J. Eur. Orch. 46 (2): 407 - 436. 2014.

Alexander V. Fateryga and Karel C.A.J. Kreutz

Checklist of the orchids of the ()

Keywords Orchidaceae, checklist of species, new nomenclature combinations, hybrids, flora of the Crimea.

Summary Fateryga, A.V. & C.A.J. Kreutz (2014): Checklist of the orchids of the Crimea (Orchidaceae).- J. Eur. Orch. 46 (2): 407-436. A new nomenclature checklist of the Crimean orchids with 49 taxa and 16 hybrids is proposed. Six new taxa are added and ten taxa are excluded from the latest checklist of the Crimean vascular flora published by YENA (2012). In addition, five nomenclature changes are proposed: Epipactis persica (Soó) Nannf. subsp. taurica (Fateryga & Kreutz) Fateryga & Kreutz comb. et stat. nov., (L.) L. var. wanjkovii (E. Wulff) Fateryga & Kreutz stat. nov., ×simorrensis (E.G. Camus) H. Kretzschmar, Eccarius & H. Dietr. nothosubsp. ticinensis (Gsell) Fateryga & Kreutz stat. nov., ×Dactylocamptis uechtritziana (Hausskn.) B. Bock ex M. Peregrym & Kuzemko nothosubsp. magyarii (Soó) Fateryga & Kreutz comb. et stat. nov., and Orchis ×beyrichii Kern. nothosubsp. mackaensis (Kreutz) Fateryga & Kreutz comb. et stat. nov. Moreover, a new variety, Limodorum abortivum (L.) Sw. var. viridis Fateryga & Kreutz var. nov. is described.

Zusammenfassung Fateryga, A.V. & C.A.J. Kreutz (2014): Eine Übersicht der Orchideen der Krim (Orchidaceae).- J. Eur. Orch. 46 (2): 407-436. Eine neue nomenklatorische Liste der Orchideen der Krim mit 49 Taxa und 16 Hybriden wird vorgestellt. Sechs Arten sind neu für die Krim. Zehn Taxa, die noch bei YENA (2012) in seiner Checklist aufgelistet wurden, kommen auf der Krim nicht vor und wurden gestrichen. Fünf nomenklatorische Änderungen

Journal Europäischer Orchideen 46 (2): 2014. 407 werden durchgeführt: Epipactis persica (Soó) Nannf. subsp. taurica (Fateryga & Kreutz) Fateryga & Kreutz comb. et stat. nov., Orchis mascula (L.) L. var. wanjkovii (E. Wulff) Fateryga & Kreutz stat. nov., Anacamptis ×simorrensis (E.G. Camus) H. Kretzschmar, Eccarius & H. Dietr. nothosubsp. ticinensis (Gsell) Fateryga & Kreutz stat. nov., ×Dactylocamptis uechtritziana (Hausskn.) B. Bock ex M. Peregrym & Kuzemko nothosubsp. magyarii (Soó) Fateryga & Kreutz comb. et stat. nov. und Orchis ×beyrichii Kern. nothosubsp. mackaensis (Kreutz) Fateryga & Kreutz comb. et stat. nov. Schließlich wird eine neue Varietät, Limodorum abortivum (L.) Sw. var. viridis Fateryga & Kreutz var. nov. beschrieben.

Резюме Фатерыга, А.В. & К.А.Й. Кройтц (2014): Чеклист орхидей Крыма (Orchidaceae).- J. Eur. Orch. 46 (2): 407-436. Предложен новый номенклатурный чеклист орхидей Крыма, насчитывающий 49 таксонов и 16 гибридов. По сравнению с последним чеклистом сосудистой флоры Крыма, опубликованным ЕНОЙ (2012), добавлено шесть таксонов и исключено десять. Предложено пять номенклатурных изменений: Epipactis persica (Soó) Nannf. subsp. taurica (Fateryga & Kreutz) Fateryga & Kreutz comb. et stat. nov., Orchis mascula (L.) L. var. wanjkovii (E. Wulff) Fateryga & Kreutz stat. nov., Anacamptis ×simorrensis (E.G. Camus) H. Kretzschmar, Eccarius & H. Dietr. nothosubsp. ticinensis (Gsell) Fateryga & Kreutz stat. nov., ×Dactylocamptis uechtritziana (Hausskn.) B. Bock ex M. Peregrym & Kuzemko nothosubsp. magyarii (Soó) Fateryga & Kreutz comb. et stat. nov. и Orchis ×beyrichii Kern. nothosubsp. mackaensis (Kreutz) Fateryga & Kreutz comb. et stat. nov. Кроме того, описана новая разновидность Limodorum abortivum (L.) Sw. var. viridis Fateryga & Kreutz var. nov.

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Introduction

The Crimean Peninsula is situated between 44°23' and 46°19' of northern latitude and 32°30' and 36°40' of eastern longitude. The distance from the most northern point of the peninsula to the most southern one is 207 km; that from 2 east to west is 324 km. The area of the Crimea is about 26,900 km (BAGROVA & al. 2001). The peninsula is divided to two main orographical parts: a large flat part on the north and mountain part on the south. The Crimean Mountains

408 Journal Europäischer Orchideen 46 (2): 2014. stretch for 180 km from south-west to north-east (from Sevastopol to Feodosiya) and consist of three main ridges. A total of 47 landscape types can be recognized in the Crimea; eight belonging to the flat part of the peninsula and 39 to the mountain one. These landscapes comprise seven major habitat zones: semi-desert steppe and saline lands, true steppes (mainly tilled), premontane forest steppes, forests of the northern mountain slopes, mountain meadows and yayla (= alpine meadow) steppes, forests of the southern mountain slopes, and submediterranean vegetation of the south coast (BIODIVERSITY SUPPORT PROGRAM 1999: map 2). According to the latest review (YENA 2012: 201-204), the vascular flora of the Crimea includes 127 families, 760 genera and 2,536 species and subspecies, including 106 endemics, 156 neophytes and 12 extinct taxa.

