Petr G. Efimov, PhD, senior researcher of the Komarov Botanical Institute (197376, Russia, Saint-Petersburg, Professor Popov Street, 2); iD ORCID:; e-mail:
Nikolay N. Panasenko, PhD, assistant professor of the Department of Biology Bryansk State University named after Acad. I.G. Petrovsky (241022, Russia, Bryansk, Bezhitskaya Street, 14); iD ORCID:; e-mail:
Aleksey V. Gornov, PhD, deputy director of the Center for Forest Ecology and Productivity of the Russian Academy of Sciences (117997, Russia, Moscow, Profsoyuznaya Street, 84/32, bldg. 14); iD ORCID:; e-mail:

Reference to article

Efimov P.G., Panasenko N.N., Gornov A.V. 2022. Remnant populations of Cypripedium macranthos (Orchidaceae) in Eastern Europe: evidence of almost complete extinction and widespread introgression with Cypripedium calceolus. Nature Conservation Research 7(2): 1–20.

Section Review articles

The distribution of Cypripedium macranthos and C. × ventricosum in Eastern Europe is described in the literature controversially. At the same time, Cypripedium species are legally protected plants of the highest conservational importance, and detailed knowledge about their historic and current distribution is essential for correct setting of conservation priorities. In the present article, we have assembled primary information about localities of C. macranthos and C. × ventricosum in Russia and Ukraine by referring to the original information sources, mostly to old herbaria and literature. As a result, 19 localities have been estimated as incorrect, 11 as doubtful and only 12 as reliable. Historical absence of C. macranthos and C. × ventricosum in the Republic of Chechnya, Republic of Mari El, Ulyanovsk Region and Kursk Region has been confirmed, whereas its presence in the Republic of Udmurtia, Nizhniy Novgorod Region, Vologda Region, Yaroslavl Region, Republic of Komi, and Voronezh/Lipetsk Regions is questioned. Original herbarium specimens confirming its locality in the Samara Region have been found in the herbarium. The existence of the species in Ukraine is being discussed in view of the information published by V.G. Sobko in 1989, but omitted in later publications. A sharp decrease in the number of localities has been proven. The only extant locality of C. macranthos is situated in the Republic of Chuvashia, and that of C. × ventricosum (1–4 subpopulations) in the Bryansk Region. The amount of hybridisation between C. macranthos and C. calceolus in Eastern Europe was previously underestimated. In the part of the species' distribution centered in Bryansk Region, only hybrids have been recorded since 1903. This fact determines the need for corresponding changes in floristic data and in documents which establish local plant protection.


flora of Russia, floristic change, hybridisation, lady's slipper orchid, old records, plant conservation, relict elements, threatened vascular plants

Artice information

Received: 18.01.2022. Revised: 24.02.2022. Accepted: 26.02.2022.

The full text of the article

Ammosov Yu.N., Belyaev V.A., Bromley G.F., Krivolutskaya G.O., Kupyanskaya A.N., Kurentsova G.E., Nechaev V.A. 1978. A.I. Kurentsov, the scientist. Tula: Priokskoe Publishers. 127 p. [In Russian]
Averyanov L. 1994. Orchids of the Caucasus. Berichte aus den Arbeitskreisen Heimische Orchideen 11(2): 4–45.
Averyanov L.V. 1999. Genus Cypripedium (Orchidaceae) in the Russia. Turczaninowia 2(2): 5–40. [In Russian]
Averyanov L.V. 2000. The genus Cypripedium (Orchidaceae) in Russia. Lindleyana 15(4): 197–221.
