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ISSN 1560–7259 (print edition)
TURCZANINOWIA
ISSN 1560–7267 (online edition)
Turczaninowia 21 (3): 29–44 (2018)
DOI: 10.14258/turczaninowia.21.3.4
http://turczaninowia.asu.ru
Поступило в редакцию 28.02.2018 Submitted 28.02.2018
Принято к публикации 09.08.2018 Accepted 09.08.2018
УДК 582.284(470)
The Meruliaceae of Russia. II. Panus
I. V. Zmitrovich1, M. A. Bondartseva1, L. G. Perevedentseva2, A. G. Myasnikov3, A. E. Kovalenko1
1 Komarov Botanical Institute, BIN RAS, Prof. Popov str., 2, St. Petersburg, 197376, Russia. Е-mail: IZmitrovich@binran.ru
2 Perm State National Research University, PGNIU, Bukireva str., 15, Perm, 614990, Russia
3 Moscow State University of Civil Engineering, MSSU, Yaroslavskoe shosse, 26, Moscow, 129337, Russia
Keywords: basidiomycetes, broadleaf wood-associated fungi, merulioid fungi, modications variability, Russia.
Summary. The history of taxonomical study of the genus Panus Fr. (Meruliaceae, Polyporales, Basidiomycota)
is considered. A current revision of the genus in Russia was carried out. Two species of the genus Panus were re-
corded in various regions of Russia, Panus conchatus (Bull.) Fr. and P. lecomtei (Fr.) Corner. For P. conchatus, the
lateral ecotype with conchiform pileus and rather dark wine-red to lilac-brown surface is more characteristic. Two
main deviations from such a neutral type were described: 1) the chromatic one, characterized by light-colored (red
or clay-yellow), usually conchiform pilei [P. conchatus var. inconstans (Pers.) Zmitr., Bondartseva, Perevedentseva,
Myasnikov et Kovalenko] and 2) the growth one, characterized by a central (often bulbous) stipe, funnel-shaped cap
and strongly inrolled margin [P. conchatus var. torulosus (Pers.) Zmitr., Bondartseva, Perevedentseva, Myasnikov et
Kovalenko]. For P. lecomtei, the ecotype having small eccentric to lateral elegant stipe is considered. However, the
stipe shape and size are variable. The stipe can be either central – rather small, with a bulbous base [P. lecomtei var.
semirudis (Singer) Zmitr., Bondartseva, Perevedentseva, Myasnikov et Kovalenko], or strongly elongated [P. lecom-
tei var. stipitata (Malk.) Zmitr., Bondartseva, Perevedentseva, Myasnikov et Kovalenko]. Four new combinations,
P. conchatus var. inconstans, P. conchatus var. torulosus, P. lecomtei var. semirudis, and P. lecomtei var. stipitata
were suggested. It was concluded that Panus represents rather well-delimited genus belonging to merulioid phylo-
genetic radiation, whose morphotype on essential features of its organization is trametoid, but supercial habitual
features make it closer to the lentinoid one. Its essential features are the abundance of brohyphae which form textura
intricata, slowly growing basidiocarps and strictly lamellate hymenophore. Apparently, such an adaptive structure
was generated at arid and warm climatic zones, and only 2 species, P. conchatus, and P. lecomtei, have been irradiated
into temperate latitudes. The substrate spectrum of these fungi is determined by their insensitivity to substrate moist-
ening and best ability to colonize hardwood, so the greatest number of their nds can be made on stumps and large
remnants of stand-formers of corresponding forest areas. In Russia, a reliable association of Panus species to Betula
spp. and Populus spp. was revealed. An ecotypic dierentiation of the genus Panus is related to the quality of sub-
strate colonized. The basidiocarps, growing over top cuts of the stumps, are characterized by strong central stipe (P.
conchatus var. torulosus, P. lecomtei var. semirudis), whereas basidiocarps with sublateral attachment are common on
fallen logs. Certain chromatic adaptations (P. conchatus var. inconstans) are associated with an insolation regime of
the habitat. During last years, the Panus representatives have attracted an interest in biomedical research development.
Their resource potential estimation should proceed from the fact that within Russian territory, such areas as Middle
Belt of European Russia, North Caucasus, Altai and other regions of Southern Siberia are promising for replenishing
the strains of P. conchatus and P. lecomtei.
Мерулиевые грибы России. II. Род Panus
И. В. Змитрович1, М. А. Бондарцева1, Л. Г. Переведенцева2, А. Г. Мясников3, А. Е. Коваленко1
1 Ботанический институт им. В. Л. Комарова, БИН РАН, ул. Проф. Попова, 2, г. Санкт-Петербург, 197376, Россия
2 Пермский государственный национальный исследовательский университет, ПГНИУ,
ул. Букирева, 15, г. Пермь, 614990, Россия
3 Московский государственный строительный университет, МГСУ, Ярославское шоссе, д. 26, г. Москва, 129337, Россия
30 Zmitrovich I. V. et al.
The Meruliaceae of Russia. II. Panus
Ключевые слова: базидиомицеты, лентиноидные грибы, мерулиевые грибы, модификационная изменчивость,
Россия.
Аннотация. Рассмотрены этапы таксономического изучения рода Panus Fr. (Meruliaceae, Polyporales, Ba-
sidiomycota). Проведена ревизия видов, известных для России. Изучены материалы Гербариев Ботанического
института им. В. Л. Комарова (LE) и Пермского государственного национального исследовательского уни-
верситета (PERM). Впервые обощены литературные и гербарные данные о внутривидовом полиморфизме
бореальных таксонов рода. Согласно имеющимся гербарным и литературным данным, на территории России
зафиксировано 2 вида рода Panus – Panus conchatus (Bull.) Fr. и P. lecomtei (Fr.) Corner. Для P. conchatus наибо-
лее характерен латеральный экотип с раковинообразной шляпкой и довольно темной лилово-коричневой по-
верхностью. Были описаны два основных отклонения от такого модального типа: 1) хроматическое, характе-
ризующееся светло-красными (красными или глиняно-желтыми) шляпками [P. conchatus var. inconstans (Pers.)
Zmitr., Bondartseva, Perevedentseva, Myasnikov et Kovalenko] и 2) ростовое, характеризующееся центральной
(часто бульбовидно вздутой) ножкой, воронковидной шляпкой и сильно подогнутым краем [P. conchatus var.
torulosus (Pers.) Zmitr., Bondartseva, Perevedentseva, Myasnikov et Kovalenko]. Для P. lecomtei наиболее харак-
терным экотипом является латеральный, с небольшой ножкой. Встречаются экотипы со вздутой центральной
[P. lecomtei var. semirudis (Singer) Zmitr., Bondartseva, Perevedentseva, Myasnikov et Kovalenko], либо сильно
удлиненной латеральной ножкой (P. lecomtei var. stipitata). Для перечисленных разновидностей было пред-
ложено 4 новые комбинации [P. conchatus var. inconstans, P. conchatus var. torulosus, P. lecomtei var. semirudis,
P. lecomtei var. stipitata Malk.) Zmitr., Bondartseva, Perevedentseva, Myasnikov et Kovalenko]. Проведенные ис-
следования позволяют характеризовать Panus как хорошо ограниченный род, относящийся к мерулиоидной
филогенетической радиации, морфотип которого в основе траметоидный, но несущий поверхностные черты
лентиноидных грибов. Его существенными особенностями являются обилие фиброгиф, образующих «пере-
путанную текстуру» ткани, медленно растущие базидиомы и строго пластинчатый гименофор. По-видимому,
такая адаптивная структура была выработана изначально в условиях теплого засушливого климата, и лишь
2 вида (P. conchatus и P. lecomtei) позднее приспособились к обитанию в умеренных широтах. Субстратный
спектр этих грибов определяется их нечувствительностью к обводненности субстрата и лучшей способно-
стью к колонизации древесины лиственных пород, поэтому наибольшее количество их находок можно сделать
на пнях и крупных остатках основных лесообразующих пород лесов умеренных и теплых широт. Гербарный
материал с территории России собран с пней и валежной древесины из родов Betula и Populus. Экотипиче-
ская дифференциация рода Panus связана с особенностями колонизированного субстрата. Базиодиомы, по-
являющиеся сверху пней, характеризуются обычно крепкой центральной ножкой (P. conchatus var. torulosus,
P. lecomtei var. semirudis), в то время как на боковых поверхностях валежных стволов развиваются латерально
прикрепленные экотипы со слабо развитой ножкой. Оценка ресурсного потенциала представителей этого рода
в пределах России должна исходить из наибольшей распространенности P. conchatus и P. lecomtei в Средней
полосе России, на Северном Кавказе, Алтае и других районах Южной Сибири, где имеет смысл организовать
работы по пополнению штаммового разнообразия этих видов.
Introduction
This paper continues a taxonomical survey on the
genera of Meruliaceae (Polyporales, Basidiomycota)
presented in Russian mycobiota (Zmitrovich et al.,
2016), and it is devoted to an interesting genus
Panus Fr., for a long time considering rst within
Pleurotaceae, later within Polyporaceae families,
but in fact being closely related to the genera
Cerrena Gray and Steccherinum Gray of merulioid
phylogenetic radiation.
The genus Panus was described by Fries in
«Epicrisis Systematis Mycologici seu Synopsis
Hymenomycetum» (Fries, 1838) where it was
characterized as follows: «Totus fungus carnoso-
coriaceus, tenax, arescens, contexto broso.
Lamellae perfectae, tenaces, rmae, inaequales,
acie acuta integerrima, saepe venoso-connexae;
trama distincta, brosa in hymenium radians.
Sporidia alba. Fungi epixyli, diormes laterales,
diu persistens». From previously described genus
Lentinus Fr. (Fries, 1825) the genus Panus was
dierentiated by Fries in the following way: «A
Lentinis genuinis, quales omnes Europaei, lamellis
tenacibus et acie integerrima mox dignoscitur».
Hereby, an entire gills edge in Panus representatives
was established as the basic discriminant character
of two genera by Fries. In total, 17 species have been
distinguished by Fries in the genus Panus, while
some of which cannot be correlated with existing
herbarium material (below such species will be
marked with an interrogation point): Panus farneus
Fr. (?), P. cyatniformis (Schae.) Fr. [= Neolentinus
cyatniformis (Schae.) Della Mag. et Trassin.],
P. pycnoticus (Klotzsch) Fr. (?), P. torulosus (Pers.)
Fr. [= P. conchatus (Bull.) Fr.], P. rudis Fr. [= P.
lecomtei (Fr.) Corner], P. velutinus (Fr.) Sacc.,
31
Turczaninowia 21 (3): 29–44 (2018)
P. hirtus Fr. (?), P. foetens Fr. (?), P. dorsalis (Bosc.)
Fr. (?), P. stipticus (Bull.) Fr. [= Panellus stipticus
(Bull.) P. Karst.], P. farinaceus (Schumach.) Fr. (=
Panellus stipticus), P. lunatus Fr. (?), P. copulatus
(Ehrenb.) Fr. [= Lentinus copulatus (Ehrenb.)
Henn.], P. patellaris Fr. [= Tectella patellaris
(Fr.) Murrill], P. delastri Fr. (?), P. lithophilus Fr.
(?). Subsequently, the character of gills edge for
dierentiation Panus from Lentinus was adopted by
many agaricologists.
An important episode in Panus taxonomy is
related to the works by Singer. In 1951, on the basis
of complex similarities between Lentinus, Panus,
and Polyporus Fr., this mycologist has united three
genera with such genera as Pseudovafolus Pat.,
Mycobonia Pat., Phyllotopsis E.-J. Gilbert et Donk
ex Singer, and Pleurotus (Fr.) P. Kumm. in the
family Polyporaceae (Singer, 1951). Considering
the Lentinus lepideus (Fr.: Fr.) Fr. as type species
for Lentinus (the more correct modern typication
is L. crinitus L.), Singer unites all the small-spored
species within the genus Panus. Thereafter he has
reissued this system three times without essential
changing of the concept (Singer, 1962, 1975, 1986).
Corner (1981) has dierentiated the genera
Lentinus and Panus on the basis of the branching
pattern of skeletal hyphae which are organized as
dendrites with an inated axial element in Lentinus
and stay unbranched in Panus. In 1983, Pegler has
published a monograph on the genus Lentinus, which
includes according to this author the Panus-union
as a subgenus, but basing on Corner’s principles.
Within the Lentinus subgenus there are considered
species with branched and swollen sclerohyphae,
whereas the Panus subgenus unites the species with
brohyphae (Pegler, 1983).
In the period of molecular taxonomy (Ko, Jung,
1999; Grand, 2004; Larsson, 2007; Lee, Lim,
2010; Miettinen, Larsson, 2011; Zmitrovich, Maly-
sheva, 2013) it was shown that the genus Panus
sensu Corner is substantially distant from Lentinus
(core Polyporaceae), but closely related to the
genus Cerrena Gray (large merulioid phylogenetic
radiation). Within the framework of “splitter’s
approach”, even two closely related families –
Cerrenaceae and Panaceae – have been described
(Justo et al., 2017), although we believe that the
preservation of the “large Meruliaceae” lies in canvas
of more balanced classication of the Polyporales.
Distracting from macromorphology, it can be
seen that Cerrena and Panus have much in common:
e. g. unbranched brohyphae, highly characteristic
scleried elements protruding the hymenium,
traditionally called as sclerocystidia or metuloids
in the genus Panus, and as pseudocystidia in the
genus Cerrena, nally, rather similar basidia and
basidiospores. It should be emphasized the absence
of any inated hyphal elements (characteristic
feature of representatives of Lentinus and Polyporus
s. l.) in all tissues of representatives of the genus
Panus. This feature, as well as rather slow growth
of the basidiomata with the formation of solid and
often twisting stipe of textura intricata, allows to
interpret Panus-like morphotype consider only
supercially similar to lentinoid one. It is rather a
unique adaptive structure combining true lamellate
hymenophore and trametoid growth and tissues
organization. Studies on hymenophore development
in Panus conchatus, P. lecomtei, and P. fulvus
(Hibbett et al., 1993) also conrm this conclusion.
The aim of the present paper is detailed
characterization of the genus Panus in Russia,
including a modern morphological elaboration,
survey of intraspecies polymorphism, substrate
preferences and resource potential, considering that
some species of this genus are known as promising
subject for biomedical research, since produce
panepoxidone and isopanepoxidone, substances that
has an inhibitory eect on a number of inammatory
chemokines secreted by cancer tissues (Erkel et al.,
1996; Shotwell et al., 2000).
Materials and Methods
The macroscopic descriptions were based on a
study of fresh and dried specimens. The materials
of the herbaria of Komarov Botanical Institute (St.
Petersburg, LE) and Perm State National Research
University (PERM) were studied. Microscopic
preparations were mounted from dried material in
Melzer’s solution, 10 % ammoniacal Congo Red
and 5 % aqueous solution of KOH, using a LOMO
Micmed-6 light microscope. The hyphal system
was revealed and described according to updated
technique (Zmitrovich et al., 2009). The size of
mature spores was measured on 30 spores in distilled
water and Melzer’s solution.
Results and Discussion
Meruliaceae Rea, 1922, British Basid.: 620.
= Podoscyphaceae D. A. Reid, 1965, Beih. Nova
Hedwigia 18: 43.
= Steccherinaceae Parmasto, 1968, Consp. syst.
Cort.: 169.
= Bjerkanderaceae Jülich, 1982, Bibl. Mycol. 85:
356.
= Hapalopilaceae Jülich, 1982, Bibl. Mycol. 85:
370.
32 Zmitrovich I. V. et al.
The Meruliaceae of Russia. II. Panus
= Hyphodermataceae Jülich, 1982, Bibl. Mycol.
85: 373.
= Mycorrhaphiaceae Jülich, 1982, Bibl. Mycol.
85: 380.
= Phanerochaetaceae Jülich, 1982, Bibl. Mycol.
85: 384.
= Phlebiaceae Jülich, 1982, Bibl. Mycol. 85:
385; Boidin, Mugnier et Canales, Mycotaxon 66:
486, 1998 illeg.
= Irpicaceae Spirin et Zmitr., 2003, Mycena 3:
48.
= Cerrenaceae Miettinen, Justo et Hibbett, 2017,
Fungal Biol. 121: 817.
= Panaceae Miettinen, Justo et Hibbett, 2017,
Fungal Biol. 121: 817.
Steccherinoideae Parmasto, 1968, Consp.
syst. Cort.: 172.
Panus Fr., 1838, Epicr. Syst. Mycol.: 396–397.
Nomen conservandum.
= Lentinopanus (Pilát) Pilát, 1941, Ann. Mycol.
39: 72 (type Agaricus conchatus Bull. : Fr.).
Basidiocarp solitary or caespitose, medium-sized
to large, slowly growing, tough and persistant, origi-
nating from a woody substratum or from sclerotium,
of lentioid habitus with gymnocarpic development.
Pileus convex, then depressed to umbilicate, tough
with dry, with hirsute, brillose-squamulose, squa-
mose, or glabrous surface. Margin mostly inrolled,
even, or radially ribbed. Hymenophore lamellate.
Gills of 2–4 levels, decurrent, rarely furcate, mod-
erately spaced to densely crowded; edge entire.
Stipe central to lateral and very reduced, sti, solid,
continuous with the pileus. Context brous, tough-
eshy to coriaceous, mostly thin. Spore print white
to cream colour.
Hyphal system dimitic with unbranched brohy-
phae. Skeletal hyphae (brohyphae) predominates
in mature basidiocarps, thick-walled, hyaline or
yellowish. Generative hyphae thin- to moderately
thick-walled, with clamp connections, hyaline. Pi-
leipellis as a repent epicutis of radially parallel
hyphae or collapsing trichoderm. Hymenophoral
trama irregular, of textura intricata, with sometimes
thickening hymenium. Gills-edge usually sterile; as
a rule, with emergent pseudocystidia (cheilocystid-
ia). Pleurocystidia absent or present pseudocystidia.
Basidia clavate with medial constriction, 4-spored,
with a basal clamp. Basidiospores cylindric, or
ellipsoid-cylindric to ovoid, hyaline, thin-walled,
smooth, lacking both a perisporium and a germ-
pore, inamyloid, acyanophilous.
On dying and dry trees, fallen logs, stumps and
large fallen branches of trees and shrubs, presum-
able angiosprems. Causes a white rot. Worldwide,
more abundant in the tropics.
Type species: Agaricus torulosus Pers.,
1801, Syn. meth. fung. 2: 475 : Fr., 1821, Syst. My-
col. 1: 181 = A. conchatus Bull., 1787, Herb. Fr. 7:
tab. 298 : Fr., 1821, Syst. Mycol. 1: 181 (selected in
Greuter et al. 2000).
Type specimen is deposited in Friesian herbari-
um of the Uppsala University Museum of Evolution
(UPS) (Ryvarden, 1991).
Dierential generic suggestions. The genus
Lentinus Fr. has a supercial resemblance, but dif-
fers by skeleto-binding hyphae with inated axial
segment (all the Panus representatives have unin-
ated skeletals). The genus Lignomyces R. H. Pe-
tersen et Zmitr. is supercially similar too, but dif-
fers by monomitic hyphal system with strongly in-
ated hyphal segments and a dorsal stem attachment
(Petersen et al., 2015). Phylogenetically related ge-
nus Cymatoderma Jungh. diers by podoscyphoid
habitus and the presence of ventricose hymenial
gloeocystidia. Phylogenetically related genus Cer-
rena diers by daedaleoid/trametoid habitus and
less elongated (in median) basidiospores.
Ecology and substrata. All the Panus repre-
sentatives are rather thermophilic, xerotolerant and
non-sensitive to substrate watercut, why they are es-
pecially often found on large logs and stumps, with-
out allocation of strict substrate specicity. On the
other hand, they have not very high enzymatic ac-
tivity and are generally not adapted to deep decom-
position of coniferous wood. This circumstance, as
well as their attraction to large-scale tree residues,
are the reasons that in zonal biomes they are most
often associated to deciduous stand formers (mostly
Betula and Populus in the temperate-boreal zone,
and Quercus and Fagus in the nemoral zone), al-
though they readily colonize many other trees.
Type of rot. All the Panus representatives cause
a white rot. Oxidative enzymes were carefully in-
vestigated in Panus lecomtei and P. conchatus
(Zhang et al., 2006; Zhou et al., 2014). It was shown
that puried enzymes of these fungi belong to the
laccases family, due to the following observations:
1) the enzyme exhibited a broad substrate pattern,
2) oxygen was used as an oxidative agent, while
there was no H2O2 to initiate the catalytic oxidation,
and 3) the determined N-terminal primary structure
of the enzyme exhibited a high degree of similarity
with the corresponding laccases sequences.
33
Turczaninowia 21 (3): 29–44 (2018)
Secondary metabolites and perspectives in
biomedical research. A metabolite of great appli-
cation value, panepoxidone has been detected in P.
lecomtei by Erkel group (Erkel et al., 1996). Such
metabolite as isopanepoxidone has been isolated
from P. conchatus by Shotwell et al. (2000), and this
substance has a similar eect reducible to the pre-
vention of degradation of inhibiting particles of NF-
κB (IκBα) that inactivate this transcriptional factor.
NF-κB chemokine represents the main pro-inam-
matory factor, constitutionally associated with can-
cer progression (Zmitrovich, 2015), therefore such
substance as panepoxidone is a prospective subject
for biomedical research, whereas the Panus species
have a great resource value.
Also, it should be mentioned the production of
pink-lilac pigment complexes by both tropical and
temperate Panus representatives which were noted
already by Miller (1967), but yet were not chemi-
cally fractioned.
Specics of Panus-like morphotype. Basidi-
ocarp development in the Panus representatives
was studied by Hibbett et al. (1993), whereas their
hyphal dierentiation was studied by Zmitrovich et
al. (2009). As it was shown, the hymenophore dif-
ferentiation in Panus involves the periclinal growth
of context hyphae below a closed surface palisade
of hymenial elements, resulting in a cantharelloid
appearance and radiate trama. This pattern is quali-
tatively dierent from that in Lentinus s. str., which
suggests that lamellae of Panus and Lentinus are not
homologous. P. conchatus and P. lecomtei basidi-
ocarps have short stipes, whereas P. fulvus basidi-
ocarps have an elongate stipe, and develop from a
pseudosclerotium. P. conchatus sporocarps devel-
oped an ephemeral partial veil that was obliterated
during basidiocarp expansion, whereas primordia
of P. lecomtei are initially gymnocarpic. Analy-
sis of hyphal system of all the tropical representa-
tives of the genus indicates the rigorous dimitism
of mature basidiocarp, whereas in P. conchatus and
P. lecomtei the hyphae sclerify slower, while mature
basidiocarps include the hyphal elements on dier-
ent stages of maturation (Zmitrovich et al., 2009).
The absence of physalohyphae determining the ac-
celerated growth of agaricoid basidiocarps makes
the growth of Panus representatives more monoto-
nous and slow, what, in combination with its hyphal
structure, brings Panus-like morphotype together
with trametoid one. However, the hymenophore of
Panus is a classical lamellate that indicates a sur-
prising convergence among agaricomycetes.
Tropical species. As a rather specic adaptive
type, the genus Panus was generated by arid and
warm climates of the planet, where the main species
diversity of the genus is concentrated. The follow-
ing species, common in various tropical regions, are
most known.
