Научная статья на тему 'AGARICOID AND BOLETOID FUNGI OF RUSSIA: THE MODERN COUNTRY-SCALE CHECKLIST OF SCIENTIFIC NAMES BASED ON LITERATURE DATA'

AGARICOID AND BOLETOID FUNGI OF RUSSIA: THE MODERN COUNTRY-SCALE CHECKLIST OF SCIENTIFIC NAMES BASED ON LITERATURE DATA Текст научной статьи по специальности «Науки о Земле и смежные экологические науки»

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AGARICOMYCETES / FUNGA / FUNGAL BIODIVERSITY AND DISTRIBUTION / REVIEW / RUSSIAN FEDERATION

Аннотация научной статьи по наукам о Земле и смежным экологическим наукам, автор научной работы — Bolshakov Sergey, Kalinina Lyudmila, Palomozhnykh Ekaterina, Potapov Kim, Ageyev Dmitry

The authors of this paper summarize the majority of published data on the distribution of agaricoid and boletoid fungi recorded in Russia, covering the period from 1824 through 2020. A comprehensive list of 6867 scientific names based on 954 publications was compiled for the first time for the whole territory of Russia. All records have been checked through Index Fungorum. The work consists of a review section and five appendices. The review section discusses the intensity of field research and accumulation of data on the distribution of agaricoid and boletoid fungi in Russia, both historically and in its current state. The authors discuss the current state of knowledge on the biodiversity of regions of Russia and point out blank spots, thus providing a reference and an “action plan” for the future. Appendix A presents a list of 6142 taxa unambiguously ascribed to 3246 accepted current names. Appendix B contains 727 names that cannot be ascribed to any accepted current names unequivocally, with reasons given (e.g., no current name, wrong authors’ citations, absence from Index Fungorum). Names from both checklists are supplemented with data on the distribution of these taxa within the Russian Federation and references to published records. Appendix C contains a list of accepted current names reported from only one region. Appendix D is an overview of the nearly 200 years of research of agaricoid and boletoid fungi for all regions of Russia. Appendix E is a list of references used for checklists and study history preparation.

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Текст научной работы на тему «AGARICOID AND BOLETOID FUNGI OF RUSSIA: THE MODERN COUNTRY-SCALE CHECKLIST OF SCIENTIFIC NAMES BASED ON LITERATURE DATA»

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FUNGAL SCIENCE

Citation: Bolshakov, S., Kalinina, L., Palomozhnykh, E., Potapov, K., Ageyev, D., Arslanov, S., Filippova, N., Palamarchuk, M., Tomchin, D., and Voronina, E. 2021. Agaricoid and boletoid fungi of Russia: the modern country-scale checklist of scientific names based on literature data. Bio. Comm. 66(4): 316-325. https://doi. org/10.21638/spbu03.2021.404

Author's information: Sergey Bolshakov, MSc, Junior Researcher, orcid.org/0000-0002-6208-7792; Lyudmila Kalinina, PhD, Researcher, orcid.org/0000-0002-7809-8891; Ekaterina Palomozhnykh, Leading Collection Manager, orcid.org/0000-0002-0789-8365; Kim Potapov, Senior Lecturer, orcid.org/0000-0002-5103-1021; Dmitry Ageyev, Senior Technical Support Specialist, orcid.org/0000-0002-0485-1908; Salavat Arslanov, orcid.org/0000-0002-6327-7870; Nina Filippova, PhD, Senior Researcher, orcid.org/0000-0002-9506-0991; Marina Palamarchuk, PhD, Senior Researcher, orcid. org/0000-0001-8538-969X; Dmitry Tomchin, PhD, Researcher, orcid.org/0000-0002-4223-373X; Elena Voronina, PhD, Associate Professor, orcid.org/0000-0002-5282-6723

Manuscript Editor: Tatiana Svetasheva, Lev Tolstoy Tula State Pedagogical University, Tula, Russia

Received: May 18, 2020;

Revised: July 17, 2021;

Accepted: October 18, 2021.

