, 23]. This genus's inherent feature is generative hyphae
encrustation at dissepiment edges by crystals resembling
rose thorns. The generative hyphae with clamps, the
skeletal hyphae hyaline, thin to slightly thick-walled,
occasionally branched. Cystidia absent, however, conical
or fusoid cystidioles are observed in most of the species.
Basidia four-spored, short clavate or barrel-shaped. Spores
hyaline, cylindrical or ellipsoid, in most species they are
somewhat curved [20, 22, 25].
Both in Ukraine and globally, most of Skeletocutis spe-
cies are saprotrophes on the dead wood of various conifers
and broadleaved trees. Such species cause white rot.
Some of them are able to develop on basidiocarps of the
other polypores [21, 22].
According to the data base Index Fungorum (request,
dated April 24, 2017), Skeletocutis genus comprises 43
species. Fungi of this genus are widely distributed in the
Northern Hemisphere. Twenty of them are known in Europe
[7, 21] and only six species have been recorded in Ukraine
[2-6]. According to the reference data, prior to our research
there had been three species known for the Left Bank Forest
Steppe: Skeletocutis nivea (Jungh.) Jean Keller, S. odora
(Peck ex Saccardo) Ginns, S. subincarnata (Peck) Jean
Keller. All of them were found in Kharkiv Oblast [4-6].
Materials and Methods
Among materials applied for this article there were
species of aphyllophoroid fungi collected in June-October,
2016 at the forestlands of Ichnia National Natural Park
(Ichnia Rayon, Chernihiv Oblast). The Natural Park's area
in terms of mycofloristic zoning of Ukraine belongs to the
Left Bank Forest Steppe [1].
The micromorphological structures of the specimens
were examined in a 5% aqueous potassium hydroxide
solution and Melzer's reagent. The nomenclature of the
species follows the "MycoBank" database
[http://www.mycobank.org/quicksearch.aspx].
Results and Discussion
Upon review of the herbarium specimens collected over
the specified area, we have reported 4 species belonging to
Skeletocutis genus. Among them, one species (Skeletocutis
brevispora Niemelä) is new for Ukraine, another one
(Skeletocutis carneogrisea A. David) was found rare for our
country and the new one for the Forest Steppe, for two
species (Skeletocutis nivea (Jungh.) Jean Keller, Skeletocutis
subincarnata (Peck) Jean Keller) the new locations were
found in the Left Bank Forest Steppe. Generally, according to
the reference data and the author's own gatherings, as of the
date, five species of fungi belonging to this genus have been
known for the Left Bank Forest Steppe. The comparative
features of these species are shown in table 1.
Table 1. Comparative Features of Skeletocutis known for Ukrainian Left Bank Forest Steppe
Species Spores dimensions, µm Number of pores per
1 mm Substrate
Skeletocutis brevispora
Niemelä
3,1(3,4)–4,1(4,6)×
1,1(1,2)–1,6 6-8(9) On basidiocarps only Phellinidium
ferrugineofuscum (P. Karst.) Fiasson & Niemelä.
Skeletocutis carneogrisea
A. David (2,7)3–3,6(4,2)×1–1,1(1,2) 4-6 On basidiocarps and wood of conifer species
affected by Trichaptum sp.
Skeletocutis nivea
(Jungh.) Jean Keller 2,8–3,3(3,7)×0,6–0,8 8-10 On dead wood of broadleaved trees.
Skeletocutis odora (Peck
ex Saccardo) Ginns
(3,4)3,9–5,2(5,7) ×
(0,8)0,9–1,4(1,6) 4-6 Most often on Picea abies (L.) H. Karst. and
Populus tremula L.
Skeletocutis subincarnata
(Peck) Jean Keller (3,2)3,5–5,5×(1,1)1,3–1,8 5-7 On dead wood of conifers (more rarely on
broadleaved trees).
© M. Shevchenko, I. Dudka , 2017
ISSN 1728-2748 БІОЛОГІЯ. 2(74)/2017 ~ 35 ~
Find below the generalizing list of Skeletocutis species
for the Left Bank Forest Steppe. For S. brevispora, listed for
Ukraine's territory for the first time, we have submitted
description of the macro- and micromorphological structures
of the examined sample and the general global distribution.