Averyanov L.V. 2006. Orchidaceae Juss. In: A.L. Takhtajan (Ed.): Conspectus Florae Caucasi. Vol. 2. Saint-Petersburg: Editio Universitatis Petropolitanae. P. 84–101. [In Russian]
Bakin O.V., Rogova T.V., Sitnikov A.P. 2000. Vascular Plants of Tatarstan. Kazan: Kazan State University. 496 p. [In Russian]
Baranova O.G. 2000. New and rare plants of the Viatka-Kama interfluve. Botanicheskii Zhurnal 85(9): 129–131. [In Russian]
Baranova O.G. 2006. The peculiarities of the Orchidaceae Juss. family species distribution in the Udmurt Republic and their protection. Bulletin of Udmurt University 3(10): 3–10. [In Russian]
Baranova O.G. (Ed.). 2012. Red Data Book of the Udmurt Republic. 2nd ed. Cheboksary: Perfectum. 458 p. [In Russian]
Baranova O.G., Adakhovskiy D.A., Borisovskiy A.G., Dedyukhin S.V., Zubtsovskiy N.E., Perevoshchikov A.A., Markova E.M., Rubtsova A.V., Tychinin V.A., Tyulkin Yu.A. 2011. Rare and endangered plants and animals of the south part of Udmurtia and their protection: results of the research undertaken in 2005–2009. Izhevsk: Udmurt State University. 272 p. [In Russian]
Baranova O.G., Ilminskikh N.G., Puzyrev A.N., Tuganaev V.V. 1992. Udmurtia Flora Conspectus. Izhevsk: Udmurt State University. 141 p. [In Russian]
Baranova O.G., Puzyrev A.N. 2012. Conspectus Florae Provinciae Udmurtiensis (Plantae Vasculares). Moscow; Izhevsk: Institute of Computer Research. 212 p. [In Russian]
Bardunov L.V., Novikov V.S. (Eds.). 2008. Red Data Book of the Russian Federation (plants and fungi). Moscow: KMK Scientific Press Ltd. 855 p. [In Russian]
Biryukova O.V., Vorotnikov V.P., Mininzon I.L. 2014. The Orchidaceae family in the flora of the Nizhni Novgorod Region. Vestnik of Lobachevsky State University of Nizhni Novgorod 3(3): 16–25. [In Russian]
Bolotova V.M., Dedov A.A., Lashchenkova A.N., Tolmachev A.I., Sholeninova T.P. 1962. Manual of the vascular plants of the Republic of Komi. Moscow; Leningrad: AS USSR. 359 p. [In Russian]
Bondartseva M.A., Prisnyi Yu.A. (Eds.). 2021. Red Data Book of the Orel Region. Orel: Papirus. 440 p. [In Russian]
Bordzilovski E.I. 1950. Orchidaceae Lindl. In: M.I. Kotov, A.I. Barbarich (Eds.): Flora URSR. 2nd ed. Vol. 3. Kiev: AS URSR. P. 312–401. [In Ukrainian]
Borodin A.M. (Ed.). 1984. Red Data Book of the USSR. 2nd ed. Moscow: Lesnaya promyshlennost. 480 p. [In Russian]
Bosek P.Z. 1975. Plants of the Bryansk Region. Bryansk: Priokskoye. 464 p. [In Russian]
Bosek P.Z. 1982. Plants. In: N.Z. Kharitonova, E.S. Murakhtanova (Eds.): Rare and protected animals and plants of the Bryansk Region. Bryansk: Bryansk Regional Publishing House. P. 133–205. [In Russian]
Bulokhov A.D., Panasenko N.N., Semenishchenkov Yu.A., Sitnikova E.F. (Eds.). 2016. Red Data Book of the Bryansk Region. 2nd ed. Bryansk: RIO BGU. 432 p. [In Russian]
Bulokhov A.D., Velichkin E.M. 1998. A manual of plants of the southeast non-Chernozem Russia (Bryansk, Kaluga, Smolensk Regions). 2nd ed. Bryansk: BGPU. 380 p. [In Russian]
Cholovsky K.A. 1882. A review of the Flora of Mohilev Province. Mohilev: Mohilev Province Administration Publishers. 188 p. [In Russian]
Claus C. 1851. Localfloren der Wolgagegenden. St. Petersburg: Buchdrukerei der Kaiserlichen Akademie der Wissenschaften. 324 p.
