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Turczaninowia 23, 4: 65-71 (2020) DOI: 10.14258/turczaninowia.23.4.6 http://turczaninowia.asu.ru
ISSN 1560-7259 (print edition)
TURCZANINOWIA
ISSN 1560-7267 (online edition)
УДК 582.594.2(575.4)
On taxonomic status of two species of orchids (Orchidaceae)
from Turkmenistan
A. V. Fateryga1*, A. V. Pavlenko2, V. V. Fateryga1
1T. I. Vyazemsky Karadag Scientific Station - Nature Reserve of RAS - Branch of A. O. Kovalevsky Institute of Biology of the Southern Seas of RAS, Nauki str., 24, Kurortnoye, Feodosiya, Republic of Crimea, 298188, Russian Federation.
E-mails:fater_84@list.ru*, valentina_vt@mail.ru
2 Serdar Branch of the Center for Prevention of Especially Dangerous Infectious Diseases, State Sanitary and Epidemiological Service, Ministry of Health and Medical Industry of Turkmenistan, O. Akmamedov str., 44, Serdar, 745150, Turkmenistan.
E-mail: alexpavlenko1974@gmail.com
Keywords. Epipactis, Middle Asia, new records, Ophrys, synonymy, taxonomy.
Summary. The orchid genera Epipactis Zinn and Ophrys L. are well-known by their complicated taxonomy and extensive debates over species richness within them. These genera are represented in Turkmenistan by two species each. Two of them, namely E. turcomanica K. P. Popov et Neshat. and O. kopetdagensis K. P. Popov et Neshat., were hitherto accepted as species endemic to Turkmenistan. In the present paper, these taxa are synonymized with broadly distributed E. persica (Soo) Hausskn. ex Nannf. and O. oestrifera M. Bieb., respectively. Thus, the genus Epipactis is represented in Turkmenistan by E. persica and E. veratrifolia Boiss. et Hohen., and the genus Ophrys is represented by O. mammosa Desf. and O. oestrifera. There are no species of orchids endemic to Turkmenistan.
1 Карадагская научная станция им. Т. И. Вяземского - природный заповедник РАН - филиал Института биологии южных морей им. А. О. Ковалевского РАН, ул. Науки, 24, пгт Курортное, г. Феодосия, Республика Крым, 298188, Россия
2 Сердарский отдел Центра профилактики особо опасных инфекций государственной санитарно-эпидемиологической службы Министерства здравоохранения и медицинской промышленности Туркменистана, ул. О. Акмамедова, 44,
г. Сердар, 745150, Туркменистан
Ключевые слова: новые находки, синонимия, Средняя Азия, таксономия, Epipactis, Ophrys.
Аннотация. Орхидеи родов Epipactis Zinn и Ophrys L. хорошо известны благодаря запутанной таксономии и бурным дебатам в отношении их видового богатства. Эти роды представлены в Туркменистане двумя видами каждый. Epipactis turcomanica K. P. Popov et Neshat. и Ophrys kopetdagensis K. P. Popov et Neshat. до настоящего времени считались эндемиками Туркменистана. В данной работе эти таксоны сведены в синонимы к широко распространенным E. persica (Soo) Hausskn. ex Nannf. и O. oestrifera M. Bieb., соответственно. Таким образом, род Epipactis представлен в Туркменистане E. persica и E. veratrifolia Boiss. et Hohen., а род Ophrys -O. mammosa Desf. и O. oestrifera. Эндемики Туркменистана среди орхидей отсутствуют.
Поступило в редакцию 01.09.2020 Submitted 01.09.2020
Принято к публикации 15.09.2020 Accepted 15.09.2020
*Corresponding author
О таксономическом статусе двух видов орхидных (Orchidaceae) из Туркменистана
А. В. Фатерыга1, А. В. Павленко2, В. В. Фатерыга1
Introduction
The orchid family (Orchidaceae) is the second largest plant family in the world after Asteraceae (Christenhusz, Byng, 2016). Most species of orchids are threatened plants due to their complex life history strategies, threats from overcollection and habitat loss, as well as climate change (Pillon, Chase, 2006; Fay, Chase, 2009; Swarts, Dixon, 2009; Efimov, 2012; Fay, 2018). Some groups of orchids are still poorly studied taxonomically even in the temperate climatic zone. Particularly, this is true for the genera Epipactis Zinn and Ophrys L., which are currently being studied phylogenetically (Breitkopf et al., 2015; Bateman et al., 2018; Zhou, Jin, 2018; Sramko et al., 2019).