The history of the study of the Crimean orchids covers more than 200 years. The first significant floristic contributions to the knowledge of the Crimean vascular flora (including orchids) were published by such famous botanists as PALLAS (1795), MARSCHALL VON BIEBERSTEIN (1808), and VON STEVEN (1857) (YENA 2012: 146-150; HAHN 2012: 9-59). However, the first comprehensive review of the Crimean orchids which should be noticed specially was published by WULFF (1930: 77-124). He listed 44 species with two additional varieties and 10 hybrids; this list was an excellent basis for further contributions. NEVSKI (1935: 589-730) listed only 38 orchid species in the Crimean flora, combining several pairs of closely related taxa with each other. Additional corrections to Wulff’s list were published by PRIVALOVA & PROKUDIN (1959: 72-80) who added one species, namely (L.) Chevall., and excluded two others which had been in their opinion incorrectly identified. The next important list of the Crimean orchids was published as an identification key of the vascular of the Crimea (RUBTZOV 1972: 96-103); this also listed 38 orchid species. Four years later SMOLYANINOVA (1976: 10-59) listed 44 species and 3 additional subspecies of Orchis mascula (L.) L. In comparison with WULFF (l.c), she also added Epipactis atrorubens (Hoffm.) Besser and excluded S. spiralis. PROKUDIN (1987: 405-412) listed 42 orchid species in a key for the vascular flora of Ukraine. Both RUBTZOV (l.c.) and PROKUDIN (l.c.) did not add any data on new findings of any taxa but obviously used mainly the data already published by WULFF (l.c.). The different numbers of species in their lists were only the result of different opinions regarding the of several problematic species. In SOBKO (1989: 73-177) 42 species of orchids were listed for the Crimean flora. A further new species for the flora of the Crimea was (L.) R.M. Bateman, Pridgeon & M.W. Chase discovered by ALEKSEEV (1993: 101- 102). All data on the Crimean orchids available up to the end of the 20th century were summarized by GOLUBEV (1996, republished in 2008: 77-78) and

Journal Europäischer Orchideen 46 (2): 2014. 409 POPKOVA (2001: 39-53). Their lists were identical and contained in total 47 species. Both lists were synoptic and made without any critical review of available herbarium material; both lists were therefore similar to the data provided by SMOLYANINOVA (1976: 10-59) but with N. ustulata and without Orchis mascula (L.) L. subsp. pinetorum (Boiss. & Kotschy) E.G. Camus which had been listed in the regional flora since WULFF (1930: 98). Other subspecies of O. mascula were regarded as separate species.

At the beginning of the 21th century three new orchid species were added to the flora of the Crimea: Himantoglossum affine (Boiss.) Schltr. (YENA & al. 2008: 472), Epipactis purpurata Smith and Epipactis condensata Boiss. ex D.P. Young (EFIMOV 2008: 72-77). However, the first was later synonymized with Himantoglossum caprinum (M. Bieb.) Spreng. (SRAMKÓ & al. 2012: 21- 24; MOLNÁR & al. 2012: 8-10). The record of N. ustulata and the erroneous record of Orchis signifera Vest = Orchis mascula subsp. speciosa (Mutel) Hegi, published by SMOLYANINOVA (1976: 44) were not included in the Red Book of Ukraine (DIDUKH 2009: 203-211) which listed 45 orchid species for the Crimea. After that, the most recent comprehensive, but still synoptic, list of the Crimean orchids was published by YENA (2012: 146-150), where a total of 50 species and subspecies were listed. This list was based on GOLUBEV (2008: 77-78) but excluding Orchis mascula (L.) L. subsp. wanjkovii (E. Wulff) Soó (which was reduced to O. mascula s. str.) and adding S. spiralis, two Epipactis species listed by EFIMOV (2008: 72-77) and Epipactis persica (Soó) Nannf. The latter species was added to the flora of the Crimea for the first time under that name although had previously been listed by WULFF (1930: 116) under the name “Epipactis latifolia (L.) All. var. latifolia f. gracilis Dager.” now considered conspecific with Epipactis albensis Nováková & Rydlo which does not grow in the Crimea. Several new floristic findings and taxonomical opinions (mainly relative to the genus Epipactis) were not included into Yena’s list because they were published later (FATERYGA & KREUTZ 2012: 201-202; HAHN 2012: 33-37; KREUTZ & FATERYGA 2012: 713-715; FATERYGA & al. 2013: 652-654; FATERYGA & al. 2014: 24) or related only to the new variety (L.) Rich. var. orientalis Kreutz (KREUTZ 2011: 276-278) (Yena’s list was restricted to species and subspecies).

The purpose of the present paper is to revise the list of Crimean orchids thereby published by YENA (2012: 146-150). In addition all known orchid hybrids occurring in the Crimea are listed (Yena’s list did not include hybrids).