Dimitriev A.V. (Ed.). 2001. Red Data Book of the Republic of Chuvashia. Cheboksary: IPK Chuvashia. 275 p. [In Russian]
Efimov P.G., Gafurova M.M., Leostrin A.V., Melnikov D.G., Senator S.A., Fateryga A.V. 2018. New data on distribution of Orchidaceae species in several Regions of Russia. Botanicheskii Zhurnal 103(7): 923–930. DOI: 10.7868/S0006813618070062 [In Russian]
Efimov P.G. 2020. Orchids of Russia: annotated checklist and geographic distribution. Nature Conservation Research 5(Suppl.1): 1–18. DOI: 10.24189/ncr.2020.018
Efimova T.P. 1972. A manual of the plants of Udmurtia. Izhevsk: Udmurtia. 224 p. [In Russian]
Fardeeva M.B. 2006. Ecological analysis of the orchid flora of Tatarstan. In: L.A. Zhukova (Ed.): Principles and methods in biodiversity conservation. Yoshkar-Ola: Mari El State University. P. 51–53. [In Russian]
Fedotov Yu.P. (Ed.). 2004. Red Data Book of the Bryansk Region. Plants. Fungi. Bryansk: Chitay-Gorod. 272 p. [In Russian]
Fedotov Yu.P., Evstigneev O.I. 2003. Records of rare species of orchids (Orchidaceae) in Bryansk Region. Botanicheskii Zhurnal 88(10): 118–121. [In Russian]
Gafurova M.M. 2014. Vascular plants of Chuvash Republic. Vol. 3 of the Flora of the Volga River Basin. Togliatti: Kassandra. 333 p. [In Russian]
Gafurova M.M. (Ed.). 2019. Red Data Book of the Republic of Chuvashia. Vol. 1, part 1. 2nd ed. Moscow: Buki Vedi. 332 p. [In Russian]
Geltman D.V., Matveeva N.V. 2018. B.S. Kharitontsev Features of the genesis of phytostroma of Russia (ecological aspect): a monograph. Tobolsk: the Division of the Tyumen State University in Tobolsk, 2015. 161 p. B. S. Kharitontsev. Floristic pressure in phytostroma of Siberia: A monograph. Tobolsk: the Division of the Tyumen State University in Tobolsk, 2016. 186 p. Botanicheskii Zhurnal 103(2): 271–277. [In Russian]
Goworuchin W.S. 1937. Flora Medio, Boreali et Polari Uralensis. Sverdlovsk: Sverdlovsk Regional Publishing House. 536 p. [In Russian]
Grossheim A.A. 1940. Flora of Caucasus. Vol. 2. 2nd ed. Baku: Azerbaijanian Branch AS USSR. 284 p. [In Russian]
Ivanov A.L. 2019. Conspectus florae Caucasi Rossicae (plantae vasculares). Stavropol: SKFU. 341 p. [In Russian]
Ivanov A.L., Kovaleva O.A. 2005. Orchids of the North Caucasus. Stavropol: SKFU. 122 p. [In Russian]
Jakubska-Busse A., Tsiftsis S., Śliwiński M., Křenová Z., Djordjević V., Steiu C., Kolanowska M., Efimov P., Hennigs S., Lustyk P., Kreutz K. 2021. How to protect natural habitats of rare terrestrial orchids effectively: a comparative case study of Cypripedium calceolus in different geographical regions of Europe. Plants 10: 404. DOI: 10.3390/plants10020404
Khitrovo V.N. 1910. Critical notes on the Flora of the Orel Province. IV. Proceedings of the Society for the study of the nature of the Orel Province 2: 157–185. [In Russian]
Khitrovo V.N. 1923. Account of the Flora of the Orel Province. A manuscript kept in the Saint-Petersburg Branch of the Archive of the Academy of Sciences. 114 p. [In Russian]
Kiseleva L.L., Parakhina E.A., Shcherbakov A.V. 2021. Checklist of vascular plants of the Orel Region. Moscow: Galleya-Print. 78 p. [In Russian]
Kleopov Yu.P. 1933. Orchidaceae. In: M. Lyaskivskiy (Ed.): Flora URSR. Part 1. Kiev: Kolkhoz and Sovkhoz Literature State Publishers. P. 327–344. [In Ukrainian]
Knyasev M.S., Kulikov P.V., Knyazeva O.I., Semerikov V.L. 2000a. Interspecific hybridization in Eurasian species of Cypripedium (Orchidaceae), and the taxonomic status of C. ventricosum. Botanicheskii Zhurnal 85(5): 94–102. [In Russian]
Knyasev M.S., Kulikow P.V., Knyaseva O.I., Semerikov V.L. 2000b. Interspecific hybridization in Northern Eurasian Cypripedium: morphometric and genetic evidence of hybrid origin of C. ventricosum. Lindleyana 15(1): 10–20.