For the rest of the species collected by the author, we have
provided details of the collecting date and localities,
substrate specialization and distribution in Ukraine.
Basidiomycota Bold ex R.T. Moore
Agaricomycotina R. Bauer, Begerow, J.P. Samp.,
M.Weiss et Oberw.
Agaricomycetes Matheny, Hibbett et Binder
Polyporales Gäum.
Polyporaceae Fr. ex Corda
Skeletocutis brevispora Niemelä,
Acta bot. fenn. 161: 10 (1998)
Basidiocarps annual, resupinate, 1–2 mm thick, waxy
when fresh and firm-ceraceous after drying. Pore surface
at first white, later straw- to lemon-yellowish, pores angular
at first 6–9 per mm. Margin initially pruinose-byssoid, white,
later lacking or very narrow. Subiculum very thin, whitish.
Hyphal system dimitic: skeletal hyphae slightly thick-walled,
dominating in all parts of basidiocarps, 3–4 µm wide,
swelling up to 5–5,5 µm wide in KOH (but not dissolving);
generative hyphae thin-walled, with clamps, 3–3,5 µm
wide, at the dissepiment edges slightly encrusted. Cystidia
none, bottle-shaped cystidioles with sharp apices present
in the hymenium, 10–13×4–5,5 µm. Basidia short-clavate
10–12×4–4,5 µm, with a basal clamp. Basidiospores 3,4–
4,1×1,2–1,5 µm, cylindrical to slightly allantoid tapering
towards the apiculus, without guttules.
Specimens examined: Ichnia National Natural Park,
Ichnia, Ichnia Rayon, Chernihiv Oblast, Ukraine,
southwestern outskirts of Ichnia, pine forest, over the dead
last year basidiocarp of Phellinidium ferrugineofuscum
(P. Karst.) Fiasson & Niemelä on the wood of the fallen
trunk Pinus sylvestris L., July 16, 2016, KW-M 70851.
Ecological peculiarities: It develops over basidiocarps
of Phellinidium ferrugineofuscum (P. Karst.) Fiasson &
Niemelä. T. Niemelä in the species description specifies
that it develops on such substrate so often that such
feature may be applied for the species identification [19].
According to L. Ryvarden and I. Melo, S. brevispora is able
to develop on the wood of conifer species affected by
Ph. ferrugineofuscum [21].
S. brevispora is deemed a rare species, however,
locally it may be found rather often [19, 22]. Globally, it is
known in certain countries of Europe (Norway, Sweden,
Final [21], Poland [14], in the European part of Russia [16,
22] and in China [11].
It is new for Ukraine.
Skeletocutis carneogrisea A. David, Naturaliste Can.
109(2): 245 (1982)
Specimens examined: Ichnia National Natural Park,
Ichnia, Ichnia Rayon, Chernihiv Oblast, Ukraine, Budy
village, standing pine forest, over basidiocarps of
Trichaptum fuscoviolaceum (Ehrenb.) Ryvarden on the
wood of the fallen trunk Pinus sylvestris L., July 17, 2016,
August 06, 2016, September 15, 2016.
Ecological peculiarities: Inherent to the wood of
conifer species affected by Trichaptum Murrill genus's
fungi. According to the reference data and the author's
observations it is able to develop both on the wood affected
by Trichaptum and the basidiocarps of these polypores.
Distribution in Ukraine: The Left Bank Gramineous-
Meadow Steppe [2, 3]. It was found for the first time in
Ukrainian Left Bank Forest Steppe and the Forest-Steppe
Zone in general.
Skeletocutis nivea (Jungh.) Jean Keller, Persoonia
10(3): 353 (1979)
Specimens examined: Ichnia National Natural Park,
Ichnia, Ichnia Rayon, Chernihiv Oblast, Ukraine, southern
outskirts of Ichnia, mixed forest, June 24, 2016, September
16, 2016, Avhustivka village, alder forest, on the fallen
branches of Acer platanoides L., Alnus glutinosa (L.)
Gaertn., Robinia pseudoacacia L., July 16, 2016, August
06, 2016, September 16, 2016.