Copyright: © 2021 Bolshakov et al. This is an open-access article distributed under the terms of the License Agreement with Saint Petersburg State University, which permits to the authors unrestricted distribution, and self-archiving free of charge.

Funding: The work of Sergey Bolshakov, Lyudmila Kalinina, and Ekaterina Palomozhnykh was supported by the institutional research project of the Komarov Botanical Institute ("Biodiversity, ecology, structural and functional features of fungi and fungus-like protists", AAAA-A19-119020890079-6) and partly supported by the Russian Foundation for Basic Research, project No. 20-04-00349 A. The work of Nina Filippova was partly supported by the Grant for organisation of a new young researcher laboratory in Yugra State University, as part of the implementation of the National Project "Science and Universities". The work of Marina Palamarchuk was carried out within the framework of the institutional research project of the Institute of Biology of the Komi Science Centre, Ural Branch of the Russian Academy of Sciences (No. AAAA-A19-119011790022-1). The work of Elena Voronina was carried out as part of the Scientific Project of the State Order of the Government of Russian Federation to Lomonosov Moscow State University No. 121032300081-7.

Ethics statement: This paper does not contain any studies involving human participants or animals performed by any of the authors.

Supplementary information:

Supplemental material to the article is available at https://doi.org/10.21638/ spbu03.2021.404. Supplementary files are published as submitted by the authors, and are not copyedited.

Competing interests: The authors have declared that no competing interests exist.

Agaricoid and boletoid fungi of Russia: the modern country-scale checklist of scientific names based on literature data

Sergey Bolshakov1, Lyudmila Kalinina1, Ekaterina Palomozhnykh1, Kim Potapov2, Dmitry Ageyev3, Salavat Arslanov4, Nina Filippova5, Marina Palamarchuk6, Dmitry Tomchin7, and Elena Voronina8

''Laboratory of Systematics and Geography of Fungi, Komarov Botanical Institute, Russian Academy of Sciences, ul. Professora Popova, 2, Saint Petersburg, 197376, Russian Federation

2Department of General Ecology, Institute of Environmental Sciences, Kazan (Volga Region) Federal University, ul. Kremlyovskaya, 18, Kazan, 420008, Russian Federation

3Signatec Ltd., ul. Inzhenernaya, 22, Novosibirsk, 630090, Russian Federation 4Saint Petersburg Mycological Society, Izmailovsky Pr., 22-1, Saint Petersburg, 190005, Russian Federation

5Department of Environmental Dynamics and Global Climate Change, Yugra State University, ul. Chekhova, 16, Khanty-Mansiysk, 628012, Russian Federation

6Department of Flora and Vegetation of the North, Institute of Biology, Komi Science Centre, Ural Branch of the Russian Academy of Sciences, ul. Kommunisticheskaya, 28, Syktyvkar, 167982, Russian Federation

7Control of Complex Systems Laboratory, Institute for Problems in Mechanical Engineering, Russian Academy of Sciences, Bolshoy Pr. V. O., 61, Saint Petersburg, 199178, Russian Federation

8Department of Mycology and Algology, Faculty of Biology,

Lomonosov Moscow State University, Leninskiye Gory, 1/12, Moscow, 119234,

Russian Federation

Address correspondence and requests for materials to Sergey Bolshakov, s.bolshakov.ru@gmail.com

Abstract

The authors of this paper summarize the majority of published data on the distribution of agaricoid and boletoid fungi recorded in Russia, covering the period from 1824 through 2020. A comprehensive list of 6867 scientific names based on 954 publications was compiled for the first time for the whole territory of Russia. All records have been checked through Index Fungorum. The work consists of a review section and five appendices. The review section discusses the intensity of field research and accumulation of data on the distribution of agaricoid and boletoid fungi in Russia, both historically and in its current state. The authors discuss the current state of knowledge on the biodiversity of regions of Russia and point out blank spots, thus providing a reference and an "action plan" for the future. Appendix A presents a list of 6142 taxa unambiguously ascribed to 3246 accepted current names. Appendix B contains 727 names that cannot be ascribed to any accepted current names unequivocally, with reasons given (e.g., no current name, wrong authors' citations, absence from Index Fungorum). Names from both checklists are supplemented with data on the distribution of these taxa within the Russian Federation and references to published records. Appendix C contains a list of accepted current names reported from only one region. Appendix D is an overview of the nearly 200 years of research of agaricoid and boletoid fungi for all regions of Russia. Appendix E is a list of references used for checklists and study history preparation. Keywords: Agaricomycetes, funga, fungal biodiversity and distribution, review, Russian Federation

Introduction

This study aims to summarize published data on agaricoid and boletoid fungi (Basidiomycota, Agaricomycetes) recorded from the modern territory of Russia (including Crimea). To our knowledge, there is no up-to-date comprehensive

summary of all, or at least the majority of the published data on this group of fungi, which is highly diverse and important, both ecologically and economically.

By "agaricoid and boletoid fungi", we mean basidi-omycete macrofungi with fleshy pileate-stipitate basidi-omata and gilled or tubular hymenophore ("agarics and boleti") (Moser, 1978; Singer, 1986; Knudsen and Vester-holt, 2008, 2012; Halbwachs, Simmel and Bassler, 2016; Sánchez-García et al., 2020). Phylogenetically related cy-phelloid, secotioid or gasteroid species as well as groups traditionally treated as "aphyllophoroid" (clavarioid, corticioid, polyporoid, etc.) were omitted. According to modern taxonomical concepts, taxa of this group belong to the Agaricales, Boletales, Gloeophyllales, Hymeno-chaetales, Polyporales, and Russulales (all orders pro parte) (He et al., 2019).

At the moment, literature containing data on the diversity and distribution of "agarics and boleti" in Russia is burdened with too many redundant names for the same organisms. We took on the task of compiling the essential data from numerous, scattered and sometimes hardly available scientific papers focused mostly on agaricoid and boletoid fungi diversity, taxonomy, and ecology (see "Data source" section) and arranging them into a literature-based all-Russian checklist of scientific names of agaricoid and boletoid fungi. The information included makes it possible to: 1) estimate the general number of agaricoid and boletoid names registered from the territory for every Russian administrative region and the Federation as a whole; 2) estimate spatial study coverage and subsequently reveal blank spots (unstudied or poorly studied regions); 3) provide researchers in the fields of mycogeography and fungal taxonomy with information on species distribution through the checklist as well as the full references list.

The presented study was planned as a "panoramic view" of the known agaricoid and boletoid fungi of Russia. We did not aim to assess the true diversity of this group, and thus limited our effort to revealing gaps in the existing knowledge, which would be of use in planning further inventorying studies of this group of fungi.

Data Gathering & Processing

Data sources. The data for the assessment were extracted from scientific publications and PhD theses containing information on the distribution of agaricoid and boletoid fungi in Russia. To choose relevant and appropriate publications, we took the following steps: 1) examination of archives of the Russian mycological journal "Mycology and Phytopathology (Mikologiya i fitopatologiya)" and "Novitates Systematicae Plantarum non Vascularium (Novosti sistematiki nizshikh rastenii)"; 2) analysis of Russian experts' profiles in the Russian Science Citation Index bibliography database as well as bibliographies of

classic mycologists (Weinmann, Thumen, Karsten, Singer, etc.); 3) analysis of reference lists in the publications selected on the previous two steps. Also, references were obtained from historical overviews of regional botanical and mycological research as well as from search queries via Google Scholar, Scopus, and Web of Science.