Many species of Epipactis in the section Epipactis s. str. are described and then synonymized with other already known species; particularly, this is true for some facultatively cross-pollinating taxa. For example, E. kartliana Kreutz et Van Domm. recently described from Georgia (Kreutz, 2019) looks morphologically almost like E. kuenkeleana (Akhalk., H. Baumann, R. Lorenz et Mosul.) P. Delforge, which, in turn, has been already synonymized with E. condensata Boiss. ex D. P. Young (Fateryga, Fateryga, 2018). R. M. Bateman (2020) states that there are just 12 "bona fide" species of Epipactis in the section Epipactis s. str. among the 65 putative species recognized by P. Delforge (2016), although only 27 of them have been studied molecularly (Sramko et al., 2019). It is evidently that further studies are required, since several taxa, which are well-recognized morphologically (e. g., E. condensata or E. tangutica Schltr.), were not included to their analysis.
Modern estimation of the species richness of the genus Ophrys varies from 9 macrospecies based on the results of molecular analysis (Bateman et al., 2018) to more than 350 microspecies based on minute morphological differences (Delforge, 2016; Baguette et al., 2019). Such a proliferation of species is merely the sort of irrational splitting. Although much more than 9 species can be recognized on the base of traditional morphology, the number of taxa well-distinguishable by this way is much less than 350. Some intermediate points of view between those of "lumpers" and "splitters" are also possible (Vela et al., 2015; Fateryga et al., 2018a).
The genera Epipactis and Ophrys are represented in Turkmenistan by two species each (Nikitin, Geldikhanov, 1988). Of them, E. veratrifolia Boiss. et Hohen. belongs to the section Arthrochilium
Irmisch; this species is not problematic. Ophrys transhyrcana Czerniak. accepted by some authors as a full species (Vlasenko, 2011; Delforge, 2016) is currently treated as a synonym of O. sphegodes subsp. mammosa (Desf.) Soo ex E. Nelson (World Flora Online, 2020), which is, however, often accepted as a full species, i. e., O. mammosa Desf. (Fateryga et al., 2018a; Efimov, 2020). Two remaining taxa were described by K. P. Popov and G. Yu. Neshataeva (1982): E. turcomanica K. P. Popov et Neshat. and O. kopetdagensis K. P. Popov et Neshat.; both are currently accepted as full species endemic to Turkmenistan (Govaerts et al., 2005-2020; World Flora Online, 2020).
Epipactis turcomanica described from Ajdere Gorge in Southwestern Kopet Dag Mountains was hitherto known just by the protologue. It was, however, included to the Red Data Book of Turkmenistan (Vlasenko, 2011) as a critically endangered and narrow endemic species. The species was described on the base of plants in fruit (Popov, Neshataeva, 1982). The authors of the protologue compared their new species with E. atrorubens (Hoffm.) Besser and E. tangutica: both had strongly pubescent rachis, while E. turcomanica had rather glabrous one (Popov, Neshataeva, 1982). At the same time, they did not denote any differences of their new species from taxa in the E. phyllanthes G. E. Sm. species group, which contained species with either glabrous or subglabrous rachis (Delforge, 2016; Fateryga, Fateryga, 2018). A similar situation is true for O. kopetdagensis as well. It was described from Pordere Gorge, also in Southwestern Kopet Dag Mountains. The authors (Popov, Neshataeva, 1982) compared it with O. transhyrcana from the O. mammosa species group (merged into O. sphegodes Mill. group in Bateman et al., 2018), while their new species actually belonged to the O. oestrifera M. Bieb. species group (merged into Ophrys fuciflora (F. W. Schmidt) Moench group in Bateman et al., 2018). For a long time O. kopetdagensis was known just by the protologue until its rediscovery in the type locality in 2015 (Pavlenko et al., 2015).
The purpose of the present study is to clarify the taxonomic statuses of both E. turcomanica and O. kopetdagensis according to the species concept proposed in the previous papers (Fateryga, Fateryga, 2018; Fateryga et al., 2018a). It was possible due to a rediscovery of E. turcomanica by the second author and Ch. Tagiev in Pordere Gorge in 2019 (Fig. 1A-B). Living plants were studied in the field and voucher specimens were
collected and preserved in the research herbarium of A. V. Pavlenko (Serdar, Turkmenistan). Specimens of O. kopetdagensis previously recorded there (Pavlenko et al., 2015) were also studied in the field and on the base of the preserved material. Type specimens of E. turcomanica and O. kopetdagensis were studied in LE.
As the result, E. turcomanica and O. kopetdagensis were found being conspecific with E. persica (Soo) Hausskn. ex Nannf. and O. oestrifera, respectively. Nomenclature of these two species and proposed new synonymies are present below, along with taxonomic notes and other relevant data.
Species account
Epipactis persica (Soo) Hausskn. ex Nannf. 1946, Bot. Not. 1946(1): 21. = Helleborine persica Soo, 1927, Repert. Spec. Nov. Regni Veg. 24: 36. = Epipactis microphylla subsp. persica (Soo) Hautz. 1976, Verh. Zool.-Bot. Ges. Wien, 115: 42. = Epipactis helleborine subsp. persica (Soo) H. Sund. 1980, Europ. Medit. Orchid. ed. 3: 41, nom. inval.
Lectotypus (Hautzinger, 1976, Verh. Zool.-Bot. Ges. Wien, 115: 42): Iran: "In dit. urb. Sultanabad. 1898. Th. Strauss s. n." (JE: JE00007753!).
= Epipactis turcomanica K. P. Popov et Neshat. 1982, Izv. Akad. Nauk Turkmensk. S. S. R., Ser. Biol. Nauk, 1982, 4: 16.
Holotypus: Turkmenistan: "Ю. Туркмения, юго-западный Копетдаг, ущелье Ай-Дере, на берегу в среднем течении р. Айдеринка. Un., чрезвычайно редко! [Southern Turkmenistan, Southwestern Kopet Dag, Ajdere Gorge, at a bank in the middle reaches of Ajderinka River. Un., very rarely!]. 19 VII 1980. K. P. Popov s. n." (LE: LE00050987!) (Fig. 2A).
= Epipactis helleborine subsp. transcaucasica A. P. Khokhr. 1991, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 96, 4: 107.
Holotypus: Georgia: "Adzharia, distr. Keden-sis, opp. pagus Zhemna, pineto-quercetum sic-cum, schistosum. 5 VI 1990. A. P. Khokhrjakov, P. A. Khokhrjakov, S. Czitanava s. n." (MW: MW0591780!).
= Epipactis taurica Fateryga et Kreutz, 2012, J. Eur. Orch. 44(1): 201. = Epipactis persica subsp. taurica (Fateryga et Kreutz) Fateryga et Kreutz, 2014, J. Eur. Orch. 46(2): 413.
Holotypus: Russia: "Крым, окр. Ялты, Ялтинский заповедник, склоны горы Лопата, сосняк коротконожковый [Crimea, vicinity of
Yalta, Yalta Reserve, slopes of Mt Lopata, pine forest with Brachypodium]. 9 VII 2011. A. V. Fateryga s. n." (CSAU!).
Specimens examined: Turkmenistan: "Ю.-З. Копетдаг, ущ. Пордере, в тени Salix [Southwestern Kopet Dag, Pordere Gorge, 38°20'N, 57°03'E, ca. 1300 m a. s. l., under a Salix shade]. 28 VI 2019. Ch. Tagiev s. n." (herbarium of A. V. Pavlenko: PAV00002) (Fig. 2B).
Distribution: Balkans, Crimea, Caucasus, Western Asia (except south of the Arabian Peninsula), Middle Asia (Turkmenistan, Tajikistan), and Southern Asia (Afghanistan, Pakistan).
Notes: The holotype of E. turcomanica has been already labeled as E. persica by D. Ruckbrodt and U. Ruckbrodt (Fig. 2A). Indeed, it perfectly fits the diagnosis of E. persica: the plant is rather small, with few leaves and flowers, and the rachis of inflorescence is glabrous. Although these characters are diagnostic for the whole E. phyllanthes species group, E. persica is the only species in this group distributed so far east (Delforge, 2016). Some doubts in the proposed synonymy, however, remained until the plants were rediscovered and studied in the field (Fig. 1A-B). Study of the living plants in flower revealed that they had epichile rather equal in length and width, and distinctly present but inefficient vis-cidim, i. e., the characters of E. persica (Fateryga et al., 2018b). One more character is a short but distinctly present pedicel.
It was recently stated that the E. phyllanthes species group consisted of a single "bona fide" species (Sramko et al., 2019; Bateman, 2020), although just three microspecies in this group were studied molecularly: E. exilis P. Delforge, E. persica, and E. phyllanthes s. str. (Sramko et al., 2019). Indeed, the differences between E. exilis and E. persica are not clear (cf. Fateryga et al., 2018b) and this is possibly true for other taxa as well. On the other hand, there are some species, which are well-recognized morphologically, e. g., E. euxina Fateryga, Popov-ich et Kreutz from the North Caucasus or the species from Crete known as E. cretica Kalop. et Robatsch, nom. inval. To better ascertain the phylogenetic relationships within the E. phyllanthes species group and to clarify the number of species in this group, such taxa should be also included in further molecular studies. Until this is done, E. persica can be also treated as a full species, not a synonym of E. phyl-lanthes (these two taxa are allopatric and, therefore, merit at least the subspecies rank).
Fig. 1. Epipactis persica (Soo) Hausskn. ex Nannf. (A-B) and Ophrys oestrifera M. Bieb. (C) from Pordere Gorge, Turkmenistan: A - flowering plant; B-C - part of an inflorescence. Photos by Ch. Tagiev (A-B) and A. V. Pavlenko (C).
Fig. 2. Holotype of Epipactis turcomanica K. P. Popov et Neshat. (LE: LE00050987) (A) and a modern gathering from Pordere Gorge, Turkmenistan (herbarium of A. V. Pavlenko: PAV00002) (B).
Ophrys oestrifera M. Bieb. 1808, Fl. Taur.-Cau- var. oestrifera (M. Bieb.) Rchb. f. in Rchb. 1851, cas. 2: 369. = Orchis oestrifera (M. Bieb.) M. Bieb. Icon. Fl. Germ. Helv. (H. G. L. Reichenbach), 1819, Fl. Taur.-Caucas. 3: 605. = Ophrys scolopax 13/14: 99. = Ophrys scolopax subsp. oestrifera
L+YS"*8
(M. Bieb.) Soo, 1970 (publ. 1971), Acta Bot. Acad. Sci. Hung. 16(3/4): 386.
Lectotypus (Averyanov, 1994, Bot. Zhurn. (St. Petersburg), 79, 10: 124): Russia: "Taur. mer. 1807. Ch. Steven s. n." (LE!).
= Ophrys bremifera Steven in M. Bieb. 1808, Fl. Taur.-Caucas. 2: 370; 1809, Mém. Soc. Imp. Naturalistes Moscou, 2: 174. = Ophrys oestrifera sub-sp. bremifera (Steven) K. Richt. 1890, Pl. Eur. 1: 264. = Ophrys oestrifera f. bremifera (Steven) Soo, 1928, Bot. Arch. 23: 32. = Ophrys scolopax subsp. bremifera (Steven) Biel, 1999, Ber. Arbeitskreis. Heimische Orchid. 16(1): 54.
Lectotypus (Averyanov, 1994, Bot. Zhurn. (St. Petersburg), 79, 10: 124): Georgia: "Ex Iberia. 1808. Ch. Steven s. n." (LE!).
= Ophrys cornuta Steven in M. Bieb. 1808, Fl. Taur.-Caucas. 2: 370; 1809, Mém. Soc. Imp. Naturalistes Moscou, 2: 175. = Ophrys bicornis var. cornuta (Steven) Nyman, 1882, Consp. Fl. Eur.: 698. = Ophrys oestrifera subsp. cornuta (Steven) K. Richt. 1890, Pl. Eur. 1: 264. = Ophrys arachnites var. cornuta (Steven) Fiori et Paol. in Fiori., Bég. et Paol. 1896, Fl. Italia [Fiori, Béguinot et Paoletti], 1: 236. = Ophrys scolopax subsp. cornuta (Steven) E. G. Camus in E. G. Camus, Bergon et A. Camus, 1908, Monogr. Orchid.: 270. = Ophrys holoserica subsp. cornuta (Steven) H. Sund. 1975, Taxon, 24(5/6): 625. = Ophrys fuciflora subsp. cornuta (Steven) H. Sund. 1980, Europ. Medit. Orchid. ed. 3: 39.