410 Journal Europäischer Orchideen 46 (2): 2014. Materials and methods

This study was made on the basis of our field observations during the period 2003-2014 and by studying of six herbarium collections in and Ukraine: the Herbarium of the Crimean Agrotechnological University in (CSAU), the Herbarium of the Nikitskiy Botanical Garden in Yalta (YALT), the Herbarium of Vernadskiy Taurida National University in Simferopol (SIMF), the Herbarium of the Karadag Nature Reserve in Feodosiya (PHEO), the Herbarium of Komarov Botanical Institute of the Russian Academy of Sciences in Saint-Petersburg (LE), and the National Herbarium of Ukraine in Kiev (KW). All possible literature sources cited above were also examined.

The taxa included in the checklist are arranged in alphabetical order. There are four additional sections after the main checklist. “Notae” contains comments to several taxa including the names used by YENA (2012: 146-150), new taxonomical and nomenclatural combinations, and other information. “Addenda” contains new taxa added in comparison with the list published by YENA (l.c.) with their relevant references. “Excludendae” contains the annotated list of excluded taxa, again in comparison with the list from YENA (l.c.). “Hybrids” contains a list of all orchid hybrids listed for the flora of the Crimea in literature or added by our field observations.

Results

A total of 44 species with five additional infraspecific taxa (two subspecies and three varieties) are today present in the Crimea.

Anacamptis coriophora (L.) R.M. Bateman, Pridgeon & M.W. Chase (L.) R.M. Bateman, Pridgeon & M.W. Chase subsp. caucasica (K. Koch) H. Kretzschmar, Eccarius & H. Dietr. Anacamptis palustris (Jacq.) R.M. Bateman, Pridgeon & M.W. Chase subsp. elegans (Heuff.) R.M. Bateman, Pridgeon & M.W. Chase Anacamptis pyramidalis (L.) Rich. var. orientalis Kreutz (Mill.) Druce Cephalanthera longifolia (L.) Fritsch (L.) Rich. viride (L.) Hartm. Comperia comperiana (Steven) Asch. & Graebn.

Journal Europäischer Orchideen 46 (2): 2014. 411 Corallorhiza trifida Châtel. L. iberica (M. Bieb. ex Willd.) Soó (L.) Soó Dactylorhiza romana (Sebast.) Soó (L.) Crantz subsp. levantina Kreutz, Óvári & Shifman Epipactis helleborine (L.) Crantz subsp. orbicularis (K. Richt.) E. Klein Epipactis krymmontana Kreutz, Fateryga & Efimov Epipactis microphylla (Ehrh.) Sw. Epipactis muelleri Godfery (L.) Crantz Epipactis persica (Soó) Nannf. subsp. persica Epipactis persica (Soó) Nannf. subsp. taurica (Fateryga & Kreutz) Fateryga & Kreutz Epipactis turcica Kreutz Sw. Goodyera repens (L.) R. Br. (L.) R. Br. Himantoglossum caprinum (M. Bieb.) Spreng. Limodorum abortivum (L.) Sw. var. abortivum Limodorum abortivum (L.) Sw. var. rubrum H. Sund. ex Kreutz Limodorum abortivum (L.) Sw. var. viridis Fateryga & Kreutz Listera ovata (L.) R. Br. (Scop.) R.M. Bateman, Pridgeon & M.W. Chase Neotinea ustulata (L.) R.M. Bateman, Pridgeon & M.W. Chase Neottia nidus-avis (L.) Rich. Huds. Ophrys mammosa Desf. subsp. taurica (Aggeenko) Soó Ophrys oestrifera M. Bieb. Orchis mascula (L.) L. var. mascula Orchis mascula (L.) L. var. wanjkovii (E. Wulff) Fateryga & Kreutz L. subsp. stevenii (Rchb. f.) B. Baumann & al. L. Orchis provincialis Balb. ex Lam. & DC. Orchis punctulata Steven ex Lindl. Huds. Orchis simia Lam. (L.) Rich. Platanthera chlorantha (Custer) Rchb. Steveniella satyrioides (Spreng.) Schltr. Traunsteinera globosa (L.) Rchb.

412 Journal Europäischer Orchideen 46 (2): 2014. Notae

Anacamptis morio (L.) R.M. Bateman, Pridgeon & M.W. Chase is only represented by A. morio (L.) R.M. Bateman, Pridgeon & M.W. Chase subsp. caucasica (K. Koch) H. Kretzschmar, Eccarius & H. Dietr. This subspecies is quite variable in its floral morphology (fig. 1-2). Different forms were obviously incorrectly misidentified for two different taxa, A. morio (L.) R.M. Bateman, Pridgeon & M.W. Chase subsp. morio and A. morio (L.) R.M. Bateman, Pridgeon & M.W. Chase subsp. picta (Loisel.) P. Jacquet & Scappat. Our observations did not confirm the presence of both taxa in the Crimea. Only A. morio subsp. caucasica was also listed by HAHN (2012: 26). Anacamptis palustris (Jacq.) R.M. Bateman, Pridgeon & M.W. Chase is only represented by its subspecies A. palustris (Jacq.) R.M. Bateman, Pridgeon & M.W. Chase subsp. elegans (Heuff.) R.M. Bateman, Pridgeon & M.W. Chase (fig. 3). Previously it was incorrectly identified for two different taxa, A. palustris (Jacq.) R.M. Bateman, Pridgeon & M.W. Chase and Anacamptis laxiflora (Lam.) R.M. Bateman, Pridgeon & M.W. Chase due to the variability of its floral morphology. Also only the subspecies A. palustris subsp. elegans is listed by HAHN (2012: 27). Anacamptis pyramidalis (L.) Rich. is only represented by A. pyramidalis (L.) Rich. var. orientalis Kreutz (KREUTZ 2011: 276-278) (fig. 4). The presence of the typical variety in the Crimea requires further investigation. Dactylorhiza romana (Sebast.) Soó requires further taxonomic investigation. There are two different forms in the Crimea: a form with white-yellowish and one with pink or purple flowers. The form with pink or purple flowers occurs only at the south coast of the Crimea while the white-yellowish form is additionally present on the northern slopes of the Crimean Mountains and in their foothills. Both the white-yellowish and pink or purple forms growing at the south coast (fig. 5-6) differ from the white-yellowish plants growing in different habitat zones (fig. 7-8) by having somewhat broader (like those of D. romana (Sebast.) Soó subsp. georgica (Klinge) Soó ex Renz & Taubenheim) as well slightly larger flowers. However, the length of the spur of all plants in the Crimea is normal for typical D. romana. Epipactis persica (Soó) Nannf. subsp. taurica (Fateryga & Kreutz) Fateryga & Kreutz comb. et stat. nov. Basionym: Epipactis taurica Fateryga & Kreutz, J. Eur. Orch. 44 (1): 201 (2012). Comments: This subspecies differs from typical E. persica only by a robust stem, rigid and erect leaves, brighter coloration of the lip, and rarely by a completely green pedicel (fig. 9-10); other important characters in the floral morphology are identical. The two taxa grow in different habitats in the

