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- ОБЗОРНЫЕ СТАТЬИ =
ORCHIDS OF RUSSIA: ANNOTATED CHECKLIST AND GEOGRAPHIC DISTRIBUTION
Petr G. Efimov
Komarov Botanical Institute of RAS, Russia e-mail: [email protected]
Received: 07.03.2020. Revised: 11.04.2020. Accepted: 13.04.2020.
A checklist of orchids is presented for Russia. Data from the project «Biodiversity Mapping of Orchidaceae of Russia» are supplemented by the literature data in order to analyse the geographical distribution of species. We consider that the orchid diversity of Russia comprises 135 species and 13 subspecies belonging to 38 genera, including several taxa with status «uncertain for Russia»: Dactylorhiza majalis, D. sambucina, Epipactispurpurata, Cephalanthera caucasica, Anacamptis laxiflora subsp. palustris, and Platanthera komarovii subsp. maximovic-ziana. Krasnodarsky Krai has the highest number of orchid species (56 species), followed by Sakhalin Region (50), Republic of Dagestan (49), Republic of Crimea (45), Primorsky Krai (42), and Khabarovsky Krai (41). Only in Astrakhan Region there are no orchids known. One species (Anacamptis laxiflora, from one locality) is known from the Republic of Kalmykia, three species from Chukotsky Autonomous Okrug, eight species from Nenetsky Autonomous Okrug. Gymnadenia conopsea is the most widely distributed species, being found in 74 regions (90%), followed by Corallorhiza trifida (73 regions), Dactylorhiza incarnata (72), and D. viridis (71). Twenty-five species are known from only one administrative region. Eleven of them have been found in Sakhalin Region, four of them in Republic of Dagestan. Cephalanthera erecta, Epipactis euxina, and E. leptochila subsp. neglecta are the rarest orchids in Russia (excluding the ones with «uncertain for Russia» status), considering the number of localities: each of these species is known from the unique locus. Two endemic species have been registered in Russia: Epipactis euxina and E. krymmontana. Both species have been described recently. So their distribution demands future studies: supposedly, they may be found in the Caucasus region outside Russia. Liparis loeselii subsp. sachalinensis is another endemic for Russia at the subspecies rank. However, this taxon is uncertain and may be a synonym of L. loeselii subsp. loeselii. Neottia krasnojarica and N. ussuriensis, earlier reported as endemics of Russia, are treated here as synonyms of N. nidus-avis and N. acuminata, respectively. Chamorchis alpina and Liparis loeselii subsp. sachalinensis are the taxa with the highest probability of extinction in Russia.
Key words: biodiversity mapping, geographical analysis, nature conservation, Orchidaceae, rare species, regions of Russia, vascular plants
Introduction
Over the last few decades, there has been a substantial increase of new data about the systematics and taxonomy of Orchidaceae, largely based on molecular polymorphism studies. This is especially true for orchids in the temperate climatic zone. Particularly, their systematics at the genus level is currently developed in detail, and much effort is put in the infra-genera classification issues. Knowledge about the orchid species of Russia is integrating step-by-step into the worldwide dataset. The composition of many taxonomically problematic orchid genera, such as Epipactis Zinn, Platanthera Rich., Liparis Rich., and some others, received in-depth studies in Russia. New catalogues of orchids of many administrative regions of Russia were published or their publication is underway, examples are the Crimea (Fateryga, 2019) or the Asian part of Russia (Chepinoga et al., unpub-
lished). However, a new catalogue of orchids of the whole Russian territory has never been compiled up to now. The current contribution aims to fill this gap.
The taxonomical background for the current catalogue comprises numerous up-to-date data on the orchid systematics and taxonomy. In our study, the priority has been given to the research performed for the Russian territory. Recently, the revisions of several genera have been performed in Russia: Cyp-ripedium L. (Averyanov, 2000), Epipactis (Fateryga & Fateryga, 2018), Liparis (Efimov, 2010), Ophrys L. (Fateryga et al., 2018), three genera, Anacamptis Rich., Neotinea Rchb.f., and Orchis L. s.str., earlier included in Orchis s.l. (Kuropatkin & Efimov, 2014), Serapias L. (Averyanova, 2019), Spiranthes Rich. (Averyanov, 1998), and Traunsteinera Rchb. (Kuropatkin, 2013). Extremely important data about the systematics of particular species belonging to the
genera Cephalanthera Rich. (Fateryga et al., 2020), Neottia Guett. (Efimov, 2019), Gymnadenia R.Br. (Kulikov & Filippov, 2011; Efimov, 2013), and Dac-tylorhiza Neck. ex Nevski (e.g. Shipunov et al., 2004; Pedersen, 2006; Shipunov & Efimov, 2015; Efimov et al., 2016) have been taken into account. Genus taxonomy and nomenclature mostly follow the «Genera Orchidacearum» project (Pridgeon et al., 1999, 2001, 2003, 2005, 2009, 2014).
The current catalogue is supplemented by a geographical analysis of distribution of orchid species in Russia in regard to the state administrative subdivision. Detailed data about distribution of particular species in certain regions of Russia were supplied by a database from a new project «Biodiversity Mapping of Orchidaceae of Russia». This project has been launched in 2017 at the Komarov Botanical Institute of the Russian Academy of Sciences (Efimov et al., 2018) and aims to analyse the orchid species dynamics and developing of a detailed «Atlas of Orchids of Russia», presenting the historical and current distribution of every orchid species as known to date.
Data collection for the project «Biodiversity Mapping of Orchidaceae of Russia» is a two-step process. First of all, the data from the 20-25 main herbarium collections of Russia have been pooled and analysed. Then the regional correspondents from particular administrative units of Russia will contribute the detailed data on a mutually beneficial basis. Currently, the first stage of the project is about to be accomplished. Orchids from the main herbarium collections of Russia have been examined either personally or after digital images, the label information has been put into the database, the data have been attributed in regard to present-day administrative units, and the geotagging is currently underway. At this stage, a special database, which includes entries of orchid localities of Russia, is developed rather in detail. It allows presenting first generalised data about the orchids of Russia in the form of a checklist and to provide the first overview on the geographic distribution of orchids of Russia in general.
Material and Methods
The main data source for this study is the ongoing database, which is a part of the project «Biodiversity Mapping of Orchidaceae of Russia». The database currently holds 73 338 entries, mainly herbarium data. The proportions of literature data and observations are still minimal (11% and 3%, respectively). The following herbarium collections were studied: LE, MW, MHA, LECB, VLA, TK, ALTB, NS, NSK, KFTA, SYKO, PKM, YALT,
LENUD, KPABG (partly), IRKU, VBGI, SVER, MOSP, UFA, IRK, SARAT, PERM, SAKH, NNSU, CSAU, UDU (partly), H (partly), KW (partly), KBHG, KUZ, SIMF, UUH, KBAI, CSR, PVB, and KHA. Small collections of less than 10 000 herbarium specimens in total are not mentioned here. Data from the two most important crowdsourcing online projects, www.inaturalist.org («iNat») and www.plantarium.ru («Plantarium»), were used as well. All images from both sources were checked and, if needed, re-determined according to the accepted methods. It should be noted that the determinations of particular entries from «iNat» and «Plantarium» accepted in the present study may have been changed by later redeterminations. However, the determinations accepted here could always be traced by the logs in «iNat» and «Plan-tarium». They have been published under the name of the author of this article.
The database uses much of original material, such as originally examined herbarium specimens, as well as crowdsourcing online datasets, original literature, personal communications, etc. The use of «sec-ondary» data, such as published maps, generalised lists of localities, lists of herbarium specimens, etc. was avoided whenever possible. The «secondary» data tend to accumulate progressively incorrect data, such as incorrect determinations and inaccurately geotagged material.
One of the important issues for using this database for correct geographic analysis lies in the incomplete data delivery from numerous small regional herbaria and other local sources. So the extraction of full lists of orchid taxa for particular regions may be unavailable from the database. Therefore, information from a local Red Data Book list or recent floristic account has been used for the analysis as well, as a temporary measure. The full list of literature sources is provided in Appendix. In case of discrepancy between our dataset and literature data, priority was given to that providing more complete data. Exceptional priority to original dataset has been given for Dactylorhiza species (except D. aristata (Fisch. ex Lindl.) Soo, D. iberica (M.Bieb. ex Willd.) Soo, and D. viridis (L.) R.M.Bateman, Pridgeon & M.W.Chase), because the literature data are often unreliable due to many incorrect determinations of these species.
The diversity of the orchid species in Russia has been analysed in regard to the 1st-level administrative subdivision principle, meaning that Russia consists of 85 «federal subjects», further referred as «regions». Federal cities (Saint-Petersburg, Moscow, and Sevastopol) have formally been included
into their larger nearest regions, Leningrad Region, Moscow Region, and the Republic of Crimea, respectively. Thus, complete lists of orchid species have been compiled and analysed for each of these 82 regions. The dividing of the Russian territory into «large geographical units» («LGUs»), which are groups of adjacent regions analysed as a whole, is a simplified, but more convenient approach. There are fourteen LGUs: north-western European Russia (Leningrad Region, Novgorod Region, and Pskov Region), «Fennoscandian» Russia (Republic of Karelia, Murmansk Region), northeastern European Russia (Nenetsky Autonomous Okrug, Arkhangelsk Region, Vologda Region, Kostroma Region, and Republic of Komi), central European Russia (Belgorod Region, Bryansk Region, Ivanovo Region, Kaliningrad Region, Kaluga Region, Kursk Region, Lipetsk Region, Moscow Region, Orel Region, Rostov Region, Ryazan Region, Smolensk Region, Tambov Region, Tula Region, Tver Region, Vladimir Region, Voronezh Region, and Yaroslavl Region), Volga region (Astrakhan Region, Nizhniy Novgorod Region, Penza Region, Samara Region, Saratov Region, Ulyanovsk Region, Volgograd Region, Republics of Chuvashia, Kalmykia, Mari El, Mordovia, Tatarstan, and Udmurtia), Crimea (Republic of Crimea), Caucasus (Krasnodarsky Krai, Stavropolsky Krai, Republics of Adygeya, Chechnya, Dagestan, Ingushetia, Kabardino-Balkaria, Karachay-Cherkessia, North Ossetia-Alania), Urals (Permsky Krai, Chelyabinsk Region, Orenburg Region, Sverdlovsk Region, Republic of Bashkortostan), western Siberia (Khanty-Mansi Autonomous Okrug, Yamalo-Nenetsky Autonomous Okrug, Kurgan Region, Omsk Region, and Tyumen Region), southern Siberia (Altaisky Krai, Kemerovo Region, Novosibirsk Region, Tomsk Region, Republic of Altai), Yenissei Siberia (Krasnoyarsky Krai, Republic of Khakassiya, Tyva Republic), eastern Siberia (Trans-Baikalsky Krai, Irkutsk Region, Republic of Buryatia), southern Far East of Russia (Jewish Autonomous Region, Khabarovsky Krai, Primorsky Krai, Amur Region, and Sakhalin Region), northern Far East of Russia (Chukotsky Autonomous Okrug, Kamchatsky Krai, Magadan Region, and Republic of Yakutia).
