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55Новости систематики высших растений 2019
Novitates Systematicae Plantarum Vascularium 50: 101-114 I i ISSN 0568-5443
Towards a taxonomic revision of the genus Trollius (Ranunculaceae) in the Asian part of Russia. I. Trollius chinensis: taxonomic and geographical reconsiderations
К таксономической ревизии рода Trollius (Ranunculaceae) в азиатской части России. I. Trollius chinensis: критический анализ таксономии и географии
M. M. Serebryanyi
Tsitsin Main Botanical Garden, Russian Academy of Sciences Herbarium
Botanicheskaya Str., 4, Moscow, 127276, Russia [email protected]
https://doi.org/10.31111/novitates/2019.50.101
М. М. Серебряный
Главный ботанический сад им. Н. В. Цицина РАН Гербарий
Ботаническая ул., 4, Москва, 127276, Россия [email protected]
Abstract. The present paper opens a series of publications by the author on taxonomy and geography of the genus Trollius L. in the Asian part of Russia. Trollius chinensis Bunge was chosen as a focus of the paper, since the current status of knowledge of taxonomy and geography of this species appeared to be inadequate. Results of the research prove a vast distribution range of T. chinensis in Russia within which there is a number of contact zones with other species of the genus, resulting in active interspecific hybridization. Detailed adjusted morphological description and list of synonyms are elaborated, T. vitalii Stepanov and T. chinensis subsp. macropetalus (Regel) Luferov are reduced to synonymy; combination T. macropetalus (Regel) F. Schmidt should be considered invalidly published and withdrawn from usage. Occurrence of T. chinensis in Mongolia (Harhiraa Mountain Area) is recorded for the first time.
Keywords: Trollius, T. chinensis, synonymy, distribution range, hybridization.
Аннотация. Настоящая работа открывает цикл публикаций автора по систематике и географии рода Trollius L. в азиатской части России. В качестве основного объекта этой статьи выбран Trollius chinensis Bunge — вид, современные представления о котором — от таксономических до географических — оказались не соответствующими действительности. Результаты исследований доказывают обширный ареал T. chinensis на территории России и наличие многочисленных зон контакта (и гибридизации) этого вида с другими купальницами. Уточнены диагноз и синонимика T. chinensis; T. vitalii Stepanov и T. chinensis subsp. macropetalus (Regel) Luferov отнесены к синонимам T. chinensis; показано, что комбинация T. macropetalus (Regel) F. Schmidt недействительно обнародована; впервые доказано нахождