CC BY
0
ISSN 1560-7259 (print edition)
TURCZANINOWIA
ISSN 1560-7267 (online edition)
УДК 582.28:581.961(470+571)
The veiled stinkhorn Dictyophora sibirica in Russia - taxonomic position, epitypification and its conspecifity with Phallus ultraduplicatus
Yu. A. Rebriev1*, N. N. Kudashova2, 4, S. I. Gashkov2, 5, I. A. Kazartsev3
1 Southern Scientific Centre of Russian Academy of Sciences, Chekhova St., 41, Rostov-on-Don, 344006, Russian Federation. E-mail: rebriev@yandex.ru; ORCID iD: https://orcid.org/0000-0001-5156-8593
2 Research Tomsk State University, Lenina St., 36, Tomsk, 634050, Russian Federation
3 All-Russian Institute of Plant Protection (FSBSI VIZR), Podbelskogo Shosse, 3, Pushkin, St. Petersburg, 196608, Russian Federation. E-mail: ikazartsev@vizr.spb.ru; ORCID iD: https://orcid.org/0000-0003-4500-0154
4E-mail: agaphnad@mail.ru; ORCID iD: https://orcid.org/0009-0003-5737-891X
5E-mail:parusmajorl@rambler.ru; ORCID iD: https://orcid.org/0000-0001-9620-5147
Keywords: gasteromycetes, ITS nrDNA sequences, molecular phylogeny, mycobiota, mycological collections, stinkhorn.
Summary. The article presents the results of studying a lectotype specimen of Dictyophora sibirica. The abolition of the genus name Dictyophora justifies the transfer of this taxon to the genus Phallus, with a new combination being Phallus sibiricus. A recently collected specimen from the locus classicus corresponds to the original description and is designated as an epitype due to the impossibility of sequencing the DNA of the holotype. Synonymy between Phallus sibiricus and P. ultraduplicatus is proposed, based on morphology and comparison of ITS sequences.
Сетконоска Dictyophora sibirica в России - таксономическое положение, эпитипификация и синонимия с Phallus ultraduplicatus
Ю. А. Ребриев1, Н. Н. Кудашова2, С. И. Гашков2, И. А. Казарцев3
1 Южный научный центр РАН, пр. Чехова, д. 41, г. Ростов-на-Дону, 344006, Россия
2 Научно-исследовательский Томский государственный университет, проспект Ленина, д. 36, г. Томск, 634050, Россия
3 Всероссийский научно-исследовательский институт защиты растений, шоссе Подбельского, д. 3, г. Пушкин, Ленинградская обл., 196608, Россия
Ключевые слова: веселка, гастеромицеты, микобиота, микологические коллекции, молекулярная филогения, ITS-последовательность ярДНК.
Аннотация. В статье изложены результаты изучения типового образца Dictyophora sibirica. Упразднение родового названия Dictyophora оправдывает перенос этого таксона в род Phallus с новой комбинацией Phallus sibiricus. Недавно собранный образец из locus classicus соответствует первоначальному описанию и обозначен как эпитип из-за невозможности секвенирования ДНК голотипа. Предложена синонимия Phallus sibiricus и P. ultraduplicatus, основанная на морфологии и сравнении последовательностей ITS.
Поступило в редакцию 28.06.2023 Принято к публикации 02.11.2023
Submitted 28.06.2023 Accepted 02.11.2023
Introduction
Phallus Junius ex L. is a widespread genus of ba-sidiomycetes, with highest species diversity in tropical and subtropical regions. The genus Dictyophora Desv., which included species with indusium-bea-ring basidiomes, is united with Phallus according to morphological (Kreisel, 1996; Calonge, 2005) and phylogenetic (Li et al., 2016; Melanda et al., 2021) studies. New species are often described in different parts of the world in recent times (Cabral et al., 2019; Li et al., 2002, 2014; Moreno et al., 2009; Song et al., 2018; etc.). There are 43 species known by now (Li et al., 2021). As pointed by Cabral et al. (2019, p. 104), "few morphological characters are available to delimit species in Phallus. In addition, most of the widely used diagnostic characters - such as colour and sizes - show high plasticity...". The difficulty of reliable identification of Phallus specimens lies in the fact that mature basidiomes are very short-lived. After the spore-bearing part (gleba) exposed, it is partially consumed within a very short period (typically less than a day) by insects attracted to a specific smell, partially adheres to their bodies and spreads radially. The remaining basidiomes are quickly destroyed by invertebrates and microorganisms. The basidiomes strongly change its shape and color during herbarization, and, therefore, it is desirable to photograph them in natural conditions. Taking a sample for DNA analysis in matured basidiomes is difficult. Only gleba and mycelial strands are suitable for analysis, but in both cases the risk of contamination of foreign DNA is extremely high. For these purposes, it is more convenient to use enclosed basidiomes in "egg" stage, but during their herbari-zation we get practically no information about mac-romorphology. Therefore, to accurately ascertain the identity of Phallus specimens, the presence of several basidiomes at various stages of maturity and their photo are often required.