Journal Europäischer Orchideen 46 (2): 2014. 413 Crimea: E. persica subsp. persica occurs mainly on northern slopes of the Crimean Mountains, while E. persica subsp. taurica grows on the southern slopes and foothills. Limodorum abortivum (L.) Sw. is represented by the typical variety (fig. 11), L. abortivum var. rubrum H. Sund. ex Kreutz (HAHN 2012: 41) (fig. 12), and by a new variety as described below. Limodorum abortivum (L.) Sw. var. viridis Fateryga & Kreutz var. nov. Diagnosis: As Limodorum abortivum var. abortivum but stem shorter, thinner, and completely green; denser, flowers slightly smaller. Holotype: Крым, окр. Алушты, близ с. Виноградное, гора Урага, дубовый лес [Crimea, vicinity of Alushta, near Vinogradnoye village, Mt. Uraga, oak forest], 29.5.2014, leg. A.V. Fateryga, CSAU, fig. 13. Isotypes (with the same labels): CSAU (1 sheet) and PHEO (2 sheets). Paratypes: Крым, окр. Ялты, Мисхор, между трассой Ялта - Севастополь, и нижней станцией канатной дороги, сосновый лес [Crimea, vicinity of Yalta, Miskhor, between the highway “Yalta - Sevastopol” and the lower station of the funicular, pine forest], 28.5.2012, leg. A.V. Fateryga, PHEO (1 sheet); ibid., 28.5.2014, leg. A.V. Fateryga, CSAU (1 sheet). Description: Terrestrial plant, erect, 15-40 cm height, with 1-2 stems. Stem 1.5-2.5 mm thick, completely green. Leaves (6-10) -like, 2-4.5 × 0.5-0.7 cm, pale-green, much paler than the stem. Inflorescence 2.5-6 cm long and 3.5- 4.5 cm wide, subdense, with 3-12 flowers. similar to the leaves but some shorter and narrower, longer than the ovaries. Ovaries 15-20 × 2-2.5 mm, green. Flowers rather small, spreading. Sepals pale, pinkish-violet, ovate-lanceolate, 16-18 × 4-5 mm; petals 11-13 × 2.5-3.5 mm, of the same color with the sepals. Hypochile 5-6 mm long, whitish. Epichile ovate, 11-13 × 6-7 mm, channeled, sides rolled up and scalloped, violet. Spur 12-14 mm long, curved at the distal end, purple (fig. 14-17). Etymology: The epithet “viridis” means “green” and refers to the green color of the stem and the bract-like leaves. Habitat: Two-layer oak forest with predomination of Quercus pubescens Willd. and three-layer pine forests with predomination of Pinus nigra J.F. Arnold subsp. pallasiana (Lamb.) Holmboe with Q. pubescens in the second layer. Altitude: The vertical distribution is 300 to 500 metres above sea level. Phenology: End of May until beginning of June, after L. abortivum var. abortivum. Distribution: Crimean Mountains (south slope of the main ridge between Alupka and Alushta). Neotinea ustulata (L.) R.M. Bateman, Pridgeon & M.W. Chase subsp. ustulata could be regarded as a neophyte. It was discovered only at the end of 20th