The publications used to compile the checklist were obtained from the library of Komarov Botanical Institute of the Russian Academy of Sciences as well as from online and private libraries.

We collected and scrutinized 954 scientific works dating from 1824 to 2020 (journal articles, monographs, book sections, conference proceedings, and PhD theses) of the following types: 1) inventorying studies with species lists for specific territories; 2) taxonomic revisions with data on the examined specimens and their localities; 3) ecological research with information on area of study locality and fungal species arranged in tables or listed in the text as a background for the main research objects. Unpublished data presented in nature reserve annals and research reports were not included. We did not include data from field guides intended for the broad public.

While there remains a possibility of some small gaps in the coverage of this publication, the multifaceted data mining approach has ensured that the vast majority of literature sources on the Russian agaricoid and bole-toid fungi has been incorporated in the checklist.

Data processing. The primary data for the agaricoid and boletoid scientific names checklist was recorded to a flat database in Google Sheets. It contains three main tables — data on species distribution, references and nomenclator, interlinked by data validation, functions, and queries by Google Visualization API Query Language (Bolshakov et al., 2017). Each database line contains basic information on the taxon name: its scientific name with authors' citation, first-level administrative region of Russia and the reference from which the record was obtained. To avoid inaccuracy of the database content, data validation was applied to the species included. Various misprints and errors in the authors' citations of scientific names were corrected when entered in the database in accordance with the correct name in Index Fungorum (IF).

IF was used for classification of scientific names applied to taxa of specific and infraspecific ranks (current names and their homotypic and heterotypic synonyms). The queries to IF were performed on 18.05.2021 through the API using package "taxize" 0.9.99 for R 4.0.5 (Chamberlain and Szocs, 2013; Chamberlain et al., 2020). Nomenclature for the Bolbitiaceae family follows E. F. Maly-sheva's monograph (2018).

Geographic scope. For the assessment, only those records were selected that could be unambiguously ascribed to any of 85 first-level administrative regions of Russia, including Crimea (as they are defined by the Constitution (Chapter 65 and the corresponding Federal

Law from March 21, 2014)). Records from the territories that were part of Russian territory in the past, but not at present, were not considered.

In order to avoid the probable misinterpretation of Russian territory administrative division, the geography of records is presented as transliteration of correspondent territorial subjects' ranks instead of its formal English translation (e.g., Krai instead of Territory, Oblast instead of Region, etc.).

As a result of state reforms in 1990-1993, a number of Russian administrative regions changed their ranks or borders. Thus, the Altai Republic was established within the Altai Krai, and the Republic of Khakassia within the Krasnoyarsk Krai; later, the two Republics became independent federal subjects. The cities of Moscow and St. Petersburg became independent federal subjects within their regions (Moscow Oblast and Leningrad Oblast, respectively); the Khanty-Mansi Autonomous Okrug and Yamalo-Nenets Autonomous Okrug (formerly parts of the Tyumen Oblast) obtained the rank of federal subjects, etc. The corresponding corrections were applied to records that had exact localities provided, making it possible to apply the current territorial nomenclature. If, for instance, the original record was made in the then-Altai Krai, with the precise locality in the Altaiskiy Nature Reserve (now located in the Altai Republic), in the database, it was ascribed to the Altai Republic. In cases when it was impossible to define the exact record localities, as in the case of the Leningrad Oblast checklist (Popov et al., 2007) based on data from Kovalenko and Morozova (1999) and containing records both for Leningrad Oblast and Saint Petersburg (authors' pers. comm.), the broader region (Leningrad Oblast) was specified. The Voronezhskiy Biosphere Reserve lies in the territories of both Lipetsk and Voronezh Oblasts. Records without exact localities were mentioned for both regions.

Bearing in mind the vastness of Russia, we divided its territory into three longitudinal sectors for ease of analysis — Eastern Europe (including the Northern Caucasus and Urals), Siberia, and the Far East. To avoid dividing the Ural Mountains, the boundary between Europe and Siberia and a region of high biodiversity, we are treating it as delimited not by administrative boundaries, but rather by the eastern foothills of the Ural Mountains (200 m. a.s.l.).