For a long time, the prevailing opinion was that specimens of stinkhorns with indusium found in Europe belongs to Dictyophora duplicata (Bosc) E. Fisch. (current name Phallus indusiatus Vent.) (Ulbrich, 1932; Flora CSR, 1958). A detailed morphological study of European specimens revealed their obvious differences from P. duplicatus leading to the description of a new variation P. impudicus var. pseudoduplicatus O. Andersson (Andersson, 1989). According to Kreisel (Kreisel, Hausknecht, 2009), this variation is also common in Africa and presumably in Asia. However, this view requires clarification using molecular genetic methods. For example,
recent studies by Cabral et al. (2019) suggest that the veiled stinkhorn P. indusiatus might conceal a significant number of undiscovered species.
The species diversity of the genus Phallus in Russia is estimated at 6 species (unpublished data). Previously, all specimens with indusium collected in Russia were identified as Dictyophora duplicata (Vasilkov, 1954; Sosin, 1973; Rtischeva, 1991; Pero-va, Gorbunova, 2001; Bulakh, 2015) or as Phallus duplicatus (Dudka et al., 2004). Based on these data, P. duplicatus was listed in the Red Data Book of the Russian Federation (Rebriev, 2008) as a rare but widely distributed species. Under the same name, the indusiate Phallus are included in many regional Red Data Books of the Russian Federation (Kolcha-nov, Maslova, 2004; Petrov, Matosova, 2010; Gorbu-nova, 2018; etc.). The name P. impudicus var. pseudoduplicatus is listed in Red Books of the Krasnodar Territory (Rebriev, 2017) and the Republic of North Ossetia - Alania (Rebriev, Nikolaev, 2022).
In 1936, Dictyophora sibirica Lavrov was described as new for science species (Lavrov, 1936). There are two syntypes were pointed: the first from vicinity of the rural locality Chemal (the Republic of Altai), and the second from vicinities of the Tomsk city (Tomsk Region). This name was previously classified as synonym of Phallus duplicatus (= Dictyophora duplicata) by B. P. Vasilkov (1954), but without any type specimens study. Later, this taxon was sometimes indicated in publications devoted to the biodiversity of the Tomsk Region (Milovidova et al., 1980; Rare and endangered., 1984). In the article about the biodiversity of the gasteroid basidiomyce-tes of the Tomsk Region indusiate stinkhorns were named as Phallus impudicus var. pseudoduplicatus (Agaphonova et al., 2011).
The accumulation of new data on the taxonomic diversity of Phallus in the Russian Federation made facilitated clarification of the taxonomic status of some specimens with indusium collected in the Asian part. Molecular genetic methods were utilized to identify specimens from Novosibirsk Region (Rebriev et al., 2020), Tomsk Region (Kudashova et al., 2023) and Primorye Territory (Rebriev et al., 2022), and they were identified as P. ultraduplicatus X. D. Yu, W. Lv, S. X. Lv, Xu H. Chen et Qin Wang. This taxon was pointed for Krasnoyarsk Territory also but without molecular data (Kryuchkova, Goncha-rova, 2021).
Clarification of the true taxonomic position of Dictyophora sibirica Lavrov has been of significant interest. In the V. L. Komarov's Mycological Herbarium of the Botanical Institute of the Russian Aca-
demy of Sciences, a specimen LE 1582 was found and the information on its label is identical to one of the two specimens mentioned in the protologue. This specimen collected in the vicinity of Tomsk in 1933 was chosen by us as a lectotype. The objective of this study was to clarify the taxonomic status of the Russian indusiate stinkhorns specimens and current taxonomic position of D. sibirica.
Despite the well-preserved basidiome, all attempts to make a molecular genetic analysis of the type specimen of D. sibirica were unsuccessful. This is probably due to the long storage of the sample and/or its drying at high temperature. But the application of molecular data in phylogenetic analyses to delimit taxa has become a widespread, beneficial practice (Aime et al., 2021). Epitypification with sequencing of newly collected specimens is the most efficient way for unambiguous species delimitation (Buyck et al., 2019). Therefore, according to 9.9 of the "International Code of Nomenclature..." (Turland et al., 2018), an epitype was chosen. Based on morphological similarity, we designate as epitype the following specimen collected in locus classicus: LE F-348635.
Methods
Morphology
The specimens examined were obtained from the Mycological Herbarium of the Komarov Botanical Institute RAS (LE F) and from Yu. Rebriev's own collection. Basidiospores mounted in 5 % KOH solution were examined under the light microscope Mikmed-6 (LM). Light microscope photography has been implemented with the equipment of the Centre for cellular and molecular technology of studying plants and fungi at the Komarov Botanical Institute RAS (St. Petersburg).
Phylogenetic analyses
DNA was extracted from the gleba of the dried immatured basidiomes (egg stage) using a standard CTAB-chloroform method (Doyle J. J., Doyle J. L., 1987). Subsequent amplification of internal transcribed rDNA spacers (ITS) was carried out using ITS1/ITS4 primer pair following the instructions provided by the authors (White et al., 1990). Amplification products were separated by electrophoresis in 1 % agarose gel stained with ethidium bromide. Fragments of the desired length were excised and purified using silica-based procedure (Malferrari et al., 2002). Sequencing PCR was performed using the Sanger method (Sanger et al., 1977) with the BigDye Terminator v. 3.1 Cycle Sequencing Kit (Applied
Biosystems, Thermo Fisher Scientific, Waltham, MA, USA), and nucleotide sequences were determined on an ABI PRISM 3500 genetic analyzer.