414 Journal Europäischer Orchideen 46 (2): 2014. century by ALEKSEEV (1993: 101-102), in spite of extensive botanical explorations carried out in suitable habitats a long time before. Until now there is only one known locality where this species grows in the Crimea (fig. 18-19). Forest reclamation works were carried out at this locality after WW II. These works provided possibly invasion of this species to the Crimea from the north of Ukraine. Ophrys mammosa Desf. is only represented by O. mammosa Desf. subsp. taurica (Aggeenko) Soó (fig. 22-25). The taxonomic isolation of this subspecies is still problematic. Ophrys oestrifera M. Bieb. requires further taxonomic investigation. One of its heterotypic synonyms, Ophrys cornuta Steven is probably a separate species. In this case, the Crimean form of O. oestrifera could be a separate, possibly endemic, taxon (cf. DELFORGE 2006: 495). Both taxa differ in flower size and flowering times: O. cornuta has slightly smaller flowers and flowers later than O. oestrifera. We did not find any differences in floral morphology between different populations of O. oestrifera s.l. growing in the Crimea. However, plants in some Crimean populations flower about 1 to 1.5 months earlier (April - early May, fig. 20) than in others (late May - early July, fig. 21). Orchis mascula (L.) L. is represented by the typical variety (fig. 26-27) and a variety known under the name Orchis mascula (L.) L. subsp. wanjkovii (E. Wulff) Soó (fig. 28-29). Orchis mascula (L.) L. var. wanjkovii (E. Wulff) Fateryga & Kreutz stat. nov. Basionym: Orchis wanjkovii E. Wulff, Fl. Taurica 1 (3): 97 (1930). Comments: Plants of both morphotypes (“mascula” and “wanjkovii”) grow often together within the same habitats. Therefore O. wanjkovii can not be regarded a separate subspecies from O. mascula. However as “wanjkovii” is quite different from the typical O. mascula this taxon should be regarded as a variety. Orchis militaris L. is only represented by O. militaris L. subsp. stevenii (Rchb. f.) B. Baumann & al. (fig. 30-33). The typical O. militaris is distributed further north and is absent in the Crimea. Platanthera bifolia (L.) Rich. requires further taxonomic investigation in the Crimea. There are various problematic forms (fig. 34-35); some of which show characteristics of P. bifolia (L.) Rich. var. latiflora (Drejer) Kreutz. The typical P. bifolia is very rare or absent (see also HAHN 2012: 58-59). Other forms can be probably attributed to P. ×hybrida Brügger (= P. bifolia (L.) Rich. × P. chlorantha (Custer) Rchb.) (fig. 36-37). See additional comments on P. ×hybrida further below.

Journal Europäischer Orchideen 46 (2): 2014. 415 Addenda

Epipactis helleborine (L.) Crantz subsp. levantina Kreutz, Óvári & Shifman (HAHN 2012: 37). Epipactis helleborine (L.) Crantz subsp. orbicularis (K. Richt.) E. Klein (KREUTZ & FATERYGA 2012: 715). Epipactis krymmontana Kreutz, Fateryga & Efimov (FATERYGA & al. 2014: 24). Epipactis muelleri Godfery (FATERYGA & al. 2013: 652-654). Epipactis persica (Soó) Nannf. subsp. taurica (Fateryga & Kreutz) Fateryga & Kreutz (FATERYGA & KREUTZ 2012: 201). See also “Notae” above. Epipactis turcica Kreutz (KREUTZ & FATERYGA 2012: 713-714).

Excludendae

Anacamptis fragrans (Pollini) R.M. Bateman, Pridgeon & M.W. Chase = A. coriophora (L.) R.M. Bateman, Pridgeon & M.W. Chase subsp. fragrans (Pollini) R.M. Bateman, Pridgeon & M.W. Chase is a misidentification of A. coriophora subsp. coriophora. This subspecies is quite variable in floral morphology and its different forms were obviously incorrectly identified for two different taxa, A. coriophora and A. fragrans. Anacamptis picta (Loisel.) R.M. Bateman, Pridgeon & M.W. Chase = A. morio (L.) R.M. Bateman, Pridgeon & M.W. Chase subsp. picta (Loisel.) Jacquet & Scappat. is a misidentification of A. morio subsp. caucasica. See also “Notae” for A. morio. Anacamptis laxiflora (Lam.) R.M. Bateman, Pridgeon & M.W. Chase is a misidentification of A. palustris subsp. elegans. See also “Notae” for A. palustris. (Rchb.) P.F. Hunt & Summerh. reported from the Crimea is almost certainly a misidentification of D. incarnata. In particular, most probably the small form of D. incarnata with broad leaves found by Ch. von Steven near Alushta (WULFF 1930: 93) was that previously identified as D. majalis. We did not find the population of the plants referred in Wulff’s paper and the herbarium specimens collected by Ch. von Steven (LE) could not be identified without doubt neither as D. incarnata nor as D. majalis. Thus, there remains a small possibility that D. majalis is present in the Crimea. However, as this species is distributed mainly much further north-westerly, and

416 Journal Europäischer Orchideen 46 (2): 2014. as we did not find it during many intensive searches in the Crimea, in our view D. majalis should be regarded as absent in the Crimea. Epipactis atrorubens (Hoffm.) Besser does not grow in the Crimea. The reports of this species were almost certainly misidentifications of E. krymmontana, E. helleborine s.l. and/or E. microphylla. Epipactis atrorubens is distributed much further north (see also EFIMOV 2008: 73). Epipactis condensata Boiss. ex D.P. Young was listed by EFIMOV (2008: 72- 76) for several localities in the Crimea. However, most of his reports did not belong to typical E. condensata; instead, they refer to the new described species, E. krymmontana (FATERYGA & al. 2014: 24). Only several plants from one locality in Karadag Mountains could be typical E. condensata; however they can be also assigned to E. turcica. Further investigation is therefore required at this site in the Karadag Mountains. Another specimen from Mt. Lapata (fig. 38) which was previously identified as E. condensata by us (FATERYGA & al. 2014: 27) was re-studied in flower (fig. 39) and then re- identified as E. krymmontana. The previous incorrect identification was due to unusual habit of the plant different from the typical one for E. krymmontana (fig. 40-41). Epipactis helleborine (L.) Crantz. is represented by E. helleborine subsp. levantina and E. helleborine subsp. orbicularis. We did not find any locality with typical E. helleborine (L.) Crantz subsp. helleborine in the Crimea. Most of the Crimean populations belong to E. helleborine subsp. levantina (fig. 42- 45), while E. helleborine subsp. orbicularis is known only from one locality near Yalta (KREUTZ & FATERYGA 2012: 715). Epipactis purpurata Smith is a misidentification of E. krymmontana (FATERYGA & al. 2014: 24). Orchis signifera Vest = Orchis mascula (L.) L. subsp. speciosa (Mutel) Hegi refers to typical Orchis mascula (L.) L. subsp. mascula. We checked the habitats of this taxon, referring to the dried specimens on the herbarium sheets that were identified as O. signifera. On these sites we only found very polymorphic specimens of typical O. mascula subsp. mascula. Spiranthes spiralis (L.) Chevall. is known only by literature. PRIVALOVA & PROKUDIN (1959: 79-80) refer to a single herbarium specimen collected “in the Crimean Mountains”. They emphasized that the specimen was identified without any doubt. However, they also suggested that confusion with labels could also have been taken place. Because of this and the fact that S. spiralis has never been confirmed for the Crimea by other botanists, and because we also were not able to find it during our field observations, we have excluded this species for the Crimea.