Study History

The research history of agarics and boleti in Russia can be tentatively divided into four periods (preliminary, early, classical, and modern). During the preliminary period, research was focused mainly on economically important fungi. In publications from the early period, scattered information on fungi can be found predominantly in reports on academic expeditions organized by the Academy of Sciences, which was established in 1724 (Vassilkov, 1953).

The classical period began with Johann A. Weinmann, the first Russian botanist to contribute significantly to myco-logical studies in the country. He summarized all existing data on macromycetes of the Russian Empire (Weinmann, 1836). Later, only scattered data on agaricoid and boletoid fungi appeared in works of both Russian (Arthur A. Jac-zewski, Fedor V. Bucholtz, etc.) and foreign (Felix von Thumen, Pier A. Saccardo, Petter A. Karsten, etc.) mycologists. The modern period began in the 1920s, when pileate fungi became an independent object of study (Vassilkov, 1953). Lidia A. Lebedeva, Rolf Singer, Ljubov N. Vassileva and Boris P. Vassilkov were the first experts to focus on the study of agaricoid fungi in the USSR. Critically important research on the role of fungi in ecosystems was performed by Vladimir Ya. Chastukhin and colleagues (Chastukhin and Nikolaevskaya, 1969). The period between 1960 and 2000 was marked by the emergence of a number of regional centers of mycological research; beginning from 1990, the number of publications on agaricoid fungi increased dramatically. For detailed information on the published data (papers, researchers, regions etc.), see Appendix D.

Until the 1960s, no more than three publications a year contained any kind of contributions to the knowledge on agaricoid fungi, with rare exceptions; in the course of the following 40 years, until the 2000s, the number of such works published annually increased to 5-8, and beginning from the 2000s, there were usually at least 20 papers published every year (Fig. 1A). The observed curve (Fig. 1C) of the number of publications not only proves the growing interest in various aspects of the biology and taxonomy of agaricoid fungi, but also reflects the explosive growth of publications, which can be considered a general trend in science since the 2000s (Fortunato et al., 2018).

By 1850, the number of published accepted names reached 414 (Fig. 1C). Until the 1970s, data on agaricoid fungi diversity were accumulating very slowly: in the course of 100 years, the number of annually published accepted current names rarely exceeded 100 (Fig. 1B). By 1900, only 541 accepted current names had been published; by 1950, this number increased to 1054. In 1920, Arthur Thesleff (1920) published 445 names for the territory of the modern Leningrad Oblast; in 1939, Vassiljeva (1939) mentioned 318 names for the Caucasus Nature Reserve (territory of modern Krasnodar Krai, Karachay-Cherkess Republic and Adygea); in 1949, Maria V. Freindling reported 228 names from the Kivach Nature Reserve (Karelia). Between 19702000, the numbers of annually published accepted current names of agaricoid fungi varied between 100-900. The majority of large lists during this period were published by Matilda I. Beglyanova (1972) for Krasnoyarsk Krai and Khakassia — 677; Ljubov N. Vassiljeva (1973) for Primor-sky Krai — 737; and Alexander E. Kovalenko and Olga V. Morozova (1999) for Leningrad Oblast and Saint Petersburg — 872 accepted current names. It is noteworthy that beginning from 1962, data were published every year. By

Fig. 1. Timelines of the agaricoid and boletoid fungi diversity research in Russia. A — Annually issued publications. B — Annually reported accepted current names. C — Accumulated numbers of publications (blue line) and detected accepted current names (red line). D — Numbers of studied regions, per year (blue columns) and accumulated (red line).

2000, 2402 accepted current names were published. Since the 2000s, the number of annually published accepted current names increased to 500-1400.