The nucleotide sequences were checked and manually edited using SeqScape V3.0 or Vector NTI Advance 11.5.1 software. Subsequently, they were compared with sequences deposited in GenBank using the BlastN search (Altschul et al., 1990). The newly obtained ITS sequences were deposited in GenBank under unique accession numbers which listed in the specimen's citation.
Five new ITS sequences of Phallus spp. were produced for our study (Table 1). Additionally, 26 sequences were retrieved from the GenBank. The specimens CCTR 6759 (GenBank sequence 0P787980) was collected together with LE F-348635 and probably formed on a common mycelium. But the sequence of the second sample turned out to be longer and of better reading quality. Itajahya galeri-culata was chosen as the outgroup. The genus Itaja-hya was recognized as sister to Phallus (Melanda et al., 2021). Sequences were aligned in MAFFT (Katoh et al., 2019), then the alignment was manually optimized with MEGA 6 (Tamura et al., 2013).
The ITS rDNA phylogenetic tree (Fig. 1) was obtained using MrBayes v. 3.2.1 (Ronquist et al., 2012) using the GTR+I+G model with 5 M generations. The best tree was visualized in FigTree v. 1.2.3 and edited in Adobe Illustrator CS5 (Fig. 1).
Results
Molecular phylogeny
0ur molecular data based on ITS sequences (Fig. 1) showed that all indusiate specimens from the Asian part of the Russian Federation are grouped with the Phallus ultraduplicatus sequences and formed own clade (PP = 1). These data are consistent with our previous phylogenetic data (Rebriev et al., 2020, 2022). The specimen YuR 3865 with indusium from European Russia is clustered with P. impudi-cus-clade.
Morphology
The indusiate specimens from the Asian part of the Russian Federation are fairly uniform in morphology. Minor variations in the size of the stipe, cup and indusium can be explained by the influence of environmental factors and growing conditions. All examined specimens correspond to the description of P. ultraduplicatus (Adamcik et al., 2015). It is very important to note that the basidiomes of stinkhorns significantly decrease in size when dried. This is evident when comparing the size of the specimen LE F-348635 in its fresh and dried states (table 2).
Table 1
Taxa, vouchers, locations, and GenBank accession numbers of DNA sequences used in the study
Taxon ID (Specimen) Country GB no. ITS Reference
Dictyophora indusiata (Vent.) ASI 32011 ? Republic of Korea AF324160 Adamclk et al., 2015
Desv.
" " ASI 32001 ? Republic of Korea AF324172 Adamclk et al., 2015
D. rubrovolvata M. Zang, D. G. YZS047 China KF939507 -
Ji et X. X. Liu
" " YZS018 China KF939513 -
D. phalloidea Desv. - ? Republic of Korea AF324162 Adamclk et al., 2015
Itajahya galericulata Moller KSRF-0014 India MF506819 Melanda et al., 2021
Phallus denigricans T. S. Cabral, INPA272383 Brazil MG678486 Cabral et al., 2019
B. D. B. Silva et Baseia*
P. dongsun T. H. Li, T. Li, Chun GDGM 75402 China NR_171851 Li et al., 2020
Y. Deng, W. Q. Deng et Zhu L.
Yang*
P. fragrans M. Zang HKAS126427 China 0Q025165 -
P. hadriani Vent. SR1659 India MT151615 -
" " GDGM 83732 China MW031862 -
" " JLF7894 USA MT101866 -
" " YuR 180 Russia, Rostov Oblast MG678525 Cabral et al., 2019
P. haitangensis H. Li Li, P. E. HKAS 88199 China KU705384 Li et al., 2016
Mortimer, J. C. Xu et K. D. Hyde
P. impudicus L. G.M. 2015-08-15.10 Luxembourg 0P603024 -
YuR 3865 Russia, Moscow Oblast OR708694 -
" " - Poland MZ410690
P. indusiatus Vent.* INPA264931 Brazil MG678502 Cabral et al., 2019
P. purpurascens T. S. Cabral, B. UFRN Fungos:2808 Brazil NR_166233 Cabral et al., 2019
D. B. Silva et Baseia*
P. rigidiindusiatus T. Li, T. H. Li GDGM 81196 China NR_175754 Li et al., 2021
et W. Q. Deng*
P. rugulosus (E. Fisch.) Lloyd TNS-F-46049 Taiwan MF372142 Pereira et al., 2017
P. serratus H. Li Li, L. Ye, P. E. HKAS:78341 China KF052623 Adamclk et al., 2015
Mortimer, J. C. Xu et K. D. Hyde
P. ultraduplicatus X. D. Yu, W. HMAS:253050 China KJ591584 Adamclk et al., 2015
Lv, S. X. Lv, Xu H. Chen et Qin
Wang*
« « HMAS:253051 China KJ591585 Adamclk et al., 2015
« « YuR 3374 Russia, Novosibirsk Oblast MK965097 Rebriev et al., 2020
« « LE-BIN 4670 Russia, Primorsky Territory 0L764905 Rebriev et al., 2022
« « CCTR 6759 Russia, Tomsk Oblast 0P787980 Kudashova et al., 2023
« « LE F-348633 Russia, Krasnoyarsk Krai OR708542 -
« « LE 324109 Russia, Irkutsk Oblast OR708543 -
« « YuR3715 Russia, Republic of Buryatia OR708544 -
" " * LE F-348635 Russia, Tomsk Oblast OR712348 -
Note. The new sequences are given in bold. Type specimens are marked with asterisk (*).