Konechnaya G.Yu., Suslova T.A. (Eds.). 2004. Red Data Book of the Vologda Region. Vol. 2. Vologda: Vologda State University and Rus' Press. 360 p. [In Russian]
Kozlovskaya N.V. 1978. Flora of Belarus, rules of its formation, scientific basis for its use and protection. Minsk: Nauka i Tekhnika. 128 p. [In Russian]
Kryshtofovich A.N. 1929. Orchidaceae. In: B.A. Fedtschenko (Ed.): Flora Rossiae Austro-Orientalis. Fasc. 3. Leningrad: Main Botanical Garden. P. 414–436. [In Russian]
Krylov P.N. 1885. Materialien zur flora des gouvernements Wjatka. Proceedings of the Society of Naturalist at the Kazan' Imperial University 14(1): 1–131. [In Russian]
Kulikov P.V., Filippov E.G. 2011. Short-spurred fragrant Orchid (Gymnadenia odoratissima (L.) Rich.) in Russia. In: I.I. Shamrov (Ed.): Protection and cultivation of orchids. Moscow: Moscow: KMK Scientific Press Ltd. P. 266–271. [In Russian]
Lebed'ko V.N. 2017. Orchidaceae Juss. In: V.I. Parfenov (Ed.): Flora of Belarus. Minsk: Belaruskaya Nauka. P. 232–310. [In Russian]
Lipschitz S. 1952. Botanicorum Rossicorum Lexicon Biographo-Bibliographicum. Vol. 5. Moscow: Editio Societatis Naturae Curiosum Mosquensis. 617 p. [In Russian]
Litvinov D.I. 1927. On some botanical-geographical proportions in our flora. Leningrad. 15 p. [In Russian]
Maltsev A.N. 1907. A review of the vegetation of Korocha County of Kursk Province. Protocols of the Society of Naturalists at the Imperial Yur'yev University 16(1): 1–56. [In Russian]
Marakaev O.A. 2015. Orchids of Yaroslavl Region: a photoatlas. Yaroslavl: Yaroslavl State University. 50 p. [In Russian]
Markov M.V. 1939. Wald und Steppe in Bedingungen des Sakamjas. II. Die Nadelwälder. Scientific Notes from the Kazan' State University 99(1–1): 67–131. [In Russian]
Martynenko V.A. 1976. Orchidaceae Juss. In: A.I. Tolmatchev (Ed.): Flora regionis Boreali-Orientalis territoriae Europaeae URSS. Vol. 2. Leninopoli: Nauka. P. 118–133. [In Russian]
Mayevskiy P.F. 1917. Flora of the Middle European Russia. 5th ed. Moscow: M. & S. Sabashnikovy. 909 p. [In Russian]
Mayevskiy P.F. 1933. Flora of the Middle European Russia. 6th ed. Moscow & Leningrad: Sel'hozgiz. 748 p. [In Russian]
Mayevskiy P.F. 1941. Flora of the Middle European USSR. 7th ed. Moscow & Leningrad: Sel'hozgiz. 824 p. [In Russian]
Mayevskiy P.F. 1954. Flora of the Middle European USSR. 8th ed. Moscow & Leningrad: State Agricultural Literature Publishing House. 910 p. [In Russian]
Mayevskiy P.F. 1964. Flora of the Middle European USSR. 9th ed. Leningrad: Kolos. 880 p. [In Russian]