Journal Europäischer Orchideen 46 (2): 2014. 417 Hybrids

Anacamptis coriophora (L.) R.M. Bateman, Pridgeon & M.W. Chase × A. morio (L.) R.M. Bateman, Pridgeon & M.W. Chase subsp. caucasica (K. Koch) H. Kretzschmar, Eccarius & H. Dietr. Listed by WULFF (1930: 108) as Orchis morio L. × O. fragrans Pollini. We did not observe this hybrid. Anacamptis ×timbali (Velen.) H. Kretzschmar, Eccarius & H. Dietr. nothosubsp. reinhardii (Ugr. ex E.G. Camus) H. Kretzschmar, Eccarius & H. Dietr. (Anacamptis coriophora (L.) R.M. Bateman, Pridgeon & M.W. Chase × A. palustris (Jacq.) R.M. Bateman, Pridgeon & M.W. Chase subsp. elegans (Heuff.) R.M. Bateman, Pridgeon & M.W. Chase). Listed by WULFF (1930: 111) as Orchis laxiflora Lam. × O. fragrans Pollini. We did not observe this hybrid. Anacamptis ×simorrensis (E.G. Camus) H. Kretzschmar, Eccarius & H. Dietr. nothosubsp. ticinensis (Gsell) Fateryga & Kreutz stat. nov. (Anacamptis coriophora (L.) R.M. Bateman, Pridgeon & M.W. Chase subsp. coriophora × A. pyramidalis (L.) Rich.) Basionym: ×Anacamptorchis ticinensis Gsell, Boll. Soc. Ticinese Sci. Nat. 43: 74 (1948) (Anacamptis pyramidalis (L.) Rich. × Orchis coriophora L.). Comments: The previously proposed name ×Anacamptorchis simorrensis E.G. Camus nothovar. ticinensis (Gsell) Del Prete & M. Tomaselli is not applied to the hybrid of A. coriophora subsp. coriophora and A. pyramidalis since A. coriophora and A. fragrans (one of the parent taxon for A. ×simorrensis) are regarding as two subspecies (not varieties) of the same species A. coriophora. This hybrid is new for the flora of the Crimea, it was found for the first time by I.S. Turbanov in Baydary Valley (fig. 46). ×Dactylocamptis uechtritziana (Hausskn.) B. Bock ex M. Peregrym & Kuzemko nothosubsp. magyarii (Soó) Fateryga & Kreutz comb. et stat. nov. (Anacamptis palustris (Jacq.) R.M. Bateman, Pridgeon & M.W. Chase subsp. elegans (Heuff.) R.M. Bateman, Pridgeon & M.W. Chase × Dactylorhiza incarnata (L.) Soó) Basionym: Orchis ×magyarii Soó, Bot. Közlem. 1934. 31: 227 (1935) (Orchis elegans Heuff. × Orchis incarnata L.). Comments: PEREGRYM & KUZEMKO (2010: 655-662) published the combination ×D. uechtritziana for the hybrid of A. palustris and D. incarnata, which had been reported from Ukraine. ×D. uechtritziana nothosubsp. magyarii is the hybrid with A. palustris subsp. elegans as the first parent species. This hybrid is new for the flora of the Crimea. It was found for the first time by I.S. Turbanov in Baydary Valley (fig. 47).