Table 2
Comparison of the sizes of basidiomes from types of Dictyophora sibirica and Phallus ultraduplicatus with some studied specimens
P. ultraduplicatus (Adamcik et al., 2015) P. sibiricus epitype LE F-348635 D. sibirica (Lavrov, 1936) P. sibiricus YuR 3714
s Immature basidiome 70-80 x 80-90 mm 45-47 x 52-70 mm 60-80 mm
e s o h di Mature basidiome / Receptacle 190-250 mm tall (130)140-160(200) mm 150-200 mm
si ra Pileus 40-50 mm high 38-44-53 mm
Ph JD Indusium 20-40 mm long 25-40(45) mm
s Immature basidiome 43-48 mm 50-60 mm 32 x 48 mm
e s di Mature basidiome / Receptacle 100-130 mm 100-200 mm 95-150
^ 1 Pileus 27-45 mm 25 mm 30-40
Indusium 32-35 mm 15-20 mm 30-43
100
100
n
a
is %
#
5
I
a
100
87
Phallus ultraduplicatus KJ591584* Phallus ultraduplicatus KJ591585 Phallus ultraduplicatus OP787980 Phallus ultraduplicatus OL764905
- LE F-348633_P/ifl//MS ultraduplicatus OR708542 LE 324109 Phallus ultraduplicatus OR708543 YuR 3715 Phallus ultraduplicatus GR708544
- LE F-348635 Phallus ultraduplicatus OR712348* Phallus idtraduplicatus MK965097
Phallus impudicus OP603024 Phallus impudicus MZ410690 YuR 3865 Phallus impudicus OR708694 Phallus hadricmi MT151615 Phallus hadriani MW031862 Phallus hadriani MT101866 Phallus hadriani MG678525 Phallus fragrans OQ025165 Phallus dongsun NR_171851 * |- Dictyophora rubrovolvata KF939507 ■ Dictyophora rubrovolvata KF939513 Dictyophoraphalloidea AF324162 Dictyophora indusiata AF324160 Dictyophora indusiata AF324172 Phallus haitangensis KU705384
gg i— Phalluspurpurascens NR_166233* '-Phallus denigricans MG67 8 4 865t
100
91
100
8C
59
67
97
100
87
Phallus indusiatus MG678502*
-Phallus sctrains KF052623 -Phallus rigidiindusiatus NR_175754* -Phallus nigidosus MF372142
Itajahya galericulata MF506819
Fig. 1. ITS phylogenetic tree generated using MrBayes v. 3.2.1 under GTR + I + G model for 5 M generations. The GenBank accession numbers are indicated after each species name. Support values are indicated on the branches (posterior probabilities). The novel species is shown in bold text. Type specimens are pointed with asterisk.
The dimensions of the fresh basidiomes LE F-348635 are in good agreement with the description of P. ultraduplicatus, while the measurements of the same basidiomes in the dried state are closer to the description of Dictyophora sibirica (Lavrov, 1936). Our measurements of the type specimen of D. sibirica correspond to the description of this species. This confirms that already dried material was used in the Latin language description (along with additional measurements of immature basidiomes and receptacle in Russian language description). Based on these data, we conclude that morphological descriptions of D. sibirica and P. ultraduplicatus exhibit notable similarities.
Dictyophora sibirica type LE 1582 study (Fig. 2).
The lectotype is one basidiome 8.8 cm high, without volva and with a pileus and upper part of receptacle covered by peridium. The pileus slightly asymmetric, up to 2.5 cm high. The receptacle is broken, the widest in the central part and tapering to both ends, 1.5 cm thick in widest upper part and 1.1 cm at the bottom. The indusium is partially visible from under the peridium, up to 2.5 cm.
Basidiospores 4.0-4.5 x 1.8-2.2 ^m, oblong, smooth, thin-walled, hyaline.
Fig. 2. The image from protologue (a), specimen (b), original label (c), and basidiospores (d) of Dictyophora sibirica lectotype LE 1582 (the coin diameter is 22 mm).
Original diagnosis:
"Tota fructificatione immatura ovoidea vel subsphaeroidea 5-6 cm. diam., ab exteriore parte alba, intus gelatinosa, infra funiculos albos radiciformibus hypharum instructa. Receptaculo cylindraceo 10-20 cm. longo, 2-3 cm crasso, albo, elastico, intus inano, superne attenuato et infra volvam obducto; indusio candido vel roseolo, conico, 15-20 mm longo, basi 50-60 mm lato, retiformi, paullum (an 3-4 mm) supra inferioris pilei marginis altitudinem e stipite oriente; indusii interstitiis subrotundis vel irregulariter polygonis, in superiore indusii parte 2.5-3 mm diam., in inferiore parte 0,5-1 mm. metientibus; pileo olivaceo-virido campanulato vel subconico, 25 mm alto, basi duplo lato, cum stipitis apice limbo annuliformi recurvato conjuncto, extus cavernoso-reticulato; cavernis pilei subrhombeis 5 x 3.3 x 2 mm; sporis ellipsoideis 34 x 2 micr. viridis, polare 2 guttulatis, levibus; epi-sporio 0.5-0.7 micr. crasso, hyalino".
Type collection's details: "Russian Federation, Tomsk Region, Stepanovka village near the Tomsk. 15 IX 1933. E. F. Markidonova" (LE 1582) (Lavrov, 1939).