Ecological peculiarities: It develops on the dead
wood of broadleaved trees.
Distribution in Ukraine: The Left Bank Polissia,
Transcarpathia, Carpathian Mountains, the Right Bank and
the Left Bank Forest Steppe, the Left Bank Gramineous
and Meadow Steppe, Crimea [6].
Skeletocutis odora (Peck ex Saccardo) Ginns,
Mycotaxon 21: 332 (1984)
We failed to support this finding with own gatherings.
We explain it by the fact that this species is rare both in
Ukraine and in the world. It is confirmed by the fact that it is
Red Listed in many European Countries (Estonia [13],
Poland [9], Slovakia and Czech Republic [8]) and is a
candidate to the list of species protected by Bern
Convention [10].
Ecological peculiarities: It develops on dead hard-
woods and conifers (most often on Picea abies (L.) H.
Karst. and Populus tremula L.)
Distribution in Ukraine: Transcarpathia, Carpathian
Mountains [6], the Left Bank Forest Steppe [5].
Skeletocutis subincarnata (Peck) Jean Keller,
Persoonia 10(3): 535 (1979)
Specimens examined: Ichnia National Natural Park,
Ichnia, Ichnia Rayon, Chernihiv Oblast, Ukraine, southern
outskirts of Ichnia, mixed forest, on the fallen branches of
Acer platanoides L., June 25, 2016.
Ecological peculiarities: It develops on conifers (more
rarely on hardwoods).
Distribution in Ukraine: Transcarpathia, Carpathian
Mountains, the Right Bank and the Left Bank Forest
Steppe, the Left Bank Gramineous-Meadow Steppe [6].
Acknowledgments
The author would like to thank kindly the directorate
members of Ichnia National Natural Park for facilitation to
the filed surveys.
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(Polyporaceae). Le Naturaliste Canadien. 1982; 109: 235–272. In French.
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Poland). Polish Botanical Journal. 2015; 60(2): 217–292.
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Cechoslovakiae III. Ceská Mykologie. 1958; 12(2):95–104. In Czech.
16. Lindgren M. Polypore (Basidiomycetes) species richness and
community structure in natural boreal forests of NW Russian Karelia and
adjacent areas in Finland. Acta Botanica Fennica. 2001; 170: 1–12.
17. Lizoň P. Red list of fungi of Slovakia. In: Balaž D., Marhold K.,
Urban P., editors. Red list of plants and animals of Slovakia. Ochr. Prir.
20 (Suppl.); 2001. p. 6–13.
18. Miettinen O., Larsson K.-H. Sidera, a new genus in
Hymenochaetales with poroid and hydnoid species. Mycol. Progress. 2011;
10:131–141.
19. Niemela T. The Skeletocutis subincarnata complex
(Basidiomycetes), a revision. Acta Botanica Fennica. 1998; 161: 1–35.
20. Ryvarden L., Gilbertson R. European polypores. Synopsis
Fungorum. 1994; 7(2): 388-743.
21. Ryvarden L., Melo I. Polypoid fungi of Europe. Synopsis
Fungorum. 2014; 31: 1–455.
22. Spirin V. Notes on some rare polypores, found in Russia 2.
Junghuhnia vitellina sp. nova, plus genera Cinereomyces and Skeletocutis.
Karstenia. 2005; 45: 103–113.
23. Tomšovský M., Menkis A., Vasaitis R. Phylogenetic relationships in
European Ceriporiopsis species inferred from nuclear and mitochondrial
ribosomal DNA sequences. Fungal biology. 2010; 114: 350–358.
24. Zhou L.-W., Qin W.-M. A new species of Skeletocutis (Polyporaceae)
on bamboo in tropical China. Mycotaxon. 2012; 119: 345–350.
25. Zíbarová L., Kout J. First record of Skeletocutis ochroalba
(Polyporales) in the Czech Republic. Czech Mycol. 2014; 66(1): 61–69.