Panus ciliatus (Lév.) T. W. May et A. E. Wood,
1995, Mycotaxon 54: 148 (Bas.: Lentinus ciliatus
Lév., 1844; Syn.: L. melanophyllus Lév., 1844;
L. setiger Lév., 1844; L. echinopus Lév., 1846;
L. braccatus Lév. in Zolling., 1854; L. dichrous Lév.
In Zolling., 1854; L. zonifer Berk. et Broome, 1873;
L. egregious Massee, 1910; Panus brunneipes Cor-
ner, 1981).
P. fasciatus (Berk.) Singer, 1962, Agaricales
mod. Tax. 2nd ed.: 172 (Bas.: Lentinus fasciatus
Berk., 1840; Syn.: L. dealbatus Fr. in Lehmann,
1847; L. fuscopurpureus Kalchbr., 1880; L. holopo-
gonius Berk. ex Cooke, 1892; L. terrestris Lloyd,
1925).
P. hookerianus (Berk.) T. W. May et A. E. Wood,
1995, Mycotaxon 54: 148 (Bas.: Lentinus hookeria-
nus Berk., 1851).
P. similis (Berk. et Broome) T. W. May et A. E.
Wood, 1995, Mycotaxon 54: 148 (Bas.: Lentinus si-
milis Berk. et Broome, 1873; Syn.: L. velulinus Fr.
var. africanus P. Henn. in Engler, 1893; L. erring-
tohnii Pat., 1900; L. samurensis Pilát, 1941).
P. strigellus (Berk.) Chardon et Toro, 1934,
Monogr. Univ. Porto Rico Ser. B 2: 315 (Bas.: Lenti-
nus strigellus Berk., 1868; Syn.: Panus guaraniticus
Speg., 1883; Lentinus crispus Pat., 1889; L. tubarius
Pat., 1899; Pocillaria palmeri Earle, 1906; Lentinus
subglaber Lloyd, 1917).
P. tephroleucus (Mont.) T. W. May et A. E.
Wood, 1995, Mycotaxon 54: 148 (Bas.: Lentinus
tephroleucus Mont., 1851; Syn.: L. leprieurii Mont.,
1854; L. siparius Berk. et M. A. Curtis, 1868;
L. dentatus Bres., 1925).
P. velutinus (Fr.) Overh., 1930, J. Dept Agric.
Porto Rico 14: 353 non Fr., 1838 (Bas.: Lentinus ve-
lutinus Fr., 1830; Syn.: Lentinus fulvus Berk., 1842;
L. coelopus Lév., 1846; L. nepalensis Berk., 1854;
L. blepharodes Berk. et M. A. Curtis, 1868; L. fas-
tuosus Kalchbr. et MacOwan, 1881; L. fallax Speg.,
1883; L. castaneus Ellis et Macbr., 1896; L. holum-
brinus De Seynes, 1897; L. ssus P. Henn., 1897;
L. natalensis Van der Byl., 1924; L. thomensis
Coutinho, 1925; L. pseudociliatus Raithelhuber,
1974).
Provisional position: Lentinus hirtiformis Mur-
rill, N. Am. Fl. 9: 293, 1915; L. courtetianus Har. et
Pat., Bull. Mus. Hist. Nat. Paris 15: 88, 1909 (Zmi-
trovich, Malysheva, 2013).
34 Zmitrovich I. V. et al.
The Meruliaceae of Russia. II. Panus
Species irradiating into temperate zones. Two
species, being widespread in subtropical and tropical
regions, have been irradiated into temperate zones
of the northern and partly southern hemispheres.
They are characterized by less xeromorphic habitus
in comparison to strictly tropical species and dem-
onstrate pseudodimiticism even at rather advanced
developmental stages. Both species are distributed
throughout Russia: P. conchatus and P. lecomtei
(see below).
Key to temperate species
1. Upperside matt-subtomentose, soon glabres-
cent and cracking with formation of ne appressed
squamules; basidia 20–35 × 4.5–6 μm; basidiospores
5–6.5 × 2.2–3.5 μm, ellipsoid-cylindrical ...............
........................................................ 1. P. conchatus
‒ Upperside tomentose, then strigose; basidia
15–20 × 3.5–5 μm; basidiospores 4.5–6 × 2.5–3.7
μm, mostly ovoid ............................. 2. P. lecomtei
1. Panus conchatus (Bull.) Fr., 1838, Epicr.
Syst. Mycol.: 396.
≡ Agaricus conchatus Bull., 1787, Herb. Fr. 7:
tab. 298 : Fr., 1821, Syst. Mycol. 1: 181.
= A. carneotomentosus L., 1753, Sp. pl. 2: 1171.
= A. abelliformis Schae., 1774, Fung. bavar.
palat. nasc. 4: 20.
= A. carneotomentosus Batsch, 1783, Elench.
fung.: 89.
= A. mesentericus Batsch, 1783, Elench. fung.
(Halle): 91.
= A. carnosus Bolton, 1792, Hist. fung. Halifax,
App. 3: 146.
= A. abellatus J. F. Gmel., 1792, Syst. Nat. 2(2):
1410.
= A. inconstans Pers., 1800, Comm. Schae.
Icon. Pict.: 17.
= A. fornicatus Pers., 1801, Syn. meth. fung. 2:
475.
= A. torulosus Pers., 1801, Syn. meth. fung. 2:
475.
= Pleuropus fornicatus Gray, 1821, Nat. Arr.
Brit. Pl. 1: 615.
= Panus monticola Berk., 1851, Hooker’s J. Bot.
Kew Gard. Misc. 3: 46.
= P. vaporarius Bagl., 1865, Comm. Soc. crittog.
Ital. 2(fasc. 2): 264.
= Lentinus percomis Berk. et Broome, 1875, J.
Linn. Soc., Bot. 14(no. 73): 42.
= L. bresadolae Schulzer, 1885, Hedwigia 24(4):
141.
= Panus abelliformis Quél., 1888, Fl. Mycol.
France (Paris): 325.
= Lentinus obconicus Peck, 1906, Bull. Torrey
bot. Club 33(4): 215.
I c o n .: Malkovský (1932: g. 1‒4, ut Panus
abelliformis); Phillips (1981: g. 267d); Pegler
(1983: g. 35); Hansen, Knudsen (1992: g. 2 ut
Lentinus conchatus); Courtecuisse, Duhem (1994:
g. 135); Zmitrovich et al. (2004: g. 25); Knudsen,
Vesterholt (2008: 73D).
Basidiocarp solitary or caespitose, medium-
sized, originating from a woody substratum, of
lentioid habitus. Pileus 1.5–15 cm diam., tough
eshy, then coriaceous, applanate or depressed, of-
ten conchoid or cyathiform. Upperside pinkish-lilac
to vinaceous-purple when young, then fading from
the centre to pale-clay, ochraceous-brown or cinna-
mon, matt-subtomentose, soon glabrescent, nally
smooth and shiny or cracking at the centre to form
indenite, appressed squamules. Margin sharp, thin,
inrolling, slightly undulate or lobed, pruinose when
young, occasionally strigose. Stipe central to lateral,
0.5–4 × 0.5–3 cm, cylindric to bulbous, sometimes
tapering at the base, solid; surface initially tinted
violaceous then fading to leave a pale grey, velu-
tinate to short strigose tomentum, pubescent at the
base, nally glabrous. Context tough eshy, then
coriaceous, 1–15 mm thick at the disk, ivory-white.
Hymenophore lamellate. Gills deeply decurrent
with a ridge extending down the stipe, often slightly
anastomosing over the stipe surface, at rst viola-
ceous or purplish then cream colour, pinkish to-
wards the edge; initially very narrow but eventually
becoming broader, 2–4.5 mm wide, very crowded,
with lamellulae of four lengths; edge entire (Fig. 1).