Results
Checklist of orchids in Russia
The diversity of orchids in Russia comprises up-to-date 135 species (Table 1). Nine species, (Anacamptis morio (L.) R.M. Bateman, Pridgeon & M.W. Chase, Dactylorhiza romana (Sebast.) Soó,
Epipactis leptochila (Godfery) Godfery, Ophrys mammosa Desf., Orchis militaris L., O. purpurea Huds., Platanthera chorisiana (Cham.) Rchb.f., P. densa Freyn, and P. komarovii Schltr.), are divided into two subspecies, two species, (Anacamptis laxiflora (Lam.) R.M. Bateman, Pridgeon & M.W. Chase, Liparis loeselii (L.) Rich.), are divided into three subspecies. Three species, (Anacamptis col-lina subsp. fedtschenkoi (Czerniak.) Kuropatkin & Efimov, A. papilionacea subsp. schirwanica (Woronow) H. Kretzschmar, Eccarius & H. Di-etr., and Serapias orientalis subsp. feldwegiana (H.Baumann & Kunkele) Kreutz), are represented in Russia by non-nominative subspecies only.
The orchid species of Russia belong to 38 genera; 22 genera are monotypic in Russia. The highest species richness is noted in the genus Dactylorhiza (18 species), followed by Epipactis (16), Platanthera (15), Neottia (8), Orchis (8), Cephalanthera (7), Anacamptis (6), Cypripedium (5), and Liparis (5 species), but the number of Liparis species may be seriously underestimated.
Regard must be paid to some taxonomical issues in the checklist, which are still ambiguous and deserve further studies. Thus, large taxonomi-cal controversy is performed by the taxa with large distribution areas, which show a geographically-correlated variability. Most often, this is a problem of taxonomic rank choice, which is usually based on a personal opinion rather than resolved on solid grounds. Absence of such taxa as Dactylorhiza umbrosa (Kar. et Kir.) Nevski and Platanthera ditmari-ana Kom., or presence of such species as Epipactis papillosa Franch. & Sav. and Neottia papilligera Schltr. in our list both represent such cases. Large-scale variation in species-level taxonomy concerns genus Dactylorhiza, which is characterised by numerous morphological forms of basic diploid species (mostly not accepted in the present study) and numerous allopolyploid taxa (accepted here as full species). Future study of orchids will inevitably shed new light on the taxonomy of such taxonomi-cally problematic groups. In some cases, more taxa may be added to the orchid flora of Russia. Currently we may underestimate the diversity in the genera that include cryptic taxa, such as Gymnadenia or Liparis. More taxa may be also discovered in genus Epipactis, if one is going to accept autogamic species that are morphologically weakly delimited, but occur over large geographical areas. In the Caucasus region, Dactylorhiza may be underestimated in regard to allotetraploid taxa other than D. urvilleana (Steud.) H.Baumann & Kunkele (Hedren, 2001).
Table 1. Checklist of orchids in Russia and their distribution along Large Geographic Units
№ Species Subspecies 1 2 3 4 5 6 7 8 9 10 11 12 13 14
1 Anacamptis collina (Banks & Sol. ex Russell) R.M.Bateman, Pridgeon & M.W.Chase subsp. fedtschenkoi (Czerniak.) Kuropatkin & Efimov +
2 A. coriophora (L.) R.M. Bateman, Pridgeon & M.W. Chase + + + +
3 A. laxiflora (Lam.) R.M. Bateman, Pridgeon & M.W. Chase subsp. dielsiana (Soó) H. Kretzschmar, Ec-carius & H. Dietr. + + +
subsp. elegans (HeufF.) Kuropatkin & Efimov +
(?) subsp. palustris (Jacq.) Kuropatkin & Efimov +
4 A. morio (L.) R.M. Bateman, Pridgeon & M.W. Chase subsp. caucasica (K. Koch) H. Kretzschmar, Eccarius & H. Dietr. + +
subsp. morio +
5 A. papilionacea (L.) R.M. Bateman, Pridgeon & M.W. Chase subsp. schirwanica (Woronow) H. Kretzschmar, Eccarius & H. Dietr. +
6 A. pyramidalis (L.) Rich. + +
7 Calypso bulbosa (L.) Oakes + + + + + + + + + + + +
8 (?) Cephalanthera caucasica Kraenzl. +
9 C. damasonium (Mill.) Druce + + +
10 C. epipactoides Fisch. & C.A.Mey. +
11 C. erecta (Thunb.) Blume +
12 C. longibracteata Blume +
13 C. longifolia (L.) Fritsch + + + + +
14 C. rubra (L.) Rich. + + + + + + +
15 Chamorchis alpina (L.) Rich. +
16 Corallorhiza trifida Châtel. + + + + + + + + + + + + + +
17 Cremastra appendiculata (D.Don) Makino +
18 Cypripedium calceolus L. + + + + + + + + + + + + +
19 C. guttatum Sw. + + + + + + + + + +
20 C. macranthos Sw. + + + + + + + + + +
21 C. shanxiense S.C.Chen + +
22 C. yatabeanum Makino + +
23 Dactylorhiza amblyoloba (Nevski) Aver. +
24 D. aristata (Fisch. ex Lindl.) Soó + +
25 D. baltica (Klinge) Nevski + + + + +
26 D. euxina (Nevski) Czerep. +
27 D. fuchsii (Druce) Soó + + + + + + + + + + +
28 D. iberica (M.Bieb. ex Willd.) Soó + +
29 D. incarnata (L.) Soó + + + + + + + + + + + + + +
30 D. maculata (L.) Soó + + + + + + + + +
31 (?) D. majalis (Rchb.) P.F.Hunt & Summerh. +
32 D. ochroleuca (Wüstnei ex Boll) Holub + + + + + + + + + +
33 D. psychrophila (Schltr.) Aver. + + + + +
34 D. romana (Sebast.) Soó subsp. romana +
subsp. georgica (Klinge) Soó ex Renz & Taubenheim +
35 D. salina (Turcz. ex Lindl.) So6 + + + +
36 (?) D. sambucina (L.) So6 +
37 D. sibirica Efimov + + + + + +
38 D. traunsteineri (Saut. ex Rchb.) So6 + + + + + + + +
39 D. urvilleana (Steud.) H.Baumann & Künkele +
40 D. viridis (L.) R.M.Bateman, Pridgeon & M.W.Chase + + + + + + + + + + + + + +
41 Dactylostalix ringens Rchb.f. +
42 Eleorchis japonica (A.Gray) Maek. +
43 Epipactis atrorubens (Hoffm.) Bess. + + + + + + + +
44 E. condensata Boiss. ex D.P. Young +
45 E. distans Arv.-Touv. +
46 E. euxina Fateryga, Popovich & Kreutz +
47 E. helleborine (L.) Crantz + + + + + + + + + + + + + +
48 E. krymmontana Kreutz, Fateryga & Efimov + +
49 E. leptochila (Godfery) Godfery subsp. leptochila + +
subsp. neglecta Kümpel +
№ Species Subspecies 1 2 3 4 5 6 7 8 9 10 11 12 13 14
50 E. microphylla (Ehrh.) Sw. + +
51 E. muelleri Godfery + +
52 E. palustris (Mill.) Crantz + + + + + + + + + + + +
53 E. papillosa Franch. & Sav. + +
54 E. persica (Soö) Hausskn. ex Nannf. + +
55 E. pontica Taubenheim +
56 (?) E. purpurata Sm. +
57 E. tangutica Schltr. +
58 E. thunbergii A. Gray +
59 Ephippianthus sachalinensis Rchb.f. +
60 Epipogium aphyllum (F.W. Schmidt) Sw. + + + + + + + + + + + + + +
61 Galearis cyclochila (Franch. & Sav.) Soö + +
62 Gastrodia elata Blume +
63 Goodyera henryi Rolfe +
64 G. repens (L.) R.Br. + + + + + + + + + + + + + +
65 G. schlechtendaliana Rchb.f. +
66 Gymnadenia conopsea (L.) R.Br. + + + + + + + + + + + + + +
67 G. densiflora (Wahlenb.) A.Dietr. +
68 G. odoratissima (L.) Rich. + +
69 Habenaria linearifolia Maxim. + +
70 H. yezoensis H.Hara +
71 Hammarbyapaludosa (L.) O. Kuntze + + + + + + + + + + +
72 Herminium monorchis (L.) R.Br. + + + + + + + + + + + +
73 Himantoglossum caprinum (M.Bieb.) Spreng. + +
74 H. comperianum (Steven) P.Delforge + +
75 H. formosum (Steven) K.Koch +
76 Limodorum abortivum (L.) Sw. + +
77 Liparis japonica (Miq.) Maxim. +
78 L. krameri Franch. & Sav. +
79 L. kumokiri F.Maek. + + + +
80 L. loeselii (L.) Rich. subsp. loeselii + + + + + + + +
subsp. orientalis Efimov + + +
subsp. sachalinensis (Nakai) Efimov +
81 L. makinoana Schltr. +
82 Malaxis monophyllos (L.) Sw. + + + + + + + + + + + +
83 Myrmechis japonica (Rchb.f.) Rolfe +
84 Neolindleya camtschatica (Cham.) Nevski + +
85 Neotinea tridentata (Scop.) R.M. Bateman, Pridgeon & M.W. Chase + +
86 N. ustulata (L.) R.M. Bateman, Pridgeon & M.W. Chase + + + + + + + +
87 Neottia acuminata Schltr. + +
88 N. camtschatea (L.) Rchb.f. + + + + +
89 N. convallarioides (Sw.) Rich. +
90 N. cordata (L.) Rich. + + + + + + + + + + + + +
91 N. nidus-avis (L.) Rich. + + + + + + + + + + + +
92 N. nipponica (Makino) Szlach. +
93 N. ovata (L.) Bluff & Fingerh. + + + + + + + + + + + +
94 N. puberula (Maxim.) Szlach. + + +
95 N. papilligera Schltr. +
96 Ophrys apifera Huds. + +
97 O. insectifera L. + + + + + +
98 O. mammosa Desf. subsp. caucasica (Woronow ex Grossh.) Soö +
subsp. mammosa + +
99 O. oestrifera M.Bieb. + +
100 Orchis mascula (L.) L. + + + + +
101 O. militaris L. subsp. militaris + + + + + + + + + +
subsp. stevenii (Rchb.f.) B. Baumann, H. Baumann, Lorenz & Ruedi Peter + +
102 O. pallens L. + +
103 O. provincialis Balb. ex Lam. & DC. + +
104 O. punctulata Steven ex Lindl. + +
Note: Large Geographic Units: 1 - north-western European Russia, 2 - «Fennoscandian» Russia, 3 - northeastern European Russia, 4 -central European Russia, 5 - Volga region, 6 - Crimea, 7 - Caucasus, 8 - Urals, 9 - western Siberia, 10 - southern Siberia, 11 - Yenissei Siberia, 12 - eastern Siberia, 13 - southern Far East of Russia, 14 - northern Far East Russia.