Lectotypus (Kreutz, 2019, Ber. Arbeitskreis. Heimische Orchid. 36(1): 156): Illustration in Steven, 1809, Mém. Soc. Imp. Naturalistes Moscou, 2: Tab. XI, Fig. 3 [based on a specimen collected in Georgia].
= Ophrys kopetdagensis K. P. Popov et Neshat. 1982, Izv. Akad. Nauk Turkmensk. S. S. R., Ser. Biol. Nauk, 1982, 4: 17.
Holotypus: Turkmenistan: "Юго-Западный Ко-петдаг, ущелье Пордере, на берегу ручья, 1200 м над ур. м. Редко [Southwestern Kopet Dag, Pordere Gorge, at a stream bank, 1200 m a. s. l. Rarely]. 25 V 1981. K. P. Popov s. n." (LE: LE01079804!).
= Ophrys oestrifera subsp. abchasica Kümpel, 1988, Ber. Arbeitskreis. Heimische Orchid. 5(1/2): 25. = Ophrys abchasica (Kümpel) P. Delforge in P. Delforge, Devillers-Tersch. et Devillers, 1991, Naturalistes Belges, 72(3): 100.
Holotypus: Russia: "Sotschi, Msymta-Tal. 17 V 1979. H. Kümpel s. n." (JE: JE00014373!).
Specimens examined: Turkmenistan: "Ю.-З. Копетдаг, ущ. Пордере (верховья) [Southwest-
ern Kopet Dag, Pordere Gorge (upper reaches), 38°15'N, 57°02'E, ca. 1200 m a. s. l.]. 1 V 2019. A. V. Pavlenko s. n." (herbarium of A. V. Pavlenko: PAV00021).
Distribution: Balkans, Crimea, Caucasus, Western Asia (except south of the Arabian Peninsula), and Middle Asia (Turkmenistan).
Notes: The studied material (Fig. 1C) well corresponds to specimens of O. oestrifera from the type locality (Crimea). No differences in flower morphology were revealed, even the shape of the appendage of the lip was tridentate like in plants from the Crimea and unlike in those from the environs of Sochi (Fateryga et al., 2018a). No differences of O. kopetdagensis from O. oestrifera were reported previously as well (Popov, Neshataeva, 1982; Pavlenko et al., 2015). Therefore, a new synonymy is proposed here. The species should be included to the next edition of the Red Data Book of Turkmenistan (Pavlenko et al., 2015) under the name O. oestrifera.
The taxonomic status of O. oestrifera was justified in one of the previous papers (Fateryga et al., 2018a). Those who accept species of the genus Ophrys in the broader sense may treat it as O. scolopax subsp. cornuta. The synonymy of O. oestrifera with O. apifera Huds. (Govaerts et al., 2005-2020; De-vey et al., 2008; World Flora Online, 2020) is incorrect and based on a mistake (cf. Bateman et al., 2018; Fateryga et al., 2018a).
Conclusions
The genus Epipactis is represented in Turkmenistan by E. persica and E. veratrifolia, the genus Ophrys is represented by O. mammosa and O. oestrifera. There are no species of orchids endemic to Turkmenistan.
Acknowledgments
Charyyar Tagiev (Sunt-Hasardag Nature Reserve, Magtymguly, Turkmenistan) supported the second author during his work in Pordere Gorge and provided some photos. Denis G. Melnikov (V. L. Komarov Botanical Institute of the Russian Academy of Sciences, Saint Petersburg, Russia) provided scans of the type specimens.
The work of A. V. Fateryga and V. V. Fateryga was a part of the State research project No. AAAA-A19-119012490044-3 of the A. O. Kovalevsky Institute of Biology of the Southern Seas of RAS.
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