418 Journal Europäischer Orchideen 46 (2): 2014. Neotinea ×dietrichiana (Bogenh.) H. Kretzschmar, Eccarius & H. Dietr. (Neotinea tridentata (Scop.) R.M. Bateman, Pridgeon & M.W. Chase × N. ustulata (L.) R.M. Bateman, Pridgeon & M.W. Chase). One specimen of this hybrid (fig. 48) was found by us on Mt. Tyrke Yayla. Orchis ×haussknechtii M. Schulze (Orchis mascula (L.) L. × O. pallens L.). Listed by WULFF (1930: 110). We did not observe this hybrid. Orchis mascula (L.) L. × O. pallens L. × O. provincialis Balb. ex Lam. & DC. Listed by WULFF (1930: 110). We did not observe this possible triple hybrid. Orchis ×penzigiana A. Camus (Orchis mascula (L.) L. × O. provincialis Balb. ex Lam. & DC.). Listed by WULFF (1930: 110). We observed this hybrid (fig. 49) near Simeiz in vicinity of Yalta. Orchis ×beyrichii Kern. nothosubsp. mackaensis (Kreutz) Fateryga & Kreutz comb. et stat. nov. (Orchis militaris L. subsp. stevenii (Rchb. f.) B. Baumann & al. × O. simia Lam.) Basionym: Orchis ×mackaensis Kreutz, Orchideeën 51 (3): 72 (1989) (Orchis simia Lam. × Orchis stevenii Rchb. f.). Comments: As O. stevenii is regarded as a subspecies of O. militaris, O. ×mackaensis should be reduced to a nothosubspecies of O. ×beyrichii (O. militaris × O. simia). This hybrid was listed for the Crimean flora by WULFF (1930: 109) as O. militaris L. × O. simia Lam. and by HAHN (2012: 54- 55). We found it (fig. 50-51) in the vicinity of Perevalnoye village. Orchis militaris L. subsp. stevenii (Rchb. f.) B. Baumann & al. × O. purpurea Huds. Listed by WULFF (1930: 110) as O. militaris L. × O. purpurea Huds. We did not observe this hybrid. Orchis ×plessidiaca Renz (Orchis pallens L. × O. provincialis Balb. ex Lam. & DC.). Listed by WULFF (1930: 110). We did not observe this hybrid. Orchis ×wulffiana Soó (Orchis punctulata Steven ex Lindl. × O. purpurea Huds.). Listed by WULFF (1930: 108), BENGUS & BENGUS (2011: 65) and HAHN (2012: 56-57). This is a very usual hybrid (fig. 52) which we observed at several localities. Orchis punctulata Steven ex Lindl. × O. purpurea Huds. × O. simia Lam. Listed by BENGUS & BENGUS (2011: 65) as a probable hybrid with O. militaris as one of the parent species. We did not observe this hybrid. Orchis ×calliantha Renz & Taubenheim (Orchis punctulata Steven ex Lindl. × O. simia Lam.). Listed by BENGUS & BENGUS (2011: 65). We did not observe this hybrid.

Journal Europäischer Orchideen 46 (2): 2014. 419 Orchis ×angusticruris Franch. (Orchis purpurea Huds. × O. simia Lam.). Listed by WULFF (1930: 109) and BENGUS & BENGUS (2011: 65). This is a very usual hybrid (fig. 53) which we observed at several localities. Platanthera ×hybrida Brügger (Platanthera bifolia (L.) Rich. × P. chlorantha (Custer) Rchb.). Listed by HAHN (2012: 58-59) as Platanthera sp. This is a very usual hybrid which we observed at numerous localities (fig. 36-37). Interestingly, both this hybrid and P. chlorantha can be found in many places in the Crimea while the typical P. bifolia is very rare. Furthermore, P. bifolia is in the Crimea represented by many forms, one of which shows characteristics similar to P. bifolia (L.) Rich. var. latiflora (Drejer) Kreutz. At other places in the Crimea P. ×hybrida completely replaces P. bifolia and thus it could be regarded as separate hybridogenous species, P. hybrida. As observed by HAHN (l.c.), this situation requires further investigation.

Acknowledgements

We thank Sergey Svirin, Pavel Yevseyenkov, Ilya Turbanov (Sevastopol, Crimea), and Prof. Sergey Ivanov (Vernadskiy Taurida National University, Simferopol, Crimea) for providing their photos and permission to publish them. We thank Leslie Lewis (Chepstow, UK) for critically reading through the text. We also thank Dr. Valentina Fateryga (Karadag Nature Reserve, Feodosiya, Crimea) as well as Sergey Svirin and Prof. Sergey Ivanov for their help during the field works.

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Authors’ addresses

Dr. Alexander V. Fateryga Karel (C.A.J.) Kreutz Karadag Nature Reserve, Naturalis Biodiversity Center Nauki str. 24, Kurortnoye, Biosystematics group Feodosiya, Republic of Crimea Wageningen University 298188, Russia Generaal Foulkesweg 37 [email protected] Wageningen, NL-6703 BL The Netherlands [email protected]

For the following figures 1-12, 14-53 their numbering follows the order from top left to bottom right on every page.

422 Journal Europäischer Orchideen 46 (2): 2014. Fig. 1-4: Species of Anacamptis. Fig. 1: A. morio subsp. caucasica, Mt. Dolgorukovskaya Yayla, meadow, 3.6.2012 (phot. AVF); Fig. 2: A. morio subsp. caucasica, Lisya Bay, slope of badland, 9.5.2011 (phot. AVF); Fig. 3: A. palustris subsp. elegans, Baydary Valley, vicinity of Tylovoye village, wet meadow, 11.6.2010 (phot. S.A. Svirin); Fig. 4: A. pyramidalis var. orientalis, vicinity of Sevastopol, Balaklava, light forest, 27.5.2011 (phot. CAJK).

Journal Europäischer Orchideen 46 (2): 2014. 423 Fig. 5-8: Dactylorhiza romana. Fig. 5-6: Vicinity of Yalta, pine forest, 29.4.2011 (phot. AVF); Fig. 7-8: Mt. Demerdji Yayla, oak forest, 3.5.2011 (phot. AVF).

424 Journal Europäischer Orchideen 46 (2): 2014. Fig. 9-12: Epipactis persica subsp. taurica and varieties of Limodorum abortivum. Fig. 9-10: E. persica subsp. taurica, type locality, vicinity of Yalta, Mt. Lapata, pine forest, 18.7.2014 (phot. AVF); Fig. 11: L. abortivum var. abortivum, Laspi Valley, 24.5.2009 (phot. S.P. Ivanov); Fig. 12: L. abortivum var. rubrum, vicinity of Alushta, near Vinogradnoye village, pine forest, 29.5.2014 (phot. AVF).