№ Species Subspecies 1 2 3 4 5 6 7 8 9 10 11 12 13 14
105 O. purpurea Huds. subsp. caucasica (Regel) B. Baumann, H. Baumann, Lorenz & Ruedi Peter +
subsp. purpurea +
106 O. simia Lam. + +
107 O. spitzelii Saut. ex W.D.J. Koch +
108 Oreorchispatens (Lindl.) Lindl. + +
109 Pecteilis radiata (Thunb.) Raf. +
110 Platanthera bifolia (L.) Rich. + + + + + + + + + + + + +
111 P. chlorantha (Cust.) Rchb. + + + + +
112 P. chorisiana (Cham.) Rchb.f. subsp. chorisiana +
subsp. elata (Finet) Efimov ined. + + +
113 P. convallariifolia (Fisch. ex Lindl.) Lindl. + +
114 P. densa Freyn + +
115 P. dilatata (Pursh) Lindl. ex L.C.Beck +
116 P. fuscescens (L.) Kraenzl. + + + +
117 P. hologlottis Maxim. + +
118 P. komarovii Schltr. subsp. komarovii +
(?) subsp. maximowicziana (Schltr.) Efimov +
119 P. metabifolia F.Maek. + +
120 P. oligantha Turcz. + + + + +
121 P. ophrydioides F.Schmidt +
122 P. sachalinensis F.Schmidt +
123 P. tipuloides (L.f.) Lindl. + + +
124 P. ussuriensis (Regel) Maxim. +
125 Pogonia japonica Rchb.f. +
126 Ponerorchis chusua (D.Don) Soö + +
127 P. cucullata (L.) X.H.Jin, Schuit. & W.T.Jin + + + + + + + + + +
128 P. kinoshitae (Makino) X.H.Jin, Schuit. & W.T. Jin +
129 Pseudorchis albida (L.) A.Löve & D.Löve + + +
130 Serapias orientalis (Greuter) H.Baumann & Künkele subsp. feldwegiana (H.Baumann & Künke-le) Kreutz +
131 Spiranthes sinensis (Pers.) Ames + + + + + + + +
132 S. spiralis (L.) Chevall. +
133 Steveniella satyrioides (Spreng.) Schltr. + +
134 Traunsteinera globosa (L.) Rchb. + +
135 T. sphaerica (M.Bieb.) Schltr. +
Hybrids are not included in the list, but those considered to be «species of hybrid origin» are included. Therefore, allotetraploid dactylorchids (D. majalis (Rchb.) P.F.Hunt & Summerh. aggregate) are listed as full taxa, whereas Cypripedium x ven-tricosum Sw. is not mentioned, although its localities may be locally important for conservation.
Orchids uncertain in Russia
Special attention should be paid to four species and two subspecies, which are included in the Flora of Russia on preliminary basis, as we do not know whether they are present here (or were historically present). These taxa are presented below.
1. Dactylorhiza majalis. It is an allotetraploid species. The eastern boundary of its distribution
range lies somewhere in between Central and Eastern Europe, i.e. approximately, coincides with the western border of Russia. Literature data about this species in Russia are numerous and confusing, because some other allotetraploid species, especially D. baltica (Klinge) Nevski, were often given this name. There are reports on D. majalis in Kaliningrad Region (Gubareva et al., 1999) and Bryansk Region (Fedotov, 2004). These are the areas of its most probable findings. The information from Kaluga Region, Kirov Region, and the Republic of Mari El (Bardunov & Novikov, 2008) is obviously mistaken. We have not seen any herbarium material of this taxon from Russia.
2. Dactylorhiza sambucina (L.) Soo. The eastern boundary of its distribution is similar
to D. majalis. It is also determined incorrectly quite often. However, D. sambucina is currently threatened with extinction, and old literature data cannot be checked when the herbarium specimens are absent. It is reported from Bryansk Region (Bossek, 1986; Fedotov, 2004). We have not seen any herbarium material of this taxon from Russia.
3. Epipactis pur pur ata Sm. The eastern boundary of its distribution is similar to D. ma-jalis. It was reported from Kaliningrad Region (Gubareva et al., 1999). We have not seen any herbarium material of this taxon from Russia.
4. Cephalanthera caucasica Kraenzl. It is reported from Republic of Dagestan historically and nowadays (Grossheim, 1940; Murtazaliev, 2009). Three herbarium specimens have been found (LE!, DAG!, LENUD!). Therefore, four possible localities are known up-to-date. However, all the reports may refer to the hybrid C. longifolia (L.) Fritsch x C. damasonium (Mill.) Druce, which is similar to C. caucasica. Two of the three collections of C. caucasica were made together with C. longifolia or C. damasonium specimens. The main part of the geographic range of C. caucasica is situated around the south part of the Caspian Sea, including Talysh.
5. Anacamptis laxiflora subsp. palustris. Kalinin [= Tver] Region, Bologoye district, near Berezayka station, 20-22 June 1975, Matsenko (MHA!). A surprising collection from a locality isolated from the main range, the nearest localities are in the Baltic countries. A specimen may be mislabeled. The original determination was Dactylorhiza baltica, re-determined correctly by Pavel Kulikov in 1993.
6. Platanthera komarovii subsp. maximovic-ziana (Schltr.) Efimov. [Sakhalin Region] Kuril Islands, Chirpoi Island, Peschanaya Bay, grassland at the sea terrace, 3 August 2000, Barkalov 20546 - LE 1040765! This taxon belongs to the taxonomically ambiguous P. mandarinorum Rchb.f. aggregate. Its systematics was discussed in detail by Efimov (2016). Earlier, P. komarovii was referred to P. maximoviczii Schltr. or P. cornu-bovis Nevski in many literature sources. In fact, P. komarovii (= P. cornu-bovis) is represented by two subspecies, subsp. komarovii and subsp. maximovicziana, the latter usually inhabits higher altitudes and has been recorded only from Japan (Efimov, 2016). Transitional forms between the two subspecies are rare. At Kuril Islands, only subsp. komarovii is known. The
specimen cited above represents its northernmost locality. Vyacheslav Yu. Barkalov collected a similar specimen on the nearby Brat Chirpoyev Island (personal communication). It is not yet clear whether the plants from Chirpoi and Brat Chirvoyev Islands in fact represent subsp. maximovicziana or subsp. komarovii, which developed a maximovicziana-like morphology in the northern climate. In addition, it is possible that both subspecies may represent ecological forms rather than true taxa.
Discussion
Our estimates of 135 species of the orchid flora in Russia is quite close to earlier values of 126 species (Vakhrameeva et al., 2014), but the true difference obviously exceeds nine species. The list presented by these authors includes many species, which we consider to be synonymous to other species of orchids found in Russia (Dactylorhiza cruenta (O.F. Müll.) Soö, D. russowii (Klinge) Holub D. umbrosa, Liparis sachalinensis Nakai, Neottia ussuriensis (Kom. & Nevski) Soö, Ophrys taurica Nevski, Orchis cassidea Biebl, O. picta Raf.) and a hybrid taxon Cypripedium x ventricosum. On the other hand, our list includes species not accepted by Vakh-rameeva et al. (2014) (Dactylorhiza amblyoloba (Nevski) Aver., D. ochroleuca (Wüstnei ex Boll) Holub, D. psychrophila (Schltr.) Aver., Gym-nadenia densiflora (Wahlenb.) A.Dietr.), discovered in Russia later (Dactylorhiza sibirica Efimov, Epipactis euxina Fateryga, Popovich & Kreutz, E. krymmontana Kreutz, Fateryga & Efimov, E. leptochila, E. muelleri Godfery, E. persica (Soö) Hausskn. ex Nannf., E. pontica Taubenheim, E. tangutica Schltr.), found only in Crimea (Epipactis distans Arv.-Touv.), or omitted by Vakhrameeva et al. (2014) for other reasons (Anacamptis collina (Banks & Sol. ex Russell) R.M.Bateman, Pridgeon & M.W.Chase, A. papilionacea (L.) R.M. Bateman, Pridgeon & M.W. Chase, Cephalanthera caucasica, Epipactis condensata Boiss. ex D. P. Young, E. pur-purata). Many species are given under different binomial names, resulting in only 82 binomials in common for the two lists. If one compares our catalogue with earlier data, the difference would be even larger. For example, only 111— 114 species are listed in the «Flora of USSR» (Nevski, 1935). The exact number is even unknown, because the distribution of taxa is given in this publication according to phytogeographic
regions, which do not correspond in many cases to the modern administrative border of Russia.