Notes: The Stepanovka village currently is a microdistrict of Tomsk city. The distance to the point of the epitype collection in the vicinity of the Kolarovo village is about 12 km.
Description of epitype, based on fresh specimen.
Immature basidiome ovoid to subglobose, 60-70 mm diam, whitish, felted, with whitish long rhizomorph. Mature basidiome with white to dirty-white, cylindrical, hollow, pliant and finely porous receptacle (130)140-160(200) mm tall and up to 35 mm diam. Pileus conical 38-54 mm high, with apical perforated disc, reticulate, whitish under the gleba. Gleba dark greenish olive, gelatinous, with strong odor. Indusium 25-40(45) mm long, white, fragile, slightly lagging behind the receptacle or sticking and then often breaking with growth, with elongated to roundish pores, about 4 mm diam under the pileus, 2-2.7 mm diam in central part and becoming gradually smaller to bottom margin entire.
Basidiospores 4.0-5.0 x 1.8-2.2 ^m, oblong, smooth, thin-walled, hyaline (Fig. 3c).
Fig. 3. The fresh basidiomes (a), dried basidiomes (b) and basidiospores (c) of Phallus sibiricus epitype LE F-348635 (the coin diameter is 18.75 mm).
Epitype collection's details: "Russian Federation, Tomsk Region, Tomsky District, vicinities of Kolarovo village, 56°21'00.0"N, 84°57'00.0"E, mixed forest with Pinus sylvestris, P. sibirica, Betula pendula, Populus tremula, and Padus avium, Sorbus sibirica, Lonicera tatarica in undergrowth, on litter and humified wood near. 27 VIII 2022. N. N. Kudashova, S. I. Gashkov, V. N. Stepanov" (LE F-348635; ITS GB OR712348, tef1-a 0R727902, ATP6 0R727901). Abundant fructification with the more 50 basidiomes (Fig. 3a).
The epitype specimen agrees with the Lavrov's original description of Dictyophora sibirica.
Additional specimens examined.
"Novosibirsk Region, Novosibirsk City, the territory of the Central Siberian Botanical Garden of the Siberian Branch of the Russian Academy of Sciences, 54°49'12"N, 83°06'36"E, planting of Quercus spp., on soil. 29 VII 2017. D. V. Ageev" (LE F-348632; ITS GenBank MK965097). Unripened basidiome on the egg stage and destroyed mature basidiome.
"Krasnoyarsk Territory, Kuraginsky District, 1 km northwest from Zharovsk village, edge of the forest, 53°46'48"N, 93°53'24"E, on soil and litter under Picea sp. 18 VIII 2016. I. Yu. Krom" (LE F-348633; ITS GB OR708542). Unripened basidiomes on the egg stage.
"Irkutsk Region, Irkutsk city, the right bank of the Irkut river, near the mouth of the Kaya river, the territory of the «Angara» resort, 52°17'24"N, 104°14'24"E, thickets of shrubs in a Pinus-domina-ted forest, on soil. 11 IX 2019. I. V. Enushchenko" (LE 324109; ITS GB 0R708543).
"Republic of Buryatia, Tunkinsky District, vicinities of Arshan village, on soil. 25 VIII 2019. E. S. Kurlov" (YuR 3715; ITS GB 0R708544). Unripened basidiomes on the egg stage.
Sequencing of the ITS1-5.8S-ITS2 region resulted in a sequence identical to that of the Phallus
ultraduplicatus holotype (GenBank KJ591584). The morphological description of Phallus ultraduplicatus (Adamcik et al., 2015) coincides with the protologue of Dictyophora sibirica as well as with the epitype and additional specimens.
Thus, the morphological and genetic identity of Phallus ultraduplicatus and P. sibiricus type material has been demonstrated. According to the principle of priority stated in the Article 11.4 of the International Code of Nomenclature (Turland et al., 2018), it is necessary to maintain P. sibiricus as the valid name for the taxon and consider P. ultraduplicatus as a synonym.
Taxonomic conclusion
According to H. Kreisel (1996), the genus Dictyophora was included in Phallus s. l. So, we introduce the following new combination. Since we consider Phallus ultraduplicatus is adequately treated as a synonym of the P. sibiricus.
Phallus sibiricus (Lavrov) Rebriev, comb. nov.
Basionym: = Dictyophora sibirica Lavrov, 1936, in Trudy Biologicheskogo nauchno-issledovateFskogo instituta 2: 46.
= Phallus ultraduplicatus X.-D. Yu, W. Lv, S. X. Lv, Xu H. Chen et Qin Wang, 2015, Crypt. Mycol. 36(2): 146.
Mycobank: MB 850579.
We designate as epitype the following collection: LE F-348635.
Mycobank: MB 254698.
Acknowledgments
The study was carried out within the frame of government assignments for the South Science Center RAS (project no. 122020100332-8).
REFERENCES / ЛИТЕРАТУРА
Adamcik S., Cai L., Chakraborty D., Chen X.-H., Cotter H. Van T., et al. 2015. Fungal Biodiversity Profiles 1-10. Cryptogamie, Mycologie 36(2): 21-166.