Journal Europäischer Orchideen 46 (2): 2014. 425 Fig. 13: Holotype specimen of Limodorum abortivum var. viridis [Vicinity of Alushta, near Vinogradnoye village, Mt. Uraga, oak forest], 29.5.2014, leg. AVF, CSAU.

426 Journal Europäischer Orchideen 46 (2): 2014. Fig. 14-17: Limodorum abortivum var. viridis. Fig. 14-15: Type locality, vicinity of Alushta, near Vinogradnoye village, oak forest, 29.5.2014 (phot. AVF); Fig. 16: Vicinity of Yalta, Miskhor, pine forest, 28.5.2012 (phot. AVF); Fig. 17: Mt. Ayudag, oak forest, 27.5.2013 (phot. P.Ye. Yevseyenkov).

Journal Europäischer Orchideen 46 (2): 2014. 427 Fig. 18-21: Neotinea ustulata and Ophrys oestrifera. Fig. 18: N. ustulata, Mt. Tyrke Yayla, meadow slope, 3.6.2012 (phot. AVF); Fig. 19: N. ustulata, ibid., 11.6.2012 (phot. CAJK); Fig. 20: O. oestrifera, vicinity of Yalta, “Mys Martyan” Nat. Res., juniper forest, 26.4.2011 (phot. AVF); Fig. 21: O. oestrifera, Simferopol district, vicinity of Perevalnoye village, Ayan Natural Landmark, shrubby meadow, 24.5.2011 (phot. AVF).

428 Journal Europäischer Orchideen 46 (2): 2014. Fig. 22-25: Ophrys mammosa subsp. taurica, Bakhchisaray district, Mt. Mangup, light forest, 23.5.2011 (phot. CAJK).

Journal Europäischer Orchideen 46 (2): 2014. 429 Fig. 26-29: Varieties of Orchis mascula. Fig. 26-27: O. mascula var. mascula, Mt. Tyrke Yayla, meadow, 17.5.2013 (phot. S.A. Svirin); Fig. 28-29: O. mascula var. wanjkovii, Mt. Baydarskaya Yayla, oak forest, 23.4.2010 (phot. S.A. Svirin).

430 Journal Europäischer Orchideen 46 (2): 2014. Fig. 30-33: Orchis militaris subsp. stevenii. Fig. 30-31: Mt. Tyrke Yayla, meadow slope, 3.6.2012 (phot. AVF); Fig. 32-33: Simferopol district, Privolnoye, forest glade, 24.5.2011 (phot. CAJK).

Journal Europäischer Orchideen 46 (2): 2014. 431 Fig. 34-37: Platanthera bifolia and P. ×hybrida. Fig. 34-35: P. bifolia, Simferopol district, vicinity of Perevalnoye village, Ayan Natural Landmark, shrubby meadow, 10.6.2012 (phot. CAJK); Fig. 36: P. ×hybrida, ibid., 10.6.2012 (phot. CAJK); Fig. 37: P. ×hybrida, Simferopol district, vicinity of Perevalnoye village, Kurlukbash Natural Landmark, oak forest, 17.6.2011 (phot. P.Ye. Yevseyenkov).

432 Journal Europäischer Orchideen 46 (2): 2014. Fig. 38-41: Epipactis krymmontana. Fig. 38: Vicinity of Yalta, Mt. Lapata, beech forest and yayla, 27.7.2013 (phot. AVF); Fig. 39: Ibid., 18.7.2014 (phot. AVF); Fig. 40: Type locality, Belogorsk district, Zemlyanichnoye, beech forest, 13.6.2012 (phot. AVF); Fig. 41: Ibid., 20.6.2012 (phot. CAJK).

Journal Europäischer Orchideen 46 (2): 2014. 433 Fig. 42-45: Epipactis helleborine subsp. levantina. Fig. 42: Vicinity of Yalta, Mt. Lapata, beech forest, 27.7.2013 (phot. AVF); Fig. 43: Ibid., 18.7.2014 (phot. AVF); Fig. 44-45: Mt. Chatyrdag, beech forest, 15.7.2014 (phot. AVF).

434 Journal Europäischer Orchideen 46 (2): 2014. Fig. 46-49: Hybrids. Fig. 46: Anacampis ×simorrensis nothosubsp. ticinensis, Baydary Valley, vicinity of Kisilovoye village, 3.6.2010 (phot. I.S. Turbanov); Fig. 47: ×Dactylocamptis uechtritziana nothosubsp. magyarii, Baydary Valley, vicinity of Tylovoye vilage, 10.6.2010 (phot. I.S. Turbanov); Fig. 48: Neotinea ×dietrichiana, Mt. Tyrke Yayla, 11.6.2012 (phot. AVF); Fig. 49: Orchis ×penzigiana, vicinity of Yalta, near Simeiz village, 30.4.2013 (phot. AVF).

Journal Europäischer Orchideen 46 (2): 2014. 435 Fig. 50-53: Hybrids. Fig. 50-51: Orchis ×beyrichii nothosubsp. mackaensis, Simferopol district, vicinity of Perevalnoye village, Ayan Natural Landmark, 19.5.2011 (phot. AVF); Fig. 52: Orchis ×wulffiana, Cape Aya, 18.4.2014 (phot. S.A. Svirin); Fig. 53: Orchis ×angusticruris, Mt. Dolgorukovskaya Yayla, 3.5.2012 (phot. S.A. Svirin).

436 Journal Europäischer Orchideen 46 (2): 2014.