Orchid species richness in regard to Large Geographic Units
Orchid species are unevenly distributed over Russia. Arid and Arctic areas are characterised by a minimum number of orchid species, whereas afforested zones, especially humid subtropical regions, by a maximum number. A presence/absence matrix of species distribution over LGUs evidences that the most diverse orchid flora is found in the southern Far East of Russia and in the Caucasus region, with 64 and 63 species, respectively, followed by 45 species in Crimea and 43 species in central European Russia (Table 1). Other LGUs comprise from 28 up to 38 species. The minimal number of species (28) is characteristic for «Fennoscandian» Russia, which is a rather small area (two regions only) situated in the northern part of the country.
Orchid species richness in various administrative regions
Krasnodarsky Krai is characterised by the highest orchid species richness of 56 species (Fig. 1), followed by Sakhalin Region (50 species), Republic of Dagestan (49), Republic of Crimea (45), Primorsky Krai (42), Khabarovsky Krai (41), and Republic of Adygeya (38). The lowest orchid species richness is characteristic for arid areas and for the extreme North. Astrakhan Region is the only region with no orchid species. There is a very old sample of Dactylo-rhiza incarnata (L.) Soo labelled «Astrachan» (MW 296034!). But it was likely collected in the northern part of the former Astrakhan Province (= «Astrakhanskaya Guberniya») outside the modern Astrakhan Region. For a long time, no orchid species were known in the neighbouring Republic of Kalmykia, until 30 Anacamptis laxi-flora plants were discovered in 2011 at Bezdon-noye bog in the Maloderbetovskiy district, near the village Plodovitoye (Baktasheva, 2014).
Three orchid species, Corallorhiza trifida Chatel., Dactylorhiza viridis, and Platanthera oligantha Turcz., are known in Chukotsky Autonomous Okrug, which is situated in the extreme North. Eight orchid species have been registered in another northern region, Nenetsky Autonomous Okrug, followed by Yamalo-Nenetsky Autonomous Okrug (10 species), Magadan Region (10), Rostov Region (12), and Volgograd Region
(14 species). The distribution of the number of administrative regions according to the number of orchid species is surprisingly very symmetrical, with maximum number of regions comprising 25-29 orchid species (Fig. 1).
Widely distributed orchid species in Russia
A diagram showing the Orchidaceae species richness per administrative region in Russia is presented in Fig. 2. The diagram is bimodal, indicating that orchid species characterised by narrow and wide distribution prevail, whereas those of intermediate values are rather few, found in 17-36 regions.
«Widely distributed» species are here arbitrarily considered to be present in more than half of the administrative regions. There are twenty-two «widely distributed» species in Russia (number of regions is shown in brackets): Dactylorhiza macu-lata (L.) Soo (43), Liparis loeselii (45), Poneror-chis cucullata (L.) X.H.Jin, Schuit. & W.T.Jin (47), Cypripedium guttatum (49), Epipogium aphyllum (F.W. Schmidt) Sw. (52), Hammarbya paludosa (L.) O. Kuntze (56), Neottia cordata (L.) Rich. (56), Malaxis monophyllos (L.) Sw. (57), Dactylorhiza fuchsii (Druce) Soo (59), Cypripedium cal-ceolus L. (61), Herminium monorchis (L.) R.Br. (62), Orchis militaris (62), Neottia nidus-avis (L.) Rich. (63), Epipactispalustris (Mill.) Crantz (65), Neottia ovata (L.) Bluff & Fingerh. (67), Epipactis helleborine (L.) Crantz (68), Goodyera repens (L.) R.Br. (69), Platanthera bifolia (L.) Rich. (69), Dactylorhiza viridis (71), D. incarnata (72), Corallorhiza trifida (73), Gymnadenia conopsea (L.) R.Br. (74). All of them are taxa characterised by a wide distribution mostly in the boreal zone, either Holarctic or only in the Old World. Gym-nadenia conopsea is the most widely distributed species, occurring in 90% of the Russian regions. This species is able to survive particularly both in arid and in cold climate. Therefore, it has probably the widest distribution in latitudinal direction. In addition, G. conopsea has a maximum number of entries (4787) in our database (6.5% of the total number of entries), significantly differing from the second most common species, Dactylorhiza fuchsii, with 4136 entries.
Rare orchid species in Russia
Twelve orchid species are found in three administrative regions, 15 species in two regions, and 25 species are registered in only one administrative region of Russia each (Fig. 2).
(M 5-9 10-14 15-19 20-24 25-29 30-34 35-39 40-44 45^9 50> number of species
Fig. 1. Orchid species richness in the regions of Russia.
Epipactis distans
Dactylorhiza sambucina (?)
Chamorchis alpina
Platanthera dilatata Neottia convallarioides
Anacamptis collina A. papilionacea
Cephalanthera caucasica (?) 15 Himantoglossum formosum io
Cephalanthera epipactoides Epipactis euxina
П n
Cephalanthera erecta Cremastra appendiculata Dactylostalix ringens Eleorchis japónica Goodyera henryi G. schlechtendaliana Habenaria yezoensis Myrmechis japónica Platanthera ophiydioides P. sachalinensis Poner or chis kinoshitae
2-4 5-9 10-16 17-25 26-36 37-49 50-64 65-74 number of regions
Liparis krameri
Penteilk rarliata
600 mi
Fig. 2. Occurrence of orchid species in regard to the number of regions. Rare taxa found in one region only are shown on the map.
Only one species within the 25 mentioned above taxa, Epipactis euxina, is possibly a narrow endemic. The distribution areas of the other 24 species continuously extend beyond Russia. The highest number of rare species (11 taxa) is noted in Sakhalin Region. In particular, Cepha-lanthera erecta (Thunb.) Blume, Daclylostalix ringens Rchb.f., Eleorchis japonica (A.Gray) Maek., Goodyera henryi Rolfe, G. schlechtendaliana Rchb.f., Habenaria yezoensis H.Hara, Myrmechis japonica (Rchb.f.) Rolfe, and Poner-orchis kinoshitae (Makino) X.H.Jin, Schuit. & W.T.Jin are all found only on the southern Kuril Islands. The others, i.e. Cremastra appendicu-lata (D.Don) Makino, Platanthera ophrydioi-des F.Schmidt, and P. sachalinensis F.Schmidt, are known both from the Kuril Islands and from Sakhalin Island. All these taxa are also registered in Japan. The Republic of Dagestan is another topical region for orchids, which do not occur anywhere else in Russia. There are four species, all of which extend southwards to the Caucasus. Among these four species, Himan-toglossum formosum (Steven) K.Koch, Ana-camptis collina, and A. papilionacea are known from a few localities, whereas the presence of Cephalanthera caucasica needs confirmation (see above). In five regions, Kamchatsky Krai, Krasnodarsky Krai, Primorsky Krai, Republic of Crimea, and Kaliningrad Region, two orchid species are found in each, which are not registered elsewhere. In Kamchatsky Krai, these are Platanthera dilatata (Pursh) Lindl. ex L.C.Beck and Neottia convallarioides (Sw.) Rich. Both of them are found only on the Commander Islands, representing the American flora with a wide distribution in the boreal zone of the New World. Liparis krameri Franch. & Sav. and Pecteilis radiata (Thunb.) Raf. are unique orchids of Pri-morsky Krai, both known from solitary localities in the southernmost part of the region. However, they are also known from China and the Korean Peninsula. In Russia, Cephalanthera epipactoi-des Fisch. & C.A.Mey. and Epipactis euxina occur only in Krasnodarsky Krai. The distribution of the latter species extends southwards beyond Russia. In Russia, Epipactis distans is currently known only from the Republic of Crimea. Epipactis purpurata s.str. has been reported in the Kaliningrad Region, but its occurrence needs confirmation. The same counts for Dactylorhiza sambucina in the Bryansk Region. If it really occurred there, it should be treated now as region-
ally extinct, similarly to Chamorchis alpina (L.) Rich., collected long ago from a small area only, which now belongs to the Murmansk Region.
What orchid is the rarest in Russia, considering the number of localities? If one takes into account the total number of localities and omits possible extinction rates (see below about regionally extinct species) and «uncertain» species (discussed above), then Cephalanthera erecta, Epipactis euxina, and E. leptochila sub-sp. neglecta Kumpel are the rarest orchid taxa in Russia according to our current knowledge. Of each of these taxa only the single record is known (Table 2). However, it is possible that the distribution of the two Epipactis taxa is underestimated, because they are hardly distinguishable in a herbarium from E. persica or E. helleborine, respectively, and further reports from the Caucasus are greatly expected. As for Cephalanthera erecta, its distribution also may be underestimated due to a superficial similarity to Epipactis papillosa (Vyshin, 1996). But since Cephalanthera erecta is found in Russia at the limit of its distribution area, substantial underestimation is hardly possible here. In the literature, one more species, Habenaria yezoensis, was reported from a single locality in Russia, dating back to the contribution by Tatewaki (1954). In Russian herbaria, no specimens were considered to exist (Vyshin, 1996; Vakhrameeva et al., 2014). But, fortunately, we discovered a nice collection of correctly determined specimens of this species dating back from the year 1975 and stored in VBGI herbarium, providing the second locality for this taxon in Russia.
A small number of localities is not the only factor which may promote the taxon extinction. Some orchid species, known from more than two localities, may be even more threatened if their abundance decreases or if the number of the plants in the local populations is too low. Such species as Anacamptis collina subsp. fedtschen-koi, A. papilionacea subsp. schirwanica, Gym-nadenia odoratissima (L.) Rich., Himantoglos-sum formosum, Orchis spitzelii Saut. ex W.D.J. Koch, Ponerorchis chusua (D.Don) Soo, Eleor-chis japonica or Goodyera schlechtendaliana may face the brink of extinction in a short time. However, a detailed analysis of this issue is out of the scope of the present study.
A species has a single locality in a region in 266 cases, two localities in 162 cases, and there are more than two localities per region in 1753 cases.
Table 2. Localities of the rarest orchids, which are known from 1-2 localities only (taxa «uncertain for Russia» excluded)
Taxon name Number of Data sources
localities
Cephalanthera erecta 1 Kuril Islands, Iturup Isl., near Kurilsk, oak forest, 07.08.1988, Barkalov, Vyshin, Yakubov, Bezdeleva (VLA!)
Epipactis euxina 1 Krasnodar Region, Gelendzhik District, near Dzhankhot, Svyataya Nina Mt, burnt pine forest, 10.06.2017, Fateryga, Fateryga, Popovich (holotype, LE!)