Agaphonova (Kudashova) N. N., Rebriev Y. A., Gashkov S. I. 2011. Gasteroid Basidiomycetes of the Tomsk Region. Mikologiya i fitopatologiya [Mycology and phytopathology] 45(3): 221-227. [In Russian] (Агафонова (Кудашо-ва) Н. Н., Ребриев Ю. А., Гашков С. И. Гастероидные базидиомицеты Томской области // Микология и фитопатология, 2011. Т. 45, вып. 3. С. 221-227).
Andersson O. 1989.The distribution and ecology of Phallus impudicus in the Nordic countries. Svensk Botanisk Tidskrift 83 (4): 219-41.
Bulakh E. M. 2015. Griby lesov Dalnego Vostoka Rossii [Mushrooms of the forests of Russian Far East]. Vladivostok: Dalnauka. 404 pp. [In Russian] (Булах Е. М. Грибы лесов Дальнего Востока России. Владивосток: Дальнаука, 2015. 404 с.).
Buyck B., Henkel T. W., Hofstetter V. 2019. Epitypification of the Central African Cantharellus densifolius and C. lu-teopunctatus allows for the recognition of two additional species. MycoKeys 49: 49-72. DOI: 10.3897/ mycokeys.49.32034
Cabral T. S., Silva B. D. B., Martín M. P., Clement C. R., Hosaka K., Baseia I. G. 2019. Behind the veil - exploring the diversity in Phallus indusiatus s. l. (Phallomycetidae, Basidiomycota). MycoKeys 58: 103-127. DOI: 10.3897/ mycokeys.58.35324
Calonge F. D. 2005. Clave provisional para identificar las especies de Phallus [A tentative key to identify the species of Phallus]. Boletín de la Sociedad Micológica Madrid 29: 9-18. [In Spanish].
Doyle J. J., Doyle J. L. 1987. A rapid DNA isolation procedure for small quantities of fresh leaf tissue. Phytochem. Bull. 19: 11-15.
Dudka I. O., Geljuta V. P., Tihonenko Ju. Ja., Andrianova T. V., Gajova V. P., Pridjuk M. P., Dzhagan V. V., Isikov
V. P. 2004. Gribi prirodnikh zon Krimu [Fungi of the Crimean Peninsula]. Ki'iv: Fitosotsiotsentr. 452 pp. [In Ukrainian] (Дудка I. О., Гелюта В. П., Тихоненко Ю. Я., Андрiанова Т. В., Гайова В. П., Придюк М. П., Джаган В. В., Шков В. П. Гриби природних зон Криму. Кшв: Фггосощоцентр, 2004. 452 с.).
Flora CSR. 1958. B. 1. Gasteromycetes. A. Pilat (ed.). Praha. 836 pp.
Gorbunova I. A. 2018. Dictyophora duplicata (Bosc) E. Fisch. In: Red Data Book of the Novosibirsk Oblast: Animals, plants and fungi. Novosibirsk: Printing house of Andrey Khristolyubov. P. 541. [In Russian] (Горбунова И. А. Сетко-носка сдвоенная - Dictyophora duplicata (Bosc) E. Fisch. // Красная книга Новосибирской области: Животные, растения и грибы. Новосибирск: Типография Андрея Христолюбова, 2018. С. 541).
Kolchanov A. F., Maslova E. V. 2005. Dictyophora duplicata (Bosc) Fischer. In: Krasnaya kniga Belgorodskoy oblasti. Redkiye i ischezayushchiye rasteniya, griby, lishayniki i zhivotnyye [Red Data Book of the Belgorod Oblast. Rare and endangered plants, fungi, lichens and animals]. Belgorod: Belgorod State University. P. 231. [In Russian] (Колчанов А. Ф., Маслова Е. В. Диктиофора сдвоенная (Сетконоска сдвоенная) Dictyophora duplicata (Bosc) E. Fisch. // Красная книга Белгородской области. Редкие и исчезающие растения, грибы, лишайники и животные. Белгород: Изд-во Белгородского гос. ун-та, 2005. С. 231).
Kreisel H. 1996. A preliminary survey of the genus Phallus sensu lato. Czech Mycology 48: 273-81.
Kreisel H., Hausknecht A. 2009. The gasteral Basidiomycetes of Mascarenes and Seychelles 3. Some recent records. Österreichische Zeitschrift für Pilzkunde 18: 149-159.
Kryuchkova O. E., Goncharova N. V. 2021. Rare gasteromycete Phallus ultraduplicatus X. D. Yu, W. Lv, S. X. Lv, Xu H. Chen & Qin Wang in the "Krasnoyarskie Stolby" National park. Conifers of the boreal area 39(6): 457-461. [In Russian] (Крючкова О. Е., Гончарова Н. В. Редкий гастеромицет Phallus ultraduplicatus X. D. Yu, W. Lv, S. X. Lv, Xu H. Chen & Qin Wang в Национальном парке «Красноярские столбы» // Хвойные бореальной зоны, 2021. Т. XXXIX, № 6. С. 457-461).
Kudashova N. N., Vaishlya O. B., Gashkov S. I., Karbysheva K. S. 2023. Phallus ultraduplicatus in the Tomsk region. Tomsk State University Journal in press. [In Russian] (Кудашова Н. Н., Вайшля О. Б., Гашков С. И., Карбышева К. С. Phallus ultraduplicatus в Томской области // Вестник ТГУ Биология, 2023. В печати).