In the 19th century, the number of studied regions increased insignificantly; by 1900, data on agaricoid and boletoid fungi existed only for 15 regions (Fig. 1D). By 1950, the number increased to 58, and by 2000 — to 77. It should be noted that here, we do not assess the intensity of agaricoid fungi research, but simply state the fact of even a single accepted name mentioned for a particular region.

Geographic Distribution

We compiled a list of 6867 names published in 954 scientific papers for 84 of 85 currently existing administrative regions. No data on the Republic of Ingushetia were found. Every name was checked for its current name in IF.

After checking the list against the IF database, all names found in published sources were divided into four groups: 1) 6142 names ascribed unambiguously to accepted current names (as given in IF); 2) 468 names

Table 1. Density of agaricoid and boletoid fungi research in Russia administrative regions

Administrative regions Code Accepted current names % total accepted names "Unique" names Number of publications

Adygea AD 441 13.58 20 26

Altai Krai ALT 473 14.57 4 17

Altai Republic AL 798 24.58 60 69

Amur Oblast AMU 466 14.35 3 39

Arkhangelsk Oblast ARK 137 4.22 2 20

Astrakhan Oblast AST 37 1.14 2 5

Bashkortostan BA 70 2.16 2 11

Belgorod Oblast BEL 431 13.27 6 22

Bryansk Oblast BRY 206 6.34 0 8

Buryatia BU 399 12.29 10 40

Chechen Republic CE 97 2.99 0 6

Chelyabinsk Oblast CHE 241 7.42 4 12

Chukotka Autonomous Okrug CHU 271 8.35 4 25

Chuvash Republic CU 49 1.51 0 5

Crimea KM 655 20.17 20 33

Dagestan DA 7 0.22 0 5

Ingushetia IN 0 0 0 0

Irkutsk Oblast IRK 787 24.24 11 60

Ivanovo Oblast IVA 21 0.65 0 6

Jewish Autonomous Oblast YEV 452 13.92 3 15

Kabardino-Balkarian Republic KB 284 8.75 6 12

Kaliningrad Oblast KGD 670 20.63 13 12

Kalmykia KL 3 0.09 0 2

Kaluga Oblast KLU 302 9.3 0 10

Kamchatka Krai KAM 201 6.19 4 26

Karachay-Cherkess Republic KC 372 11.46 8 23

Karelia KR 781 24.05 22 51

Kemerovo Oblast KEM 45 1.39 0 7

Khabarovsk Krai KHA 683 21.03 15 43

Khakassia KK 437 13.46 13 12

Khanty-Mansi Autonomous Okrug KHM 671 20.67 36 27

Kirov Oblast KIR 329 10.13 0 16

Komi Republic KO 720 22.17 22 36

Kostroma Oblast KOS 67 2.06 0 3

Krasnodar Krai KDA 609 18.76 32 48

Krasnoyarsk Krai KYA 1158 35.66 47 99

Kurgan Oblast KGN 92 2.83 0 6

Kursk Oblast KRS 175 5.39 0 15

Leningrad Oblast LEN 1193 36.74 35 88

Lipetsk Oblast LIP 518 15.95 4 15

Magadan Oblast MAG 487 15 8 29

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Mari El Republic ME 72 2.22 0 9

Administrative regions Code Accepted current names % total accepted names "Unique" names Number of publications