Epipactis leptochila subsp. neglecta 1 Krasnodar Region, left bank of Khosta River, 123 m a.s.l., Fagus - Carpinus forest with Taxus baccata, 01.07.2017, Averyanova - LE!, PHEO!. Photographs from the same locality were published on «Plan-tarium» by A. Fateryga: near Khleborob, 12.06.2017, entry no. 516125 (Fateryga, 2017).
Chamorchis alpina 2 1) Lps, ad peninsulam Karelsgammen districtu Kolaensi, 1829, Fellman (H 1746362); «Karelsgam-men» (Fellman, 1831 - cited after Blinova & Uotila, 2011) - according to Blinova & Uotila (2011), actual place is Murmansk Region, Rybachiy Peninsula, Vajda-Guba. 2) Lps, ad alpis Peisen, districtu Kolaensi (Pechenga, in alpes Petchenga), 09.1829, Fellman (H 1746361) - according to Blinova & Uotila (2011), collected in Murmansk Region, near Pechenga.
Habenaria yezoensis 2 1) «Kunashiri, Tomari» [Golovnino] (Tatewaki, 1954). Reference herbarium possibly deposited at herbarium SAP. 2) Kunashir Isl., near Yuzhno-Kurilsk, bog, 11.08.1975, Verkholat, Krivenko 5309 (VBGI!).
Climatic limits of the orchid distribution in the northern areas and in the arid zone
Corallorhiza trifida is a species characterised by the northernmost localities in Russia. However, its northernmost locality is doubtful (ad fl. Taimyr pr. Petrowkrest [?], 27.07.1843, Middendorf - LE 1030927!). There is no correct locality name on the label of the herbarium specimen. According to the collection date, our colleague Igor Pospelov has found that it should be collected at the mouth of the River Verkhnyaya Taimyra (Kupffer Island or Sablera Cape) at 74.50° N. However, the herbarium specimens seem to be too large for the harsh climate of this location, showing evidence that it may be mislabeled. In this case, another sample of the same species from Taimyr, at 73.25° N refers to the northernmost point of Orchidaceae distribution in Russia: «West Taimyr, terrace of the left bank of the River Pyasina in its middle reach in front of the village Tareya, 27.07.1965, Boch» (LE 1030933!). It is followed by the sample obtained from the River Zakharova Rassokha (MW 156299!). In addition to Corallorhiza trifida, Dac-tylorhiza viridis, and Platanthera oligantha also stand the harsh conditions of tundra zone successfully. At last, some localities of Neottia cordata, Pseudorchis albida (L.) A.Love & D.Love, Dac-tylorhiza psychrophila, Goodyera repens, Gym-nadenia conopsea, and Chamorchis alpina also refer formally to the tundra zone.
In the arid zone, no orchids cope with true arid environment. But they grow here locally in damp places which represent a kind of extrazonal distribution. Anacamptis morio or A. laxi-flora are possibly the most drought-resistant species, since their localities in Krasnodarsky Krai, Stavropolsky Krai, Republic of Crimea, Republic
of Dagestan, Republic of Kalmykia, and Rostov Region protrude the most far into the arid zone compared to other species. Another species of the same genus, A. pyramidalis (L.) Rich., is notable for its isolated locality in the central Crimea, if not resulted of mislabeling. Possibly, Neotinea tridentata (Scop.) R.M. Bateman, Pridgeon & M.W. Chase, Dactylorhiza incarnata, and Ophrys mammosa also stand arid environment more successfully than the other taxa.
Endemism
There are only a few Orchidaceae endemics in Russia. If they do not exist in fact, this is not a surprise in regard to the biogeography. Currently, only two taxa at the rank of species, Epipactis euxina and E. krymmontana, should be considered as endemics. Both were described only recently. Given that Epipactis species of the helleborine-complex are often hardly distinguishable in a herbarium, the true distribution area of E. euxina and E. krymmontana cannot be outlined by herbarium material and, therefore, may be seriously underestimated. It is highly possible that in future, both species will be discovered in the Caucasian countries adjacent to Russia, and the accumulation of online crowd-source data confirmed by photographs may strongly contribute to this.
Liparis loeselii subsp. sachalinensis (Nakai) Efimov may be another possible endemic taxon. In most of the literature sources, it is treated at species level (L. sachalinensis). However, its taxonomy is poorly understood now. So this taxon is either identical or very closely related to L. lo-eselii subsp. loeselii (Efimov, 2010), just representing an isolated locality of L. loeselii outside its main distribution range. All Liparis specimens
from Sakhalin were earlier determined as «L. sa-chalinensis» (Barkalov & Taran, 2004). However, except for the type specimen of L. sachalinensis (TI!) and one other collection, all the other Liparis specimens from Sakhalin belong to the group of L. japonica (Miq.) Maxim. - L. kumokiri F.Maek. -L. makinoana Schltr., which is very different morphologically. Only one, poorly collected specimen in fruit (Nevelsk district, Southern Sakhalin State Nature Reserve, Kuznetsovo, River Kuznetsovka, 07.08.1950, Kravchenko - LE 1034591!) may belong to subsp. sachalinensis, along with its holo-type. The scarce material does not allow judging on the degree of difference between subsp. sachalinensis and subsp. loeselii.
Local endemism may be typical for Dactylorhi-za, if one accepts stabilised allotetraploids at species rank. The recently described D. sibirica may serve as an example, if it has been not found in adjacent Kazakhstan. Therefore, it may be classified only as sub-endemic of Russia. It should be noted that the size of the distribution area of D. sibirica in Russia is incomparably larger (more than 200 loci), whereas only a few localities are currently known in Kazakhstan (vouchers in herbaria TK! and SVER!, data from the website www.plantarium.ru).
Liparis loeselii subsp. orientalis Efimov is another sub-endemic of Russia. It has also been described recently. Similarly to D. sibirica, it is a Siberian flora element with a single locus in Kazakhstan (voucher in herbarium MOSP!). Its final taxonomi-cal position is not established yet. It may be a variant of geographic polymorphism of L. loeselii, or it may represent the consequence of introgression of L. loeselii with Far East species of the genus, with the predominance of the traits of the former.
Epipactis euxina, E. krymmontana, Dacty-lorhiza sibirica, and Liparis loeselii subsp. orientalis have been described recently (in 2018, 2014, 2016, and 2010, respectively) and they deserve further studies, which, in turn, may alter their endemic or subendemic status.
In the literature, two other endemic species are mentioned from time to time: Neottia ussuriensis (Vakhrameeva et al., 2014) and N. krasnojarica E.M. Antipova (Varlygina, 2011; Vakhrameeva et al., 2014). These species are not accepted here for several reasons.
Neottia ussuriensis was described in 1935 as a member of a new genus, Holopogon Kom. & Nevski, which was characterised by such impressive features as semi-actinomorphic flowers and a specific column structure with some traits in pre-
sumable ancestral state (Chen & Tsi, 1987). Ho-lopogon was not accepted by most of the authors. It was assumed that all species of this genus represented the transitional stages to peloric forms of the genus Neottia (Wu et al., 2009). This was later confirmed by molecular methods for Chinese Ho-lopogon smithianus (Schltr.) S.C.Chen (Zhou & Jin, 2018). Here, we support the reduction of Ho-lopogon to Neottia and suggest to bring Neottia us-suriensis into synonymy of N. acuminata. The latter was also proposed earlier by Skvortsov (2004) on the basis of morphological studies. This syn-onymisation may explain the unusual rarity of N. ussuriensis, because it was found only twice after its discovery in 1935: in 1984, at a new place, and in 2003, at the type locality again (Vyshin, 1996; see Skvortsov, 2004 for a detailed description of the species morphology and localities).
Neottia krasnojarica was described in 2003 as a third component of a vicarious pair N. nidus-avis - N. papilligera. It was reported to occur very locally within the gap between the distribution ranges of N. nidus-avis and N. papilligera. However, in fact N. krasnojarica is neither geographically, nor morphologically isolated from N. nidus-avis. In particular, its papillose pubescence in the inflorescence is not a discriminative feature of N. krasnojarica, because it may occur in all three species. Moreover, N. papilligera itself is a dubious species, possibly, better referred as a subspecies of N. nidus-avis.
By summarising, only two endemic species of orchids, Epipactis euxina and E. krymmontana, are currently accepted in Russia. However, new localities outside Russia are highly expected. The subspecies Liparis loeselii subsp. sachalinensis may also be endemic, if not taxonomically identical with subsp. loeselii.
Orchids «regionally extinct» in Russia
A problem of species' dynamics is one of the main objectives of the project «Biodiversity Mapping of Orchidaceae of Russia». A detailed study of their dynamics will be possible after all the geotag-ging data will be put into the database. However, the conclusions about the possible extinct orchids of Russia may be drawn right now. The list of taxa with the highest extinction probability in Russia or that may become extinct in the nearest future (taxa «un-certain for Russia» excluded) is presented below.
1. Chamorchis alpina. In 1829, it was recorded from two localities of the current area of Murmansk Region (Blinova & Uotila, 2011). And it has
never been seen again over there, despite visiting its localities by botanists several times, including a small expedition in 2014 designed by the author especially for a Chamorchis search. In Russia, the localities represented the easternmost point of the species' distribution area. By comparing with the other orchids of Russia, the extinction of Chamor-chis is certainly of the highest probability.
2. Liparis loeselii subsp. sachalinensis. If this taxon is not taxonomically identical with subsp. loeselii (see above), it may be extinct because its habitat has been destroyed after it was collected in 1906; nowadays, the former site is possibly located within the Yuzhno-Sakhalinsk town. The second locality is doubtful (determination unsure). On the other hand, new findings are still possible in other localities of Sakhalin Island.
3. Habenaria yezoensis was recorded in Russia twice. The most recent record is dated back to 1975. Nothing is known about this species since that time. But the presence of this species in some place(s) on Kunashir Island is very possible.