Hyphal system dimitic with rather prolonged
pseudodimitic stage. Generative hyphae 2–4.5
μm diam., non-inating, hyaline, thin-walled,
branched, with large clamp connections. Skeletal
hyphae 2–5 μm diam., unbranched, sinuose, hya-
line, thick-walled, the pseudoskeletal hyphae of the
same diameter, with clamp connections, and refrac-
tive contents predominate in young basidiocarps.
Pileipellis an epicutis, 45–90 μm thick, of repent,
radially parallel generative hyphae 3–5 μm diam.
with a golden-brown wall. Hymenophoral trama
irregular, hyaline, of radiate construction, similar
in structure to the context. Gills-edge sterile, with
conspicuous, crowded, clavate to sublageniform
cheilocystidia 24–60 × 7–16 μm, hyaline, thin- to
thick-walled. Pleurocystidia (pseudocystidia) abun-
dant, 25–70 × 5–10(12) μm, narrowly clavate, often
sinuous and constricted, thick-walled, originating
deep in the subhymenial layer and projecting 5–20
μm above the basidia. Basidia 20–35 × 4.5–6 μm,
35
Turczaninowia 21 (3): 29–44 (2018)
Fig. 1. The most typical laterally attached morphotype of
Panus conchatus (Kalinovskaya 4552M/12): 1 – an up-
perside view; 2 – a hymenophore. Scale bar – 1 cm. Fig. 2. Association of Panus conchatus with Trametes
multicolor-coll. (Zmitrovich 2014-12).
clavate-cylindrical, 4-spored, with a basal clamp.
Basidiospores 5–6.5 × 2.2–3.5 μm, ellipsoid-cylin-
drical, hyaline with few contents, thin-walled.
On dying trees, fallen logs and branches, stumps
of many hardwoods, causing a white rot.
S u b s t r a t a: on many hardwoods, especially
Betula spp. and Populus tremula, rarely on conifers
(Pinus sylvestris).
Cultural characteristics: Hibbett et al.
(1993); Johnson, Methven (1994); Grand (2004).
General distribution: EUROPE (Austria,
Belarus, Belgium, Bulgaria, Denmark, Estonia,
Finland, France, Germany, Latvia, Lithuania,
Norway, Russia, Scotland, Spain, Sweden, Ukraine);
AFRICA (Ethiopia); ASIA (Armenia, Georgia,
Japan, Korea, Russia); NORTH AMERICA
(Canada, Mexico, USA), CENTRAL AMERICA
(Costa Rica); SOUTH AMERICA (Equador);
AUSTRALIA and OCEANIA (Australia) (Pegler,
1983; Panus conchatus.., 2018).
Distribution in Russia: see Table 1.
Exsiccates examined. Panus torulosus:
“Sweden, Bohuslan, Uddevalla, Sarven Lake, on
stump of Betula sp., X 1947. S. Woldmar LE 3722 (S.
Lundell et J. A. Nannfeldt. Fungi exsiccati Suecici
praesertim Upsalienses, N 1771)”. – “Sweden,
Västergötland, Göteborg, Naturparken, on stump
of Betula sp., 20 IX 1960. F. Karlvall LE 3723 (S.
Lundell et J. A. Nannfeldt. Fungi exsiccati Suecici
praesertim Upsalienses, N 2865)”. – “Sweden,
Småland, Femsjö parish, on stump of Betula sp.,
28 IX 1959. F. Karlvall LE 3721 (S. Lundell et J.
A. Nannfeldt. Fungi exsiccati Suecici praesertim
Upsalienses, N 2866)”. – “Russia, Khabarovsk
Region, Ragozhino vicinities, on Quercus sp.,
03 VI 1910. M. Korotkiy (M. Korotkiy. Museum
Botanicum Academiae Scientiarum Petropolitanae,
N 73)”. – “England, 15 III 1989. F. B. Delange
LE 24071, LE 24074 (R. B. G. K. Richmond.
Herbarium path/mixed debris England, N 73). –
“USSR, Bashkortostan Republic, Bashkirsky
Reserve, Drozdov Log vicinities, on stump of Pinus
sylvestris, VIII 1948. E. A. Selivanova-Gorodkova
LE 3703 (E. A. Selivanova-Gorodkova. Plantae
australiuralenses, N 459)”.
Association with Trametes multicolor.
According to our observations made on the
Karelian Isthmus clear cuttings, Panus conchatus
often settles on stumps, primarily colonized by
Trametes multicolor (Schae.) Jülich (Fig. 2).
Within ten records made for Betula stumps on the
Karelian Isthmus clear cuttings, seven ones have
contained an indication of the joint presence of
Panus conchatus and Trametes multicolor. The
latter species is an active producer of laccase and is
characterized by high growth rate (Zmitrovich et al.,
2017). Apparently, their primary delignication and
moistening of wood by T. multicolor creates a niche
for Panus conchatus, carrying out a deeper substrate
delignication. The pair in question certainly echoes
with another pair, Antrodiella pallescens/Fomes
36 Zmitrovich I. V. et al.
The Meruliaceae of Russia. II. Panus
fomentarius, described for dead wood in boreal
forests (Spirin, 2002).
Nomenclature. Two names sanctioned by
Fries (1821) were considered in the literature as a
t basionyms of this species, Agaricus conchatus
(Bulliard, 1787) and A. torulosus (Persoon, 1801).
Since both names are sanctioned by Fries and there
is no any doubt in their synonymy, the combination
based on A. conchatus name, i. e. Panus conchatus
(Fries, 1838), has a priority.
Intraspecic variability. As a neutral type
of this species (P. conchatus var. conchatus) the
more or less lateral ecotype with conchiform pileus
and rather dark wine-red to lilac-brown surface is
considered by default. Two main deviations from
such neutral type were described: 1) the chromatic
one, characterized by light-colored (red or clay-
yellow), usually conchiform pilei (P. conchatus var.
inconstans) and 2) the growth one, characterized
by entral (often bulbous) stipe, funnel-shaped cap
and strongly inrolled margin (P. conchatus var.
torulosus).
Panus conchatus var. inconstans (Pers.) Zmitr.,
Bondartseva, Perevedentseva, Myasnikov et
Kovalenko, comb. nov. (MB 824338). – Basionym:
Agaricus inconstans Pers., 1800, Comm. Schae.
Icon. Pict.: 17.
= A. conchatus var. carneotomentosus Fr., 1832,
Syst. Mycol. (Index): 11.
Persoon (1800) has characterized this taxon
as follows: «Subcespitosus, pileo carnoso tenaci
depresso integro, aut dimidiato lobato exuoso, ex
alutaceo subrufescente, lamellis subramosis basi
crispis albis subrutilesque, stipite brevi sublaterali».
I c o n .: Malkovský (1932: g. 5, ut Panus
abelliformis).
From type variety diers by clay-bu, carneous
of rufescent color of the upperside. Basidiocarps of
conchoid appearance, the stipe often reduced and
then the hymenophore is subporoid at the base. The
microstructures vary as in a neutral type.
Panus conchatus var. torulosus (Pers.) Zmitr.,
Bondartseva, Perevedentseva, Myasnikov et
Kovalenko, comb. nov. (MB 824339). – Basionym:
Agaricus torulosus Pers., 1801, Syn. meth. fung. 2:
475.
I c o n .: Malkovský (1932: g. 2, 3, 6 ut Panus
abelliformis).
From type variety diers by funnel-shaped
basidiocarps on a central stipe, often with bulbous
base, and inrolled undulating margin. The
microstructures vary as in a neutral type (Fig. 3).
Fig. 3. Panus conchatus var. torulosus (Kalinovskaya
3850M/15): 1 – an upperside view; 2 – a hymenophore;
3 – generative hypha; 4 – pseudoskeletal hypha; 5 – -
brohyphae; 6 – pseudocystidium; 7 – cheilocystidia; 8 –
basidia; 9 – basidiospores. Scale bars: 1, 2 – 1 cm, 3–9 –
10 μm.