Mordovia MO 283 8.72 4 35

Moscow MOW 240 7.39 8 19

Moscow Oblast MOS 836 25.75 21 46

Murmansk Oblast MUR 451 13.89 12 52

Nenets Autonomous Okrug NEN 16 0.49 1 4

Nizhny Novgorod Oblast NIZ 28 0.86 0 6

North Ossetia SE 135 4.16 1 4

Novgorod Oblast NGR 492 15.15 9 35

Novosibirsk Oblast NVS 649 19.99 10 30

Omsk Oblast OMS 16 0.49 0 6

Orenburg Oblast ORE 322 9.92 8 15

Oryol Oblast ORL 72 2.22 1 6

Penza Oblast PNZ 744 22.91 25 36

Perm Krai PER 848 26.12 17 29

Primorsky Krai PRI 1313 40.44 166 92

Pskov Oblast PSK 590 18.17 7 21

Rostov Oblast ROS 419 12.9 12 18

Ryazan Oblast RYA 530 16.32 10 11

Saint Petersburg SPE 833 25.65 35 54

Sakha Republic SA 180 5.54 17 45

Sakhalin Oblast SAK 331 10.19 12 33

Samara Oblast SAM 496 15.28 16 35

Saratov Oblast SAR 99 3.05 1 11

Sevastopol SEV 210 6.47 3 5

Smolensk Oblast SMO 173 5.33 1 8

Stavropol Krai STA 45 1.39 3 20

Sverdlovsk Oblast SVE 762 23.47 12 27

Tambov Oblast TAM 15 0.46 0 2

Tatarstan TA 564 17.37 12 36

Tomsk Oblast TOM 783 24.11 24 38

Tula Oblast TUL 686 21.13 10 25

Tver Oblast TVE 429 13.21 5 29

Tyumen Oblast TYU 51 1.57 2 11

Tyva Republic TY 42 1.29 1 3

Udmurt Republic UD 481 14.81 3 6

Ulyanovsk Oblast ULY 349 10.75 1 9

Vladimir Oblast VLA 42 1.29 0 8

Volgograd Oblast VGG 172 5.3 16 19

Vologda Oblast VLG 461 14.2 6 26

Voronezh Oblast VOR 322 9.92 1 18

Yamalo-Nenets Autonomous Okrug YAN 317 9.76 7 32

Yaroslavl Oblast YAR 412 12.69 2 12

Zabaykalsky Krai ZAB 53 1.63 0 5

Fig. 2. Choropleth map of accepted current names of agaricoid and boletoid fungi by administrative regions of Russia. Names of administrative regions are abbreviated following the international standard codes ISO 3166-2:RU. The Republic of Crimea and Sevastopol are abbreviated as KM and SEV, respectively. The complete list of region codes is given in Table 1.

Fig. 3. Choropleth map of accepted names of agaricoid and boletoid fungi reported from only one administrative region of Russia ("unique" for this region).

U) K) U)

FUNGAL SCIENCE

that have no current name; 3) 72 correct binomials with incorrect author citations, i.e., such that do not match any records in IF; 4) 187 genus-species or infraspecific combinations absent from IF.

Names unambiguously interpreted according to IF are included in the first list. Infraspecific autonyms without current names also belong here. Such names are given in Appendix A as a list of current names with synonyms.

The second list given in Appendix B enumerates taxa with missing current names as well as correct binomials with wrong author citations. It is impossible to interpret them unambiguously and to bring into correlation with certain current names. Names absent from IF are also included in the second list.

The compiled list of fungal names cannot represent the real structure of the mycobiota, because it contains scientific names of different levels of reliability. Therefore, it can only be subjected to very basic analysis. For the following summary and analyses, we only used information on names and regions from the first list (Appendix A). We consider the authors of the original papers to be in charge of the accuracy of reported species identification.

By 2020, a total of 3246 accepted current names of agaricoid and boletoid fungi were reported for Russia. The general statistics for regions is presented in Table 1. A choropleth map of research density is shown in Fig. 2. The numbers of accepted names recorded for each region differ dramatically. 918 accepted scientific names are known from only one region, 1381 are reported from 2-10 regions, 847 — from 11-50 regions. Only 85 species are reported from more than 50 regions. 15 accepted current names were published as occurring in Russia without the exact region of collection.