4. Ponerorchis chusua. This species was recorded in Amur Region (most of the localities), Jewish Autonomous Okrug, Khabarovsky Krai, Primorsky Krai, and Trans-Baikalsky Krai (in total, ca. 30 databased localities). But it has possibly undergone the most severe decline if compared with all other orchid species of Russia. The most recent finding dated back to 1987, although there are some data on its survival at least in the Khin-gansky State Nature Reserve in the Red Data Book of Amur Region (Senchik, 2019). In the meantime, there are some doubts whether this species survived in Russia generally (Kozhevnikov, 2008). In Bardunov & Novikov (2008), there is an estimation of the total number of plants («500-1000»), but any factual data are absent. So any estimation like this cannot be taken seriously. Most possibly, it survived up to the present days, but the number of the localities is obviously minimal.
Conclusions
A checklist and some short statistical data about the orchids of Russia represent the first results of the project «Biodiversity Mapping of Orchidaceae of Russia». The taxonomical treatment of orchid species in Russia proposed here may be recommended for various local floristic accounts and checklists, as well as for developing of the lists of protected taxa. To date, there is a substantial inconsistency for certain groups of orchids between their taxonomical treatments and revisions presented in
local Russian publications. Especially out-of-date taxonomic concepts may often be encountered in Red Data Books and Red Lists, and as a result they are confusing nature protection actions.
Acknowledgements
The study was financially supported by the Russian Fund for Basic Research, project no. 20-04-00561 (data collection and analysis in the year 2020). The present study was also carried out within the framework of the institutional research project no. АААА-А19-119031290052-1 of the Komarov Botanical Institute of RAS (initial herbarium data accumulation in the period of 2018-2019). The authors thank many colleagues who provided data from numerous herbarium collections in Russia: Ilnur Bikbayev, Olga Biryukova, Alexander Fateryga, Valentina Fateryga, Margarita Gafurova, Tatyana Ivchenko, Olga Kalmykova, Elena Kazmina, Mikhail Kozhin, Marina Legchenko, Ar-tyom Leostrin, Valeriy Linkevich, Svetlana Litvinskaya, Vasiliy Martynenko, Denis Melnikov, Vera Merker, Albert Muldashev, Evgeniy Philippov, Elena Pushay, Stepan Senator, Alexey Seregin, Svetlana Sheremetova, Alla Verkhozi-na, Natalya Zenkova, Ekaterina Zheleznaya, and to those who helped with interpretation of «complicated» localities from herbarium labels. Special thanks are to Igor Pospelov for providing data about Middendorf's collections, and to Alexander Fateryga for providing unpublished data about the systematics of Epipactis, Cephalanthera, and Serapias. Sincere thanks are also due to two anonimous reviewers, who performed a detailed analysis of the manuscript, and to Daria Martynova for linguistic editing.
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Efimov P.G. 2013. Sibling species of Fragrant Orchids (Gymnadenia: Orchidaceae, Magnoliophyta) in Russia. Russian Journal of Genetics 49(3): 299-309. DOI: 10.1134/S102279541302004X Efimov P.G. 2016. A revision of Platanthera (Orchidaceae; Orchidoideae; Orchideae) in Asia. Phytotaxa 254: 1-233. DOI: 10.11646/phytotaxa.254.1.1 Efimov P.G., Filippov E.G., Krivenko D.A. 2016. Allo-polyploid speciation of Dactylorhiza (Orchidaceae, Orchidoideae) in Siberia (Russia). Phytotaxa 258(2): 101-120. DOI: 10.11646/phytotaxa.258.2.1 Efimov P.G. 2019. Taxonomic notes on some photosyn-thetic species of Neottia (Orchidaceae) from North Temperate Asia. Phytotaxa 411(1): 84-88. DOI: 10.11646/phytotaxa.411.1.8 Efimov P.G., Leostrin A.V, Fateryga A.V, Fateryga VV, Senator S.A., Gafurova M.M., Kozhin M.N., Kalmyko-va O.G., Zheleznaya E.L., Pushay E.S., Filippov E.G. 2018. Mapping orchids of Russia: a starting point. In: A.I. Shirokov (Ed.): Protection and cultivation of orchids: materials of XI International Conference (Nizhniy Novgorod, May 25-28, 2018). Nizhniy Novgorod: Nizhniy Novgorod State University. P. 36-37. [In Russian] Fateryga A. 2017. Image of Epipactis leptochila ssp. ne-glecta Kumpel. In: Plantarium: open on-line atlas and key to plants and lichens of Russia and neighbouring countries, 2007-2020. Available from: https://www. plantarium.ru/page/image/id/516125.html Fateryga A.V 2019. New checklist of orchids (Orchidaceae) in the flora of the Crimea. Ekosistemy 17: 38-43. [In Russian]
Fateryga A.V., Efimov P.G., Fateryga V.V. 2018. Taxonomic notes on the genus Ophrys L. (Orchidaceae) in the Crimea and the North Caucasus. Turczaninowia 21(4): 9-18. DOI: 10.14258/turczaninowia.21.4.2 Fateryga A.V., Fateryga V.V. 2018. The genus Epipactis Zinn (Orchidaceae) in the flora of Russia. Turczaninowia 21(4): 19-34. DOI: 10.14258/turczaninow-ia.21.4.3 [In Russian] Fateryga A.V, Popovich A.V, Fateryga V.V., Kreutz C.A.J. 2020. Cephalanthera epipactoides (Orchidaceae) in Russia. Nature Conservation Research 5(Suppl. 1): 69-76. DOI: 10.24189/ncr.2020.014 Fedotov Yu. P. (Ed.). 2004. Red Data Book of Bryansk Region. Plants. Fungi. Bryansk: Chitay-Gorod. 272 p. [In Russian]
Grossheim A.A. 1940. Flora of Caucasus, vol. 2. 2nd ed. Baku: Azerbaijanian Branch of AS USSR. 284 p. [In Russian] Gubareva I.Yu., Dedkov V.P., Napreenko M.G., Petrova N.G., Sokolov A.A. 1999. Conspectus of vascular
plants of Kaliningrad Region: a manual. Kaliningrad: Kaliningrad University. 107 p. [In Russian] Hedren M. 2001. Systematics of the Dactylorhiza euxi-na/incarnata/maculata polyploid complex (Orchi-daceae) in Turkey: evidence from allozyme data. Plant Systematics and Evolution 229: 23-44. DOI: 10.1007/s006060170016 Kozhevnikov A.E. (Ed.). 2008. Red Data Book of Primor-sky Krai. Plants. Vladivostok: AVK Apelsin. 688 p. [In Russian]
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Vol. 4. Makhachkala: Epokha. 231 p. [In Russian] Nevski S.A. 1935. Orchidaceae Lindl. In: V.L. Komarov (Ed.): Flora USSR. Vol. 4. Leningrad: Editio Aca-demiae Scientiarum USSR. P. 589-730, 750-754. [In Russian]
Pedersen H.^. 2006. Systematics and evolution of the Dactylorhiza romana/sambucina polyploid complex (Orchidaceae). Botanical Journal of the Lin-nean Society 152(4): 405-434. DOI: 10.1111/j.1095-8339.2006.00573.x Pridgeon A.M., Cribb P.J., Chase M.W., Rasmussen F.N. (Eds.). 1999. Genera Orchidacearum. Vol. 1: General Introduction, Apostasioideae and Cypripedioideae. Oxford: Oxford University Press. 197 p. Pridgeon A.M., Cribb P.J., Chase M.W., Rasmussen F.N. (Eds.). 2001. Genera Orchidacearum. Vol. 2: Orchidoideae (part 1). Oxford: Oxford University Press. 416 p.
Pridgeon A.M., Cribb P.J., Chase M.W., Rasmussen F.N. (Eds.). 2003. Genera Orchidacearum. Vol. 3: Orchidoideae (part 2). Oxford: Oxford University Press. 358 p.
Pridgeon A.M., Cribb P.J., Chase M.W., Rasmussen F.N. (Eds.). 2005. Genera Orchidacearum. Vol. 4: Epi-dendroideae (part 1). Oxford: Oxford University Press. 672 p.
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Appendix. Literature sources for Russian regions.