37
Turczaninowia 21 (3): 29–44 (2018)
2. Panus lecomtei (Fr.) Corner, 1981, Beih.
Nova Hedwigia 69: 90.
≡ Lentinus lecomtei Fr., 1825, Syst. Orb. Veg. 1:
77.
= Agaricus strigopus Pers. in Gaudichaud-
Beaupré in Freycinet, 1827, Voy. Uranie., Bot.: 167.
= A. hirtus Secr., 1833, Mycogr. Suisse 2: 452.
= A. macrosporus Mont., 1837, Annls Sci. Nat.,
Bot., sér. 2 8: 370.
= Panus rudis Fr., 1838, Epicr. Syst. Mycol.:
398.
= Lentinus capronatus Fr., 1838, Epicr. Syst.
Mycol.: 389.
= A. sainsonii Lév. in Demidov, 1842, Voyage
dans la Russie Meridionale et la Crimeé, par la
Hongrie, la Valachie et la Moldavie 2: 85.
= Lentinus chaetophorus Lév., 1844, Annls Sci.
Nat. Bot. 2: 177.
= L. melanophyllus Lév., 1844, Annls Sci. Nat.
Bot. 2: 175.
= Panus lamyanus Mont., 1856, Syll. gen. sp.
crypt.: 147.
= P. homannii Fr. in Homann, 1867, Icon.
Anal. Fung., Abbild. Beschr. Pilz. 1(4): 94.
= Lentinus sparsibarbis Berk. et M. A. Curtis,
1869, J. Linn. Soc., Bot. 10(no. 45): 301.
= L. substrigosus Henn. et Shirai in Hennings,
1900, Bot. Jb. 28(3): 270.
= Panus semirudis Singer, 1936, Beih. Botan.
Centralbl. B 56: 142.
= P. fragilis O. K. Mill., 1965, Mycologia 57(6):
943.
= P. neostrigosus Drechsler-Santos et Wartchow,
2012, J. Torrey bot. Soc. 139(4): 438.
I c o n .: Malkovský (1932: g. 10‒12, ut
P. rudis); Zerova (1974: tab. 90, 1 ut P. rudis); Pegler
(1983: g. 31, ut Lentinus strigosus); Zmitrovich
et al. (2004: Tab. 4, a, b ut Panus rudis); Bulakh
(2015: g. 418).
Basidiocarp solitary or caespitose, medium-
sized, originating from a woody substratum, of
lentioid habitus. Pileus 1.5–12 cm diam., tough
eshy, then coriaceous, convex, then depressed to
infundibuliform, or laterally attached and abelli-
form to spathulate. Upperside ivory-white to stra-
mineous with prominent lilac or vinaceous tints,
fading pale-ochraceous or grayish-brown, at rst to-
mentose, then strigose (hairs 1–2 mm long), without
a clear zonation. Margin thin, inrolling, slightly un-
dulate or lobed, strongly strigose. Stipe eccentric to
lateral, 0.5–3.8 × 0.3–1.5 cm, cylindric to bulbous,
solid, sometimes reduced; surface concolorous
with the pileus, tomentose-strigose. Context tough
eshy, then coriaceous, 1–7 mm thick at the disk,
ivory-white. Hymenophore lamellate. Gills deeply
decurrent, ivory-white to ochraceous-bu, some-
times with violaceous tints; initially very narrow but
eventually becoming broader, 1–2 mm wide, very
crowded, with lamellulae of four lengths; edge en-
tire (Fig. 4).
Hyphal system dimitic with expressed
pseudodimitic stage. Generative hyphae 2–4 μm
diam., non-inating, hyaline, thin-walled, branched,
with large clamp connections. Skeletal hyphae
2–7.5 μm diam., unbranched, sinuose, hyaline,
thick-walled to subsolid, the pseudoskeletal hyphae
of the same diameter, bearing clamp connections
and refractive contents, are abundant in young ba-
sidiocarps. Pileipellis a trichodermal epicutis, 25–
45 μm thick, of repent, radially parallel generative
hyphae 3–7.5 μm diam. with a golden-brown wall.
Hymenophoral trama irregular, hyaline, of radi-
ate construction, similar in structure to the context.
Fig. 4. The most typical laterally attached morpho-
type of Panus lecomtei (Myasnikov 4472A/15): 1 –
an upperside view; 2 – a hymenophore. Scale bar – 1
cm.
38 Zmitrovich I. V. et al.
The Meruliaceae of Russia. II. Panus
Gills-edge sterile, with conspicuous, crowded, cla-
vate to sublageniform cheilocystidia 18–35 × 4–6
μm, hyaline, thin- to thick-walled. Pleurocystidia
(pseudocystidia) abundant, 25–55 × 9–13 μm,
narrowly clavate, often sinuous and constricted,
thick-walled, originating deep in the subhymenial
layer and projecting up to 40 μm above the basid-
ia. Basidia 15–20 × 3.5–5 μm, clavate-cylindrical,
4-spored, with a basal clamp. Basidiospores 4.5–6 ×
2.5–3.7 μm, ovoid to ellipsoid-cylindrical, hyaline
with few contents, thin-walled.
On dying trees, fallen logs and stumps, causing
a white rot.
Substrata: On many hardwoods, especially
Carpinus betulus, Betula spp., Fagus sylvatica,
Quercus spp., rarely on conifers (Larix spp.).
Cultural characteristics: Hibbett et al.
(1993); Grand (2004); Vargas-Isla, Ishikawa (2008);
Petre, Tănase (2013).
General distribution: EUROPE
(Bulgaria, Estonia, France, Germany, Hungary,
Portugal, Romania, Russia, Serbia, Slovakia,
Slovenia, Spain, Ukraine); AFRICA (Madagascar,
Mayotte, Uganda, Zaire); ASIA [India, Iran, Japan,
Malaysia, Myanmar (Burma), Nepal, Pakistan,
Philippines, Russia, Sri Lanka, Taiwan, Thailand,
Turkey]; NORTH AMERICA (Canada, Mexico,
USA), CENTRAL AMERICA (Costa Rica, Cuba,
Guatemala, Honduras, Nicaragua, Panama, Puerto
Rico); SOUTH AMERICA (Argentina, Brazil,
Colombia, Guiana, Paraguay, Peru, Venezuela);
AUSTRALIA and OCEANIA (Galapagos, Norfolk
Island, Papua New Guinea, Australia) (Pegler, 1983;
Panus neostrigosus .., 2018).
Distribution in Russia: see Table 2.
Exsiccates examined. Agaricus sain-
sonii: “Russia, 1842. A. Demido LE 5848 (J.
Leveillé. Voyage dans la Russie Meridionale et la
Crimeé, N 85)”. – Lentinus lecomtei: “USA, Ohio,
VII 1883. W. A. Kellermann LE 5849 (Rabenhorst–
Winter. Fungi Europaei, N 2940)”. – Panus rudis:
“Austria, trunk of Fagus sylvaticus. P. P. Strasser
LE 5854 (P. P. Strasser. Kryptogamae exsiccatae, N
1422)”. – “Georgia, Tiis Hortus Botanicus, 12 X
1923. G. Woronow LE 208198 (G. Woronow. Fungi
Caucasici, N 2127)”. – “Russia, Tomsk Region, on
stump of Betula sp., 3 III 1926. Lavrov LE 5803
(A. A. Jaczewski. Ex Herbario Instituti Mycologici
et Phytopathologici, N 81)”. – “Austria, Salzburg,
on fallen trunk of Fagus sylvatica, 1914 C. Keissler
LE 5851 (C. Keissler. Kryptogamae exsiccatae, N
1422)”. – “Baton Range, 23 II 1960. B. Lowy LE
5845 (B. Lowy. Ex Mycological Herbarium of Lou-
isiana State University)”. – “Russia, Bashkortostan
Republic, Bashkirsky reserve, Drozdov Log vicini-
ties, on stump of Betula sp., 8 IX 1946 E. A. Seliva-
nova-Gorodkova LE 5838 (E. A. Selivanova-Goro-
dkova. Plantae australiuralenses, N 2427).” – “Rus-
sia, Bashkortostan Republic, Bashkirsky reserve,
Drozdov Log vicinities, on stump of Betula sp., 8
IX 1946 E. A. Selivanova-Gorodkova LE 5834 (E.