Mayevskiy P.F. 2006. Flora of the Middle European Russia. 10th ed. Moscow: Moscow: KMK Scientific Press Ltd. 600 p. [In Russian]
Mayevskiy P.F. 2014. Flora of the Middle European Russia. 11th ed. Moscow: KMK Scientific Press Ltd. 635 p. [In Russian]
Montrezor V. 1881. A review of the most beautiful plants in the flora of Kiev Academic District: Kiev, Podol'sk, Volhyn', Chernigov and Poltava Provinces. Kiev: Kiev Society of Horticulturalists. 49 p. [In Russian]
Montrezor V. 1886. A review of the plants in the Flora of Kiyev Academic District: Kiev, Podol'sk, Volhyn', Chernigov and Poltava Provinces. Notes of the Kiev Society of Naturalists 7(1): 1–144. [In Russian]
Mosyakin S.L., Fedoronchuk N.M. 1999. Vascular plants of Ukraine: A nomenclatural checklist. Kiev: M.G. Kholodny Institute of Botany. 345 p. DOI: 10.13140/2.1.2985.0409
Nazirov A.A. (Ed.). 2016. Red Data Book of the Republic of Tatarstan. 3rd ed. Kazan: Idel-Press. 760 p.
Nevski S.A. 1935. Orchidaceae Lindl. In: V.L. Komarov (Ed.): Flora URSS. Vol. 4. Leningrad: Editio Academiae Scientiarum URSS. P. 589–730, 750–754. [In Russian]
Nevski S.A. 1949. Orchidaceae Lindl. In: B.K. Shishkin (Ed.): Flora of BSSR. Vol. 1. Moscow: State Agricultural Literature Publishing House. P. 377–414. [In Russian]
Nyankovskiy M.A. (Ed.). 2015. Red Data Book of the Yaroslavl Region. Yaroslavl: Akademia 76. 472 p. [In Russian]
Okhapkin A.G. (Ed.). 2005. Red Data Book of the Nizhniy Novgorod Region. Vol. 2. Nizhniy Novgorod: Committee for Nature Conservation and Land-Use Management of the Nizhniy Novgorod Region. 328 p. [In Russian]
Protopopova V.V. 1987. Orchidaceae. In: Yu.N. Prokudin (Ed.): Manual of the Higher Plants of Ukraine. Kiev: Naukova Dumka. P. 405–412. [In Russian]
Perfiljev I.A. 1934. Flora of the Severnyi Krai. Part 1. Arkhangelsk: Sevkraigiz. 160 p. [In Russian]
Plaksina T.I. 2001. Conspectus of the flora of the Volga-Ural Region. Samara: Samara State University. 388 p. [In Russian]
Prisnyi A.V. (Ed.). 2005. Red Data Book of the Belgorod Region. Belgorod: Ecological Fund of the Belgorod Region. 532 p. [In Russian]
Privalova L.A., Prokudin Yu.N. 1959. Addenda et corrigenda ad vol. I «Florae Tauricae». Transactions of the State Nikitskiy Botanical Garden 31: 5–127. [In Russian]
Rakov N.S., Saksonov S.V., Senator S.A., Vasjukov V.M. 2014. Vascular plants of the Ulyanovsk Region. Vol. 2 of the Flora of the Volga River Basin. Togliatti: Kassandra. 295 p. [In Russian]
Rännbäck L.M. 2007. Propagation, cultivation and breeding of terrestrial temperate orchids, with focus on Cypripedium spp. Bachelor project in the Danish-Swedish Horticulture programme. Alnarp: SLU. 50 p.