ALTAISKY KRAI: Kamelin R.V., Shmakov A.I. (Eds.). 2006. Red Data Book of Altaisky Krai. Barnaul: IPP Altay. 262 p. [In Russian]
AMUR REGION: Senchik A.V. (Ed.). 2019. Red Data Book of Amur Region: Rare and endangered animals, plants and fungi. Blagoveshchensk: Far East State Agrarian University. 499 p. [In Russian]
ARKHANGELSK REGION: Novoselov A.P. (Ed.). 2008. Red Data Book of the Arkhangelsk Region. Arkhangelsk: Committee of Ecology of Arkhangelsk Region. 351 p. [In Russian]
BELGOROD REGION: Prisnyi A.V. (Ed.). 2005. Red Data Book of Belgorod Region. Belgorod: Ecological Fund of Belgorod Region. 532 p. [In Russian]
BRYANSK REGION: Fedotov Yu.P. (Ed.). 2004. Red Data Book of Bryansk Region. Plants. Fungi. Bryansk: Chi-tay-Gorod. 272 p. [In Russian]
CHELYABINSK REGION: Korytin N.S. (Ed.). 2005. Red Data Book of Chelyabinsk Region. Ekaterinburg: Ural University Press. 362 p. [In Russian]
CHUKOTSKY AUTONOMOUS OKRUG: Chereshnev I.A. (Ed.). 2008. Red Data Book of the Chu-kotsky Autonomous Pkrug. Vol. 2. Magadan: Department of Industrial and Agricultural Politics of Chukotsky Autonomous Okrug. 217 p. [In Russian]
IRKUTSK REGION: Gaykova O.Yu., Popov V.V. (Eds.). 2010. Red Data Book of Irkutsk Region. Irkutsk: Vremya Stranstviy Press. 480 p. [In Russian]
IVANOVO REGION: Borisova E.A., Kurganov A.A., Mishagina D.A. 2014. Patterns of orchid species distribution in the Ivanovo Region. Vestnik of Lobachevsky Nizhnii Novgorod State University 3(3): 26-29. [In Russian]
JEWISH AUTONOMOUS OKRUG: Shlotgauer S.D., Rubtsova T.A. (Eds.). 2006. Red Data Book of the
Vakhrameeva M.G., Varlygina T.I., Tatarenko I.V 2014. Orchids of Russia (biology, ecology and protection). Moscow: KMK Scientific Press Ltd. 437 p. [In Russian] Varlygina T.I. 2011. Preservation of Orchidaceae in Russia. In: I.I. Shamrov (Ed.): Protection and cultivation of orchids: materials of IX International Conference (26-30 September 2011). Moscow: KMK Scientific Press Ltd. P. 76-80. [In Russian] Vyshin I.B. 1996. Orchidaceae. In: S.S. Charkevicz (Ed.): Plantae vasculares orientis extremi sovietici, vol. 8. Saint Petersburg: Nauka. P. 301-339. [In Russian] Wu Z.-Y., Raven P.H., Hong D.-Y. (Eds.). 2009. Flora of China. Vol. 25 (Orchidaceae). Beijing & St. Louis: Science Press & Missouri Botanical Garden Press. 570 p. Zhou T., Jin X.-H. 2018. Molecular systematics and evolution of mycoheterotrophy of tribe Neottieae (Orchidaceae, Epidendroideae). In: X.-H. Jin, Y.-M. Shui, Y.-H. Tan, M. Kang (Eds.): Plant diversity in Southeast Asia. PhytoKeys 94: 39-49. DOI: 10.3897/phy-tokeys.94.21346
Jewish Autonomous Okrug. Novosibirsk: Arta Publishing. 247 p. [In Russian]
KALININGRAD REGION: Gubareva I.Yu., Dedkov V.P., Napreenko M.G., Petrova N.G., Sokolov A.A. 1999. Conspectus of vascular plants of Kaliningrad Region: a manual. Kaliningrad: Kaliningrad University. 107 p. [In Russian]; Dedkov V.P., Grishanov G.V. (Eds.). 2010. Red Data Book of Kaliningrad Region. Kaliningrad: Immanuil Kant Russian State University. 334 p. [In Russian]
KALUGA REGION: Reshetnikova N.M., Krylov A.M., Sidorenkova E.M., Voronkina N.V., Shmytov A.A., Popchenko M.I., Mayorov S.R., Romanova R.A. 2015. Materials for Red Data Book of Kaluga Region: registration of vascular plants in the past 150 years with maps. Kaluga: Vash Dom. 448 p. [In Russian]
KAMCHATSKY KRAI: Chernyagina O.A. (Ed.). 2018. Red Data Book ofKamchatsky Krai. Vol. 2. Petropav-lovsk-Kamchatskiy: Kamchatpress. 388 p. [In Russian]
KEMEROVO REGION: Kupriyanov A.N. (Ed.). 2012. Red Data Book of the Kemerovo Region. Kemerovo: Asia Print. 208 p. [In Russian]
KHABAROVSKY KRAI: Voronov B.A. (Ed.). 2008. Red Data Book of the Khabarovsky Krai. Khabarovsk: Pri-amurskie Vedomosti Publishing House. 632 p. [In Russian] KHANTY-MANSI AUTONOMOUS OKRUG: Vasin A.M., Vasina A.L. (Eds.). 2013. Red Data Book of Khanty-Mansi Autonomous Okrug. Ekaterinburg: Basko. 460 p. [In Russian]
KIROV REGION: Bolshakov V.N. (Ed.). 2001. Red Data Book of Kirov Region. Ekaterinburg: Ural University Press. 215 p. [In Russian]
KOSTROMA REGION: Elenevskiy A.G., Konstanti-nov V.M. (Eds.). 2009. Red Data Book of Kostroma Region. Kostroma. 387 p. [In Russian]
KRASNODARSKY KRAI: Litvinskaya S.A. (Ed.). 2017. Red Data Book of Krasnodarsky Krai. Krasnodar. 850 p. [In Russian]
KRASNOYARSKY KRAI: Stepanov N.V. (Ed.) 2012. Red Data Book of the Krasnoyarsky Krai. Krasnoyarsk: Siberian Federal University. 576 p. [In Russian]
KURGAN REGION: Naumenko N.I. 2008. Flora and vegetation of South Trans-Ural Area. Kurgan: Kurgan State University. 512 p. [In Russian]
KURSK REGION: Zolotukhin N.I. (Ed.) 2001. Red Data Book of Kursk Region. Tula: Centralno-Chernozemny Zapovednik. 165 p. [In Russian]
LENINGRAD REGION: Efimov P.G. 2012. Orchids of North-West European Russia (in the limits of Leningrad, Pskov and Novgorod Regions). 2nd ed. Moscow: KMK Scientific Press Ltd. 220 p. [In Russian]
LIPETSK REGION: Shcherbakov A.V. (Ed.). 2014. Red Data Book of Lipetsk Region. Vol. 1. Lipetsk: Committee for Ecology and Nature Resources of Lipetsk Region. 696 p. [In Russian]
MAGADAN REGION: Khoreva M.G., Dokuchaev N.E. (Eds.). 2019. Red Data Book of Magadan Region. Magadan: Okhotnik. 356 p. [In Russian]
MOSCOW REGION: Varlygina T.I., Zubakin V.A., Sobolev N.A. (Eds.). 2008. Red Data Book of Moscow Region. Moscow: KMK Scientific Press Ltd. 828 p. [In Russian]
MURMANSK REGION: Konstantinova N.A., Ko-ryakin A.S., Makarova O.A., Bianki V.V. (Eds.). 2014. Red Data Book of Murmansk Region. Kemerovo: Asia-Print. 584 p. [In Russian]
NENETSKY AUTONOMOUS OKRUG: Matveeva N.V. (Ed.). 2020. Red Data Book of Nenetsky Autonomous Okrug. Naryan-Mar. 456 p. [Manuscript] [In Russian]
NIZHNIY NOVGOROD REGION: Biryukova O.V., Vorotnikov V.P., Mininzon I.L. 2014. The Orchidaceae family in the flora of the Nizhniy Novgorod Region. Vestnik of Lobachevsky Nizhnii Novgorod State University 3(3): 16-25. [In Russian]
NOVGOROD REGION: Efimov P.G. 2012. Orchids of North-West European Russia (in the limits of Leningrad, Pskov and Novgorod Regions). 2nd ed. Moscow: KMK Scientific Press Ltd. 220 p. [In Russian]
NOVOSIBIRSK REGION: Marchenko Yu.Yu., Yudkin V.A., Shaulo D.N. (Eds.). 2008. Red Data Book of Novosibirsk Region. Novosibirsk: Arta. 528 p. [In Russian]
OMSK REGION: Sidorov V.N., Rusakov G.N. (Eds.). 2005. Red Data Book of Omsk Region. Omsk: Omsk State University. 460 p. [In Russian]
OREL REGION: Ogureeva G.N., Khitsova L.N. (Eds.). 2007. Red Data Book of Orel Region. Orel: A.V. Vo-robyev. 264 p. [In Russian]
ORENBURG REGION: Belov V.S. (Ed.). 2019. Red Data Book of Orenburg Region. Voronezh: Mir. 488 p. [In Russian]
PENZA REGION: Ivanov A.I. (Ed.). 2002. Red Data Book of Penza Region. Vol. 1. Penza: Committee for Nature Resources of Penza Region. 159 p. [In Russian]
PERMSKY KRAI: Ovesnov S.A. 2009. Local Flora. Flora of Perm Krai and its analysis. Perm: Perm State University. 215 p. [In Russian]
PRIMORSKY KRAI: Kozhevnikov A.E. (Ed.). 2008. Red Data Book Primorsky Krai, Plants. Vladivostok: AVK Apelsin. 688 p. [In Russian]
PSKOV REGION: Efimov P.G. 2012. Orchids of North-West European Russia (in the limits of Leningrad, Pskov and Novgorod Regions). 2nd ed. Moscow: KMK Scientific Press Ltd. 220 p. [In Russian]
REPUBLIC OF ADYGEYA: Sirotyuk (Kuvaeva) E.A., Akatova T.V., Lipka O.N. (Eds.). 2012. Red Data Book of the Republic of Adygeya. Part 2. Maykop: Kachestvo. 340 p. [In Russian]
REPUBLIC OF ALTAI: Gerasimovich L.V. 2012. Orchids (Orchidaceae) of the Altai Mountains. Novosibirsk: Geo. 119 p. [In Russian]
REPUBLIC OF BASHKORTOSTAN: Mirkin B.M. (Ed.). 2011. Red Data Book of the Republic of Bashkortostan. Vol. 1. Ufa: Media Print. 384 p. [In Russian]
REPUBLIC OF BURYATYA: Pronin N.M. (Ed.).
2013. Red Data Book of the Republic of Buryatia. Ulan-Ude: Buryat Branch of RAS. 688 p. [In Russian]
REPUBLIC OF CHECHNYA: Taysumov M.A., Tochiev T.Yu., Umarov R.M., Gayrabekova R.Kh., Kushalieva Sh.A., Shoipov R.Kh. (Eds.). 2007. Red Data Book of the Republic of Chechnya. Groznyi: Ecological Committee of the Republic of Chechnya. 158 p. [In Russian]
REPUBLIC OF CHUVASHIA: Gafurova M.M.
2014. Vascular plants of Chuvash Republic. Flora of the Volga River Basin. Vol. 3. Togliatti: Kassandra. 333 p. [In Russian]
REPUBLIC OF CRIMEA: Fateryga A.V., Efimov P.G., Svirin S.A. 2019. Orchids of the Crimean Peninsula. Simferopol: Arial. 224 p. [In Russian]
REPUBLIC OF DAGESTAN: Murtazaliev R.A. 2009. Daghestan Flora Conspectus. Vol. 4. Makhachkala: Epokha. 231 p. [In Russian]
REPUBLIC OF INGUSHETIA: Martazanov A.M. (Ed.). 2007. Red Data Book of the Republic of Ingushetia. Magas: Serdalo Press. 368 p. [In Russian]
REPUBLIC OF KABARDINO-BALKARIYA: Za-likhanov M.Ch. (Ed.). 2018. Red Data Book of the Republic of Kabardino-Balkariya. Nalchik: Pechatnyi Dvor. 494 p. [In Russian]
REPUBLIC OF KALMYKIA: Baktasheva N.M. (Ed.). 2014. Red Data Book of Kalmykia. Vol. 2. Elista: Dzhangar. 199 p. [In Russian]
REPUBLIC OF KARACHAY-CHERKESSIA. Onipchenko V.G. (Ed.). 2013. Red Data Book of the Republic of Karachay-Cherkessia. Cherkessk: Nartizdat. 360 p. [In Russian]
REPUBLIC OF KARELIA: Kravchenko A.V. 2007. Conspectus of Karelian Flora. Petrozavodsk: Karelian Scientific Centre RAS. 403 p. [In Russian]
REPUBLIC OF KHAKASSIA: Ankipovich E.S. (Ed.)