Надійшла до редколегії 22.11.17
М. Шевченко, асп., І. Дудка, д-р біол. наук
Інститут ботаніки ім. М.Г. Холодного НАН України, Київ, Україна
ГРИБИ РОДУ SKELETOCUTIS (POLYPORACEAE) ЛІВОБЕРЕЖНОГО ЛІСОСТЕПУ УКРАЇНИ
На основі аналізу літературних даних та власних гербарних матеріалів було встановлено, що для Лівобережного Лісостепу Украї-
ни відомо п'ять видів грибів із роду Skeletocutis. Один вид – Skeletocutis brevispora – є новим для України, ще один – Skeletocutis carne-
ogrisea – вперше виявлений у Лівобережному Лісостепу. Для двох видів – Skeletocutis nivea, Skeletocutis subincarnata – виявлені нові міс-
цезнаходження в межах регіону дослідження.
Ключові слова: рід, Skeletocutis brevispora, поширення, Ічнянський національний природний парк, Україна.
М. Шевченко, асп., И. Дудка, д-р биол. наук
Институт ботаники им. Н.Г. Холодного НАН Украины, Киев, Украина
ГРИБЫ РОДА SKELETOCUTIS (POLYPORACEAE) ЛЕВОБЕРЕЖНОЙ ЛЕСОСТЕПИ УКРАИНЫ
На основе анализа литературных данных и собственных гербарных материалов было установлено, что для Левобережной
Лесостепи Украины известно пять видов грибов из рода Skeletocutis. Один вид – Skeletocutis brevispora – является новым для
Украины, еще один – Skeletocutis carneogrisea – впервые обнаружен в Левобережной Лесостепи. Для двух видов – Skeletocutis nivea, Skel-
etocutis subincarnata – обнаружены новые местонахождения в пределах региона исследования.
Ключевые слова: род, Skeletocutis brevispora, распространение, Ичнянский национальный природный парк, Украина.
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The complex of Skeletocutis subincarnata (Peck) Jean Keller comprises more species than what it is generally considered to have. Skeletocutis subincarnata itself is taxonomically problematic because of great variation and misinterpretations. Skeletocutis subincarnata sensu typi bears a close resemblance to S. papyracea A. David: both are short-spored and their hyphae soften and swell in KOH. Skeletocutis biguttulata (Romell) Niemelä, comb. nova, is a related, in Europe commoner taxon, whose hyphae are negative in KOH; the name 'subincarnata' has traditionally been addressed to it. Another species, S. brevispora Niemelä. spec. nova, grows in old-growth forests of North Europe. It differs from S. subincarnata and S. biguttulata by its waxy white and when dry bright yellow colour, shorter, thicker and strongly curved spores, glued-together tramal hyphae, and by its habit to grow on dead basidiocarps of Phellinidium ferrugineofuscum (P. Karsten) Fiasson & Niemelä. All these species have spores well over 1 μm thick. Externally similar S. kuehneri A. David has very narrow (under 1 μm) spores and grows on trees decayed by species of Trichaptum. Another species in the narrow-spored group is S. chrysella Niemelä, spec. nova, which grows on dead basidiocarps of Phellinus chrysolama (Fr.) Donk. Being confined to virgin forests and dependent on other rare polypores, both S. brevispora and S. chrysella are evidently threatened. Skeletocutis boreal is Niemelä, spec. nova is resupinate and perennial; it resembles S. stellae (Pilát) Jean Keller, but its spores are twice as thick, lower tubes bear only solitary skeletals, and hyphal tips are often inflated at dissepiment edges. Skeletocutis friabilis Niemelä & Saarenoksa, spec. nova grows on hardwood in rich sites; it has rare skeletal hyphae in an otherwise monomitic tissue. For comparison S. odora (Sacc.) Ginns and S. stellae are redescribed, and nine related species are discussed on the basis of type studies. The genus Skeletocutis Kotl. & Pouzar is divided into three subgenera, and a key is given to subg. Incrustoporia. Affinities with other genera are discussed.
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Skeletocutis bambusicola sp. nov. is described and illustrated from the tropical region of Yunnan, southern China. The species is characterized by the annual, resupinate cream-colored basidiocarps, small pores, a dimitic hyphal system, small ellipsoid basidiospores, and habit on dead bamboo. The combination of characters distinguishes S. bambusicola from other known species of Skeletocutis.