Lavrov N. N. 1936. Civis novus mycoflorae sibiricae - Dictyophora sibirica n. sp. Trudy biologicheskogo nauchno-issledovatelskogo instituta [Proceedings of the Biological Research Institute] 2: 41-47. [In Russian] (Лавров Н. Н. Новый представитель сибирской микологической флоры Dictyophora sibirica n. sp. // Труды Биологического научно-исследовательского института, 1936. Т. 2. С. 41-47).
Li H., Ma X., Mortimer P. E., Karunarathna S. C., Xu J., Hyde K. D. 2016. Phallus haitangensis, a new species of stinkhorn from Yunnan province, China. Phytotaxa 280: 116-128. DOI: 10.11646/phytotaxa.280.2.2
Li H., Mortimer P. E., Karunarathna S. C., Xu J., Hyde K. D. 2014. New species of Phallus from a subtropical forest in Xishuangbanna, China. Phytotaxa 163(2): 91-103. DOI: 10.11646/phytotaxa.163.2.3
Li H., Tian Ya., Menolli N., Ye L., Karunarathna S. C., Perez J., et al. 2021. Reviewing the world's edible mushroom species: A new evidence-based classification system. Compr. Rev. Food Sci. Food Saf. 20: 1982-2014. DOI: 10.1111/15414337.12708
Li T., Deng W.-Q., Song B., ZhangM., WangM., Li T.-H. 2021. Two new species of Phallus (Phallaceae) with a white indusium from China. MycoKeys 85: 109-125. DOI: 10.3897/mycokeys.85.75309
Li T., Li T., Deng W., Song B., Deng C., Yang Z. L. 2020. Phallus dongsun and P. lutescens, two new species of Phallaceae (Basidiomycota) from China. Phytotaxa 443: 19-37. DOI: 10.11646/phytotaxa.443.1.3
Li T. H., SongB., Liu B. 2002. Three taxa of Phallaceae in HMAS, China. Fungal Diversity 11: 123-127.
Malferrari G., Monferini E., DeBlasio P., Diaferia G., Saltini G., Del Vecchio E., Rossi-Bernardi L., Biunno I. 2002. High-quality genomic DNA from human whole blood and mononuclear cells. BioTechniques 33(6): 1228-1230. DOI: 10.2144/02336bm09
Melanda G. C. S., Silva-Filho A. G. S., Lenz A. R., Menolli N. Jr, Lima A. A., Ferreira R. J., Assis N. M., Cabral T. S., Martín M. P., Baseia I. G. 2021. An Overview of 24 Years of Molecular Phylogenetic Studies in Phallales (Basidiomycota) with Notes on Systematics, Geographic Distribution, Lifestyle, and Edibility. Front. Microbiol. 12: 689374. DOI: 10.3389/fmicb.2021.689374
Milovidova L. S., PlatzM. S., Tolstova N. Yu. 1980. Homobasidial fungi of Tomsk near Ob' region. In: Voprosy biolo-gii [Questions of biology]. Tomsk: Tomsk University Press. Pp. 65-68. [In Russian] (Миловидова Л. С., Плац М. Ш., Толстова Н. Ю. Гомобазидиальные грибы Томского Приобья // Вопросы биологии. Томск: Изд-во Томского ун-та, 1980. С. 65-68).
Moreno G., Khalid A. N., Alvarado P. 2009. A new species of Phallus from Pakistan. Mycotaxon 108: 457-462.
Pereira L. T., De Meijer A. A. R., Reck M. A., Hosaka K., Silveira R. M. B. 2017. Phallus aureolatus (Phallaceae, Agaricomycetes), a new species from the Brazilian Atlantic Forest. Phytotaxa 327(3): 223-236.
Perova N. V., Gorbunova I. A. 2001. Makromitsety yuga Zapadnoy Sibiri [Macromycetes of the South of Western Siberia]. Novosibirsk: SO RAN. 157 pp. [In Russian] (Перова Н. В., Горбунова И. А. Макромицеты юга Западной Сибири. Новосибирск: Изд-во СО РАН, 2001. 157 с.).
Petrov A. N., Matosova E. A. 2010. Dictyophora duplicata (Bosc) E. Fischer. In: Red Data Book of the Irkutsk Oblast. Irkutsk. P. 33. [In Russian] (Петров А. Н., Матосова Е. А. Сетконоска сдвоенная - Dictyophora duplicata (Bosc) E. Fischer // Красная книга Иркутской области. Иркутск, 2010. С. 33).
Rare and endangered species of animals and plants of the Tomsk region. 1984. I. P. Laptev (ed.). Tomsk: Tomsk University Press. 136 pp. [In Russian] (Редкие и исчезающие виды животных и растений Томской области. Под ред. И. П. Лаптева. Томск: Изд-во Томского ун-та, 1984. 136 с.).
Rebriev Yu. A. 2008. Dictyophora duplicata (Bosc) E. Fisch. In: Krasnaya kniga Rossiyskoy Federatsii (Rasteniya i griby) [Red Data Book of the Russian Federation (Plants and Fungi)]. Moscow: KMK Scientific Press Ltd. P. 775. [In Russian] (Ребриев Ю. А. Сетконоска сдвоенная - Dictyophora duplicata (Bosc) E. Fisch. // Красная книга Российской Федерации (Растения и грибы). М.: Тов-во науч. изд. КМК, 2008. С. 775).