A. Selivanova-Gorodkova. Plantae australiuralens-
es, N 2429). – Panus sainsonii: “Austria, Salzburg,
IX 1868 Dr. Sauter LE 5855 (Rabenhorst–Winter.
Fungi Europaei, N 1207)”. – “Austria, Salzburg, on
fallen trunk of Fagus. Dr. Sauter LE 5850 (F. Thü-
men. Fungi austriaci, N 212)”.
Nomenclature. Two names in application to this
peculiar species persisted in the literature for a long
time, Lentinus strigosus and Panus rudis. Since, as
it was shown, this species does not belong to the
Lentinus s. str. (Zmitrovich, Malysheva, 2013; Zmi-
trovich, Kovalenko, 2016), it became necessary a
nomenclatural adaptation of species name within the
Panus. Because of the name P. strigosus Berk. et M.
A. Curtis was preoccupied by Berkeley and Curtis
(1859) in application to another species, the name
P. rudis (Fries, 1838) continued to be persisting.
Drechsler-Santos et al. (2012) proposed new name
P. neostrigosus without any nomenclature analysis,
and this name was taken as a basis for GBIF (Pa-
nus neostrigosus .., 2018). At the same time, Cor-
ner (1981) already made a combination P. lecomtei,
basing on the description by Fries Lentinus lecomtei
in his earlier work (Fries, 1825).
Intraspecic variability. As a neutral type of
Panus lecomtei (P. lecomtei var. lecomtei), the eco-
type having small eccentric to lateral minute and
often spalled stipe is considered by default. How-
ever, the stipe shape and size are variable. The stipe
can be either central – minute, with a bulbous base
(P. lecomtei var. semirudis), or strongly elongated
(P. lecomtei var. stipitata).
Panus lecomtei var. semirudis (Singer) Zmitr.,
Bondartseva, Perevedentseva, Myasnikov et Kova-
lenko, comb. nov. (MB 824340). – Basionym: Pa-
nus semirudis Singer, 1936, Beih. Botan. Centralbl.,
Abt. B 56: 142.
From type variety is distinguished by short, but
strong and often bulbous stipe of central or sub-
central position. Upperside is moderately strigose
to glabrous at the center, color usually fading to
ochraceous-brown. The microstructures vary as in a
neutral type (Fig. 5).
39
Turczaninowia 21 (3): 29–44 (2018)
Panus lecomtei var. stipitatus (Malk.) Zmitr.,
Bondartseva, Perevedentseva, Myasnikov et Kova-
lenko, comb. nov. (MB 824341). – Basionym: Pa-
nus rudis f. stipitata Malk., 1932, Annls mycol.
30(1/2): 40.
I c o n .: Malkovský (1932: g. 12, ut Panus ru-
dis f. stipitata).
From type variety is distinguished by long lateral
or eccentric stipe and spathulate pilei. Upperside is
strongly strigose, color usually fading to cinnamon.
The microstructures vary as in a neutral type (Fig. 6).
Fig. 5. Panus lecomtei var. semirudis (LE 3660): 1 – hy-
menophore; 2 – upperside view; 3 – brohypha; 4 – gen-
erative hyphae; 5 – cheilocystidia; 6 – pseudocystidium;
7 – basidia; 8 – basidiospores. Scale bars: 1, 2 – 1 cm,
3–8 – 10 μm.
The problem of Lentinus martianoanus.
This problematic species was described by Thü-
men (1877) with reference to Kalchbrenner, and
its authentic material kept in Kew Herbarium (K).
There is a single specimen, collected on Populus
balsamifera by Martiano near Minussinsk in 1880
and distributed within exsiccate series by Thümen
(Fung. Exot. Dec. 21).
The Kew material was studied by Pegler (1983)
and described as follows: “Imbricate. Pileus later-
ally attached, 3–5 cm diam., thin, coriaceous, irreg-
ularly abelliform, depressed; surface pale yellow-
ish rufous, uniformly velutinate-strigose becoming
hispid towards the margin; margin irregular, deeply
lobed, faintly sulcate, densely ciliate with hair-like
squamules, 1–2 mm long. Lamellae decurrent, pal-
lid, narrow, about 1 mm wide, very crowded, with
lamellulae of three lengths; edge entire. Stipe lat-
eral, 1–1.5 cm × 8–12 mm, short, cylindric or com-
pressed, solid; surface concolorous with the pileus,
densely strigose, glabrescent. Context up to 2 cm
Fig. 6. Panus lecomtei var. stipitatus (Myasnikov
4350A/16): 1 – upperside view; 2 – hymenophore. Scale
bar – 1 cm.
40 Zmitrovich I. V. et al.
The Meruliaceae of Russia. II. Panus
Table 1
Herbarium data on distribution of Panus conchatus over Russian territory and its substrate preferences
Region Substrata Date of collection Collector Herbarium
numbers
European part
Karelia Republic Betula pubescens 08 X 1950 A. S. Bondartsev LE 3704
Leningrad Region unidentied substrate 19 VIII 1960 M. A. Bondartseva LE 3702
Leningrad Region Betula pubescens 20 VII 2014 I. V. Zmitrovich LE 287527
Leningrad Region Populus tremula 05 VIII 2001 I. V. Zmitrovich LE 212955
Leningrad Region Betula sp. 23 VI 1997 O. V. Morozova LE 215053
Leningrad Region unidentied substrate 07 X 2007 N. V. Psurtseva LE 265028
Leningrad Region unidentied substrate VIII 1918 V. P. Savich LE 3716
Mari El Republic Betula sp. 14 VI 1938 B. P. Vasilkov LE 3708
Mari El Republic Populus sp. 03 VII 1935 B. P. Vasilkov LE 3713
Moscow Region Betula sp. 19 VIII 1925 L. A. Lebedeva LE 3715
Moscow Region unidentied substrate 17 VI 2016 O. V. Anisimova LE 315401
Orel Region Betula sp. 06 VIII 1912 A. S. Bondartsev LE 3718
Pskov Region unidentied substrate 23 VII 2002 O. V. Morozova LE 217599
Saint Petersburg unidentied substrate 27 IX 1994 O. V. Morozova LE 227995
Stavropol Territory unidentied substrate 15 VIII 1925 A. I. Lobik LE 3711
Tver Region unidentied substrate VII 1924 L. A. Lebedeva LE 3712
Vologda Region Betula sp. 25 VIII 2002 E. S. Popov LE 246368
Urals
Perm Territory Populus tremula 15 VIII 1994 L. G. Perevedentseva PERM 118-3
Siberia
Irkutsk Region Populus tremula 15 VIII 1983 A. E. Kovalenko LE 18116
Irkutsk Region Betula sp. 20 VIII 1947 B. P. Vasilkov LE 3710
Irkutsk Region Betula sp. 26 VIII 1947 B. P. Vasilkov LE 3714
Irkutsk Region Betula sp. 27 VIII 1947 B. P. Vasilkov LE 3707
Krasnoyarsk Territory Betula sp. 11 IX 1965 A. L. Yavorskiy LE 3705
thick at the base, very thin over the hymenophore,
white, consisting of a dimitic hyphal system with
skeletal hyphae. Generative hyphae 2–5 μm diam.,
not inating, hyaline, branched, thin-walled or
occasionally with a slightly thickened wall, and
clamp-connexions. Skeletal hyphae 2–6 μm diam.,
unbranched, hyaline, with a thickened wall (up to 2
μm) and a narrow lumen, intercalary and terminal in
origin, nally tapering to an obtusely rounded apex.