Reshetiuk O.W. 2003. The genus Cypripedium L. (Orchidaceae Juss.) in Ukraine (classification, horology, phytocenology and its species protection ex situ and in situ). PhD Thesis. Kyiv: M.M. Gryshko National Botanical Gardens, National Academy of Sciences of Ukraine. 20 p. [In Ukrainian]
Rogowitsch A.S. 1855. A review of vascular and semivascular plants in the flora of Kiev, Chernigov and Poltava. In: Natural History of the provinces of Kiev Academic District. Kiev. 146 p. [In Russian]
Rogowitsch A.S. 1869. A review of vascular and higher cryptogamic plants in the flora of the Provinces of the Kiev Academic District: Volhyn', Podol'sk, Kiev, Chernigov and Poltava. Kiev. 308 p. [In Russian]
Ruprecht F. 1866. Geobotanical studies of Chernozem. Saint-Petersburg: Imperial Academy of Sciences. 131 p. [In Russian]
Saksonov S.V., Senator S.A. 2012. Guide to the Samara flora 1851–2011. Vol. 1 of the Flora of the Volga River Basin. Togliatti: Kassandra. 511 p. [In Russian]
Saksonov S.V., Koneva N.V. 2006. The abstract of Orchidaceae family the Samara Area. Bulletin of Udmurt University 10: 43–50. [In Russian]
Schmalhausen I. 1897. Flora of the Middle and South Russia, Crimea and North Caucasus. Vol. 2. Kiev: I.N. Kushnerev i Ko. 752 p. [In Russian]
Shchepovskikh A.I. (Ed.). 1995. Red Data Book of the Republic of Tatarstan. Kasan: Priroda; Star. 454 p. [In Russian]
Shchepovskikh A.I. (Ed.). 2006. Red Data Book of the Republic of Tatarstan. 2nd ed. Kasan: Idel-Press. 832 p. [In Russian]
Smirnov A.G. 1979. Orchidaceae. In: M.V. Markov (Ed.): A manual of plants of the Tatar Soviet Republic. Kazan: Kazan State University. P. 108–115. [In Russian]
Smirnov V. 1904. Zur Flora des Gouv. Simbirsk. Supplement to the Protocols of the Proceedings of the Society of Naturalist at the Kazan' Imperial University 231: 1–24. [In Russian]
Smolyaninova L.A. 1976. Orchidaceae Juss. In: A.A. Fedorov (Ed.): Flora Partis Europaeae URSS. Vol. 2. Leningrad: Nauka. P. 10–59. [In Russian]
Sobko V.G. 1989. Orchids of Ukraine. Kiev: Naukova Dumka. 190 p. [In Ukrainian]
Sobolev N.A., Belonovskaya E.A. (Eds.). 2013. Emerald Data Book of the Russian Federation. Conservationally Important Protected Areas of European Russia and their elucidation. Part 1. Moscow: Institute of Geography. 308 p. [In Russian]
Thiers B.M. 2022. Index Herbariorum. Available from
Tikhomirov V.N. (Ed.). 1986. A manual of the flora of the Yaroslavl Region. Yaroslavl: Verkhnyaya Volga. 182 p.
Tikhomirov V.N., Kharitontsev B.S. 1984. New data about the flora of the left-bank of Desna River within the Bryansk Region. Biological Sciences 8(248): 73–76. [In Russian]
Ueda K. (Ed.). 2021a. Cypripedium macranthos. In: iNaturalist: A community for Naturalists. Available from
Ueda K. (Ed.). 2021b. Cypripedium macranthos. In: iNaturalist: A community for Naturalists. Available from
Ueda K. (Ed.). 2021c. Cypripedium macranthos. In: iNaturalist: A community for Naturalists. Available from
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]
Wirzen J.E.A. 1839. De geographica plantarum per partem provinciae Casanensis distributione. Helsingforsiae: Frenckelliana. 130 p.
Yudin U.P. 1963. Relict flora of the limestone outcrops of the North-East of the European part of the USSR. In: V.N. Sukaczev (Ed.): Materials on the history of the flora and vegetation of the USSR. Fasc. 4. Moscow; Leningrad: AS USSR. P. 493–587. [In Russian]
Zinger V.Ya. 1886. A Compendium of the Middle Russian Flora. Moscow: M. Katkov. 520 p. [In Russian]