Rebriev Yu. A. 2017. Phallus impudicus var. pseudoduplicatus O. Andersson. In: Krasnaya kniga Krasnodarskogo kraya. Rasteniya i griby. Ill-e izd. [Red Book of Krasnodar territory. Plants and Fungi, 3-d ed.]. Krasnodar. Pp. 732-733. [In Russian] (Ребриев Ю. А. Веселка обыкновенная ложносдвоенная - Phallus impudicus var. pseudoduplicatus O. Andersson // Красная книга Краснодарского края. Растения и грибы. III-е изд. Краснодар, 2017. С. 732-733).
Rebriev Yu. A., Ageev D. V., Kokaeva L. Yu., Yaroslavtseva O. N. 2020. The first record of Phallus ultraduplicatus (Phallaceae, Basidiomycetes). Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. [Bull. Moscow Soc. Natur. Biol. Ser.] 125, 1: 66-70. [In Russian] (Ребриев Ю. А., Агеев Д. В., Кокаева Л. Ю., Ярославцева О. Н. Первая находка Phallus ultraduplicatus (Phallaceae, Basidiomycetes) в России // Бюл. МОИП. Отд. биол., 2020. Т. 125, вып. 1. С. 66-70).
Rebriev Yu. A., Bogacheva A. V., Bulakh E. M., Bukharova N. V., Erofeeva E. A., Popov E. S., Psurtseva N. V., Sazanova N. A., Shiryaev A. G., Zvyagina E. A. 2022. New species of macromycetes for regions of the Russian Far East. 3. Mikologiya i fitopatologiya [Mycology and phytopathology] 56(4): 254-263. DOI: 10.31857/S0026364822040080
Rebriev Yu. A., Nikolaev I. A. 2022. Phallus impudicus var. pseudoduplicatus O. Andersson. In: Krasnaya kniga Respubliki Severnaya Osetiya - Alaniya: redkiye i nakhodyashchiyesya pod ugrozoy ischeznoveniya vidy gribov, rasteniy i zhivotnykh [Red Data Book of the Republic of North Ossetia - Alania: rare and endangered species of fungi, plants and animals]. Vladikavkaz: Pero i kist. Pp. 40-41. [In Russian] (Ребриев Ю. А., Николаев И. А. Веселка обыкновенная ложносдвоенная - Phallus impudicus var. pseudoduplicatus O. Andersson // Красная книга Республики Северная Осетия - Алания: редкие и находящиеся под угрозой исчезновения виды грибов, растений и животных. Владикавказ: Перо и Кисть, 2022. С. 40-41).
Rtischeva A. I. 1991. Rare species of fungi of the Central Chernozem region and their protection. Mikologiya i fitopatologiya [Mycology and phytopathology] 25(3): 218-219. [In Russian] (Ртищева А. И. Редкие виды грибов Центрального Черноземья и их охрана // Микология и фитопатология, 1991. Т. 25, вып. 3. С. 218-219).
Sanger F., Nicklen S., Coulson A. R. 1977. DNA sequencing with chain-terminating inhibitors. Proc. Natl. Acad. Sci. USA 74(12): 5463-5467. DOI: 10.1073/pnas.74.12.5463
SongB., Li T., Li T., Huang Q., Deng W. 2018. Phallus fuscoechinovolvatus (Phallaceae, Basidiomycota), a new species with a dark spinose volva from southern China. Phytotaxa 334(1): 19-27. DOI: 10.11646/phytotaxa.334.1.3
Sossin P. E. 1973. Opredelitel gasteromitsetov SSSR [Handbook of the gasteromycetes of the U.S.S.R.]. Leningrad: Nauka. 164 pp. [In Russian] (Сосин П. Е. Определитель гастеромицетов СССР. Л.: Наука, 1973. 164 с.).
Tamura K., Stecher G., Peterson D., Filipski A., Kumar S. 2013. MEGA6: Molecular Evolutionary Genetics Analysis Version 6.0. Mol. Biol. Evol. 30(12): 2725-2729. DOI: 10.1093/molbev/mst197
Turland N. J., Wiersema J. H., Barrie F. R., Greuter W., Hawksworth D. L., Herendeen P. S., et al. (eds.). 2018. International Code of Nomenclature for algae, fungi, and plants (Shenzhen Code) adopted by the Nineteenth International Botanical Congress Shenzhen, China, July 2017. Regnum Vegetabile 159. Glashütten: Koeltz Botanical Books. DOI: 10.12705/Code.2018
Ulbrich E. 1932. Dictyophora duplicata (Bosc) E. Fischer, ein für Europa neuer Vertreter der Phallaceae. Ber. Deutsch. Bot. Ges. 50: 359-366.
Vasilkov B. P. 1954. About some interesting and new species of gasteromycetes in the USSR. Trudy Bot. Inst. Akad. Nauk S.S.S.R., Ser. 2, Sporov. Rast. 9: 447-464. [In Russian] (Васильков Б. П. О некоторых интересных и новых видах гастеромицетов в СССР // Труды Ботанического института им. В. Л. Комарова Академии наук СССР, 1954. Серия 2, вып. 9. С. 447-464).
White T. J., Bruns T., Lee S., Taylor J. W. 1990. Amplification and direct sequencing of fungal ribosomal RNA genes for phylogenetics. In: M. A. Innis et al. (eds.). PCR Protocols: A guide to Methods and Applications. USA: Academic Press, San Diego. Pp. 315-322.