Spores 4.7–6.2 × 1.7–2.5 μm, Q = 2.37, narrowly
cylindric, at times arcuate, hyaline, thin-walled,
with few contents. Basidia 16‒22 × 4.5‒5.5 μm,
clavate, bearing four sterigmata. Lamella-edge ster-
ile, forming a narrow zone of small, inconspicuous
cheilocystidia, together with scattered metuloids.
Cheilocystidia 16‒24 × 3.5×5 μm, basidioid, sinu-
ous fusoid, often constricted, with a rounded apex,
hyaline, thin-walled. Metuloids scattered to numer-
ous, on both sides and edge of lamellae, 26‒45 ×
7‒12 μm, inated clavate to fusoid-submucronate,
usually with a thickened wall (1‒2.5 μm) and res-
inous, yellowish brown contents, projecting to 25
μm beyond the basidia, sometimes surrounded by
a sheath of thin-walled, generative hyphae, 2‒5 μm
diam. Hymenophoral trama irregular, hyaline, of ra-
diate construction, comprising mostly of generative
hyphae. Subhymenial layer well developed, 14‒20
μm wide, tightly interwoven. Pileipellis a disrupted
and irregular trichodermial palisade, forming fas-
cicles of unbranched, scleried generative hyphae,
4‒6 μm diam., with an obtusely rounded apex”
(Pegler, 1983).
Since the molecular testing of any Kew material
is prohibited, only the morphological description
may be considered on, which unambiguously indi-
cates the close relationships of this taxon with Pa-
nus lecomtei-coll. Only basidiospores are something
diverse, since are not ovoid, but rather ellipsoidal-
cylindrical. Malkovský (1932) has considered Len-
tinus martianoanus as a synonym of Panus rudis,
whereas Pegler (1983) has abstained from synony-
mization procedure. In any case, this taxon belongs
to the genus Panus, although there have been at-
tempted to associate this name with the Lentinus
41
Turczaninowia 21 (3): 29–44 (2018)
Table 2
Herbarium data on distribution of Panus lecomtei over Russian territory and its substrate preferences
Region Substrata Date of collection Collector Herbarium
numbers
European part
Adygeya Republic Fagus sylvatica V 1910 N. Shestunov LE 5815, LE
5817, LE 5827
Adygeya Republic Fagus sylvatica 18 IX 2003 N. V. Psurtseva LE 241942
Kirov Region unidentied substrate 23 VII 1921 M. K. Khokhryakov LE 5799
Krasnodar Territory unidentied substrate 13 VIII 2003 N. V. Psurtseva LE 227998
Krasnodar Territory unidentied substrate 31 VII 1979 A. E. Kovalenko LE 5837
Krasnodar Territory unidentied substrate 17 VII 1976 A. E. Kovalenko LE 5829
Leningrad Region Betula pubescens 06 VII 1998 I. V. Zmitrovich LE 214737
Mari El Republic unidentied substrate 10 VI 1932 B. P. Vasilkov LE 5832
North Ossetia–Alainia
Republic
Carpinus betulus 15 V 1925 Z. Chernetskaya LE 5813
Penza Region Betula sp. 27 VII 1921 Shtukenberg LE 5841
Ryazan Region unidentied substrate IX 1960 G. K. Milberg LE 5828
Stavropol Territory unidentied substrate 24 VIII 1915 A. I. Lobik LE 5836
Voronezh Region unidentied substrate 1946 V. Ya. Chastukhin LE 5833
Urals
Perm Territory Betula pendula 25 VIII 1980 L. G. Perevedentseva PERM 118-1
Perm Territory Betula pendula 30 VIII 1985 L. G. Perevedentseva PERM 118-2
Perm Territory Betula pendula 22 VIII 1999 L. G. Perevedentseva PERM 118-4
Perm Territory Betula pendula 04 VIII 2004 L. G. Perevedentseva PERM 118-5
Siberia
Altai Republic Betula sp. 17 VIII 2008 N. V. Psurtseva LE 254518
Altai Republic Betula sp. 18 VIII 2008 N. V. Psurtseva LE 254519
Altai Republic Populus sp. VIII 1937 R. Singer LE 5842
Chita Region unidentied substrate V 08.1910 no data LE 5812
Irkutsk Region Betula sp. 20 VIII 1921 T. Smirnov LE 5804
Irkutsk Region Betula sp. 04 IX 1947 B. P. Vasilkov LE 5805
Tumen Region unidentied substrate 20 VI 1914 Varentsov LE 5831
Yakutia Republic Larix sp. 10VIII1908 N. A. Palchevskiy LE 5844
Far East
Amur Region Quercus sp. 17VII1959 B. A. Tomilin LE 5791
Amur Region unidentied substrate 13VI1910 M. Korotkiy LE 3663
Primorye Region unidentied substrate 1913 V. L. Komarov LE 3664
Primorye Region unidentied substrate 29VI1913 V. L. Komarov LE 5840
Primorye Region Betula dahurica 19VII1934 B. Kolesnikov LE 5797
Primorye Region Quercus sp. 15 VII 1952 E. V. Volkova LE 5808
Sakhalin Region Larix decidua 1960 B. P. Vasilkov LE 5839
Sakhalin Region Sorbus aucuparia 29 VIII 1954 M. G. Tarabaev LE 5820
piloso-squamulosus Lj. Vassilieva (current name is
Lignomyces vetlinianus) (Bulakh, 2015).
Сonclusion
The Panus represents rather well-delimited ge-
nus belonging to merulioid phylogenetic radiation,
whose morphotype on essential features of its or-
ganization is trametoid, but supercial habitual fea-
tures make it closer to the lentinoid one. One of its
essential features is the abundance of brohyphae
which forms a textura intricata, slowly growing
basidiocarps and strictly lamellate hymenophore.
Apparently, such an adaptive structure was gener-
ated under the inuence of arid and warm climate
conditions at the dierent regions of the planet, and
only 2 species, Panus conchatus, and P. lecomtei,
have been irradiated into temperate latitudes, in-
cluding Russian territory. The substrate spectrum
of these fungi is determined by their insensitivity to
the substrate moistening and best ability to colonize
42 Zmitrovich I. V. et al.
The Meruliaceae of Russia. II. Panus
hardwood, so the greatest number of their nds can
be made on stumps and large remnants of stand-
formers of corresponding forest areas. In Russia, a
reliable association of Panus species to Betula spp.
and Populus spp. was revealed. An ecotypic dier-
entiation of the genus Panus is related to the qual-
ity of substrate colonized. The basidiocarps, grow-
ing over top cuts of the stumps, are characterized
by strong central stipe (P. conchatus var. torulosus,
P. lecomtei var. semirudis), whereas basidiocarps
with sublateral attachment are common on fallen
logs. Certain chromatic aberrations (P. conchatus
var. inconstans) are associated with an insolation
regime of the habitat. During last years, the Panus
representatives have attracted an interest in biomed-
ical research development. Their resource potential
estimation should proceed from the fact that within
Russian territory, such areas as Middle Belt of Eu-
ropean Russia, North Caucasus, Altai and other re-
gions of Southern Siberia are promising for replen-
ishing the strains of P. conchatus and P. lecomtei.
Acknowledgements
The authors are very grateful to N. I. Kalinovs-
kaya for kindly provided photos of Panus concha-
tus. The work by I. V. Zmitrovich, M. A. Bondartse-
va, and A. E. Kovalenko was carried out in canvas
of the State Task of Komarov Botanical Institute of
the Russian Academy of Sciences named as “Bio-
diversity and spatial structure of fungi and myxo-
mycetes communities in natural and anthropogenic
ecosystems” (АААА-А18-118031290108-6). All
experimental work has been fully implemented on
the equipment of the Center for collective use of sci-
entic equipment “Cellular and molecular technol-
ogy of studying plants and fungi” at the Komarov
Botanical Institute of the Russian Academy of Sci-
ences (Saint Petersburg).
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