2012. Red Data Book of the Republic of Khakassia. Novosibirsk: Nauka. 288 p. [In Russian]
REPUBLIC OF KOMI: Teteryuk L., Kirillova I. 2011. Rare and protected orchids of the Komi Republic. Berichte Arbeitskreise Heimische Orchideen 28(1): 133-179.
REPUBLIC OF MARIY EL: Bogdanov G.A., Abramov N.V., Urbanavichus G.P., Bogdanova L.G. (Eds.).
2013. Red Data Book of the Republic of Mariy El. Yoshkar-Ola: Mariy El State University. 324 p. [In Russian]
REPUBLIC OF MORDOVIA: Silaeva T.B., Kiijukhin I.V., Chugunov G.G., Ljovin V.K., Majorov S.R., Pismarki-na E.V., Ageeva A.M., Vargot E.V. 2010. Plantae Vasculares Rei Publicae Mordoviensis (Conspectus Florae). Saransk: Publisher of the Mordovia State University. 352 p. [In Russian]
REPUBLIC OF NORTH OSETIYA-ALANIYA: Vagin V.S., Olisaev V.A. (Eds.). 1999. Red Data Book of the Republic of Severnaya Osetiya-Alaniya. Vladikavkaz: Proekt Press. 248 p. [In Russian]
REPUBLIC OF TATARSTAN: Shchepovskih A.I. (Ed.). 2006. Red Data Book of the Republic of Tatarstan. Kasan: Idel-Press. 832 p. [In Russian]
REPUBLIC OF TYVA: Ondar S.O., Shaulo D.N. (Eds.). 2019. Red Data Book of the Republic of Tyva. Kyzyl. 564 p. [In Russian]
REPUBLIC OF UDMURTIA: Baranova O.G. 2006. The peculiarities of the Orchidaceae Juss. family species distribution in the Udmurt Republic and their protection. Vestnik of Udmurt State University. Series Biology 10: 3-10. [In Russian]
REPUBLIC OF YAKUTIA: Danilova N.S. (Ed.). 2017. Red Data Book of the Republic of Yakutia. Vol. 1. Moscow: Reart Press. 412 p. [In Russian]
ROSTOV REGION: Minoranskiy V.A. (Ed.). 2004. Red Data Book of Rostov Region. Vol. 2. Rostov-on-Don: Malysh Publishing. 333 p. [In Russian]
RYAZAN REGION: Ivanchev V.P., Kazakova M.V. (Eds.). 2011. Red Data Book of Ryazan Region. Ryazan: Golos Gubernii. 626 p. [In Russian]
SAKHALIN REGION: Eremin V.M. (Ed.). 2005. Red Data Book of Sakhalin Region. Plants. Sakhalin: Sakhalin-skoye Publishing House. 348 p. [In Russian]
SAMARA REGION: Saksonov S.V., Koneva N.V. 2006. The abstract of Orchidaceae family [of] the Samara Area. Vestnik of Udmurt State University. Series Biology 10: 43-50. [In Russian]
SARATOV REGION: Shlyakhtin G.V. (Ed.). 2006. Red Data Book of Saratov Region. Saratov: Torgovo-Promyshlen-naya Palata Saratovskoy Oblasti Press. 526 p. [In Russian]
SMOLENSK REGION: Kruglov N.D. 1997. Red Data Book of Smolensk Region. Smolensk: Smolensk State Pedagogical Institute. 294 p. [In Russian]
STAVROPOLSKY KRAI: Ivanov A.L. (Ed.). Red Data Book of Stavropolsky Krai. Vol. 1. Stavropol: Andreev
1.V. 374 p. [In Russian]
SVERDLOVSK REGION: Knyazev M.S., Tretyakova A.S., Podgaevskaya E.N., Zolotareva N.V., Kulikov P.V. 2017. An annotated checklist of the flora of Sverdlovsk Region. Part II: Monocotyledonous plants. Phytodiversity of Eastern Europe 11(3): 4-108. [In Russian]
TAMBOV REGION: Sukhorukov A.P. (Ed.). 2010. Manual of vascular plants of Tambov Region. Tula: Grif i K. 349 p. [In Russian]
TOMSK REGION: Adam A.M. (Ed.). 2013. Red Data Book of Tomsk Region. Tomsk: Pechatnaya Manufactura Press. 504 p. [In Russian]
TRANS-BAIKALSKY KRAI: Polyakov O.A., Popo-va O.A., Afonina A.M., Gileva M.V., Makryi T. V., Pershina N.A., Petrov A.N., Tkachuk T.E., Kharchenko N.A. (Eds.). 2017. Red Data Book of Trans-Baikalsky Krai. Plants. Novosibirsk: Dom Mira. 384 p. [In Russian]
TULA REGION: Filatov V.P. (Ed.). 2010. Red Data Book of Tula Region. Tula: Grif i K. 393 p. [In Russian]
TVER REGION: Pushay E.S., Dementyeva S.M. 2008. Biology, ecology and distribution of species of the family Orchidaceae Juss. in Tver Region. Tver: Tver State University. 206 p. [In Russian]
TYUMEN REGION: Petrova E.A. (Ed.) 2004. Red Data Book of Tyumen Region. Ekaterinburg: Ural University Press. 496 p. [In Russian]
ULYANOVSK REGION: Rakov N.S., Saksonov S.V., Senator S.A., Vasjukov V.M. 2014. Vascular plants of Ulyanovsk Region. Flora of the Volga River Basin. Vol. 2. Togliatti: Kassandra. 295 p. [In Russian]
VLADIMIR REGION: Seregin A.P., Borovichev E.A., Glazunova K.P., Kokoshnikova Y.S., Sennikov A.N. 2012. Flora of Vladimir Oblast, Russia: Checklist and atlas. Tula: Grif i K. 620 p. [In Russian]
VOLGOGRAD REGION: Baranova O.G., Sagalaev V.A. (Eds.). 2017. Red Data Book of Volgograd Region. Vol. 2. Voronezh: Izdat-Print. 268 p. [In Russian]
VOLOGDA REGION: Konechnaya G.Yu., Suslova T.A. (Eds.). 2004. Red Data Book of Vologda Region. Vol.
2. Vologda: Vologda State University and Rus' Press. 360 p. [In Russian]
VORONEZH REGION: Agafonov V.A., Starodubt-seva E.A., Negrobov V.V., Barabash G.I., Bedenko A.B., Kazmina E.S., Kirik A.I., Kobzeva E.V., Chernyshova T.N. 2019. The cadastre of vascular plants protected in Voronezh Region. Voronezh: Tsifrovaya poligraphiya. 440 p. [In Russian]
YAMALO-NENETSKY AUTONOMOUS OKRUG: Ektova S.N., Zamyatin D.O. (Eds.). 2010. Red Data Book of Yamalo-Nenetsky Autonomous Okrug. Ekaterinburg: Basko. 308 p. [In Russian]
YAROSLAVL REGION: Voronin L.V. (Ed.). 2004. Red Data Book of Yaroslavl Region. Yaroslavl: Alexandr Rutman. 384 p. [In Russian]
ОРХИДНЫЕ РОССИИ: ЧЕК-ЛИСТ И КРАТКИЙ АНАЛИЗ ГЕОГРАФИЧЕСКОГО РАСПРОСТРАНЕНИЯ
П. Г. Ефимов
Ботанический институт им. В.Л. Комарова РАН, Россия e-mail: [email protected]
В статье представлен чек-лист орхидных России. При помощи данных проекта «Картирование орхидных России», а также сведений из литературы осуществлен географический анализ распространения орхидных России. Общее разнообразие орхидных России оценено в 38 родов, 135 видов и 13 подвидов, включая виды, предположительно встречающиеся в России - Dactylorhiza majalis, D. sambucina, Epipactis purpurata, Cephalanthera caucasica, Anacamptis laxiflora subsp. palustris, Platan-thera komarovii subsp. maximovicziana. Наиболее богатым административным регионом страны по числу видов орхидных оказался Краснодарский край, в котором встречается 56 видов, за ним следуют Сахалинская область (50), Республика Дагестан (49), Республика Крым (45), Приморский край (42), Хабаровский край (41). Ни одного вида орхидных неизвестно только из Астраханской области; один вид (Anacamptis laxiflora, всего одно местонахождение) известен из Республики Калмыкия, три вида встречаются в Чукотском автономном округе, восемь - в Ненецком автономном округе. Наиболее широко распространенным видом оказался Gymnadenia conopsea, встречающийся в 74 регионах (90% от общего количества), за ним следуют Corallorhiza trifida (73), Dactylorhiza incarnata (72) и D. viridis (71 регион). 25 видов встречаются в России всего в одном административном регионе, из них 11 - в Сахалинской области и четыре - в Республике Дагестан. Наиболее редкими орхидными России, если судить только по числу местонахождений (исключая виды, предположительно встречающиеся в России), оказываются Cephalanthera erecta, Epipactis euxina и E. leptochila subsp. neglecta, каждый из которых известен из единственного местонахождения. В России отмечаются и два эндемика в ранге вида - Epipactis euxina и E. krymmontana, но оба были описаны лишь недавно, их распространение требует дальнейшего изучения, оба впоследствии могут быть обнаружены на Кавказе за пределами России. В России имеется также один эндемик в ранге подвида, Liparis loeselii subsp. sachalinensis. Но самостоятельность этого таксона подвергается сомнению и возможно, что он является синонимом L. loeselii subsp. loeselii. Neottia krasnojarica и N. ussuriensis, ранее указывавшиеся в качестве эндемиков для территории России, рассматриваются нами как синонимы N. nidus-avis и N. acuminata, соответственно. Таксоны, с наибольшей вероятностью исчезнувшие в России - это Chamorchis alpina и Liparis loeselii subsp. sachalinensis.
Ключевые слова: Orchidaceae, географический анализ, картографирование биоразнообразия, охрана природы, регионы России, редкие виды, сосудистые растения