The most thoroughly studied regions (more than 1000 accepted names of agaricoid and boletoid fungi reported) are Primorsky Krai — 1313, Leningrad Oblast — 1193, Krasnoyarsk Krai — 1158. The number of well-studied regions (500-1000 names reported) is 21, and there are lists of 100-500 names for 38 regions. The median for all regions is 340.

At the same time, there are three regions with less than 10 accepted names known: Ingushetia — 0, Kalmykia — 3, Dagestan — 7. There are 23 understudied regions in Russia with less than 100 accepted names published. In most cases, there are no published studies of agaricoid and boletoid fungi in these territories, and data for the checklist were obtained from Soviet floras and regional Red lists.

The distribution of 919 currently accepted names known only from a single region also substantially differs from region to region. The following territories are the richest in such "unique" taxa: Primorsky Krai — 166, Altai Republic — 60, Krasnoyarsk Krai — 47, Khanty-Mansi Autonomous Okrug — 36, Leningrad Oblast — 35, Saint Petersburg — 35, Krasnodar Krai — 32.

Eastern Europe

Siberia Far East

Fig. 4. Comparison of lists of accepted current names for longitudinal sectors.

At least 30 such names are recorded in 59 regions, while in 19 regions, there are no unique finds (Fig. 3, Table 1). The full lists of such names for each region are given in Appendix C arranged by longitudinal sectors.

The lists of accepted current names have been compared by longitudinal sectors. In the Eastern European part of Russia 2629 names are recorded, in Siberia — 1959, in the Russian Far East — 1647. A total of 1105 accepted current names are shared by the three sectors. A total of 790 names are unique for the Eastern European Sector, 269 for Siberia and 277 for the Far East (Fig. 4).

The distribution of "unique" accepted names between sectors is as follows: 468 in Eastern Europe, 235 in Siberia and 215 in the Far East (Appendix C).

Concluding remarks

We thoroughly checked all available libraries to find as many publications dealing with agaricoid fungi of Russia as possible. We found 954 works by hundreds of authors in five languages dating from 1824 to 2020 in order to compile the most comprehensive bibliography to date and summarize the existing data, bringing it to the "common denominator" by updating the nomenclature (the accumulated bulk of taxonomical changes gravely complicates the use of these data in their original form); at the same time, the checklist also contains all original information, including synonyms, ambiguous names, etc. The resulting checklist clearly demonstrates that despite more than 150 years of agaricoid fungi research in Russia, there are regions that are in desperate need

of inventorying studies, revealing unexpectedly massive blank spots on the "agaricological" map of Russia. There have been no surveys whatsoever in over twenty regions (Fig. 2); while Northern Caucasus, South Siberia and the Far East are in need of more detailed surveying. We hope that our work will inspire scientists to invest more effort into inventorying surveys, which are a cornerstone in biodiversity and biogeography research.

Acknowledgements

The work is dedicated to Prof. Alexander E. Kovalenko who passed away in December 2021. His hard and fruitful labour was instrumental in the advances of Russian mycological research. This project was initiated with his blessing. We are cordially grateful to our colleagues Olga S.Shiryaeva (Institute of Plant and Animal Ecology, Ural Branch of the Russian Academy of Sciences) and Eugene S.Popov (Komarov Botanical Institute of the Russian Academy of Sciences) for their helpful assistance in compiling data and valuable discussion and suggestions, to the Manuscript Editor Tatiana Yu. Sveta-sheva (Lev Tolstoy Tula State Pedagogical University) as well as the reviewers Henning Knudsen (Professor Emeritus of the Natural History Museum of Denmark, University of Copenhagen) and Slavomir Adamcik (Institute of Botany, Plant Science and Biodiversity Center, Slovak Academy of Sciences) for their valuable remarks and suggestions that greatly improved the quality of this publication. Also, we would like to thank Tatiana M. Bulyonkova (A. P. Ershov Institute of Informatics Systems, Siberian Branch of the Russian Academy of Sciences) for improving the quality of the English text, and for her comments on the subject of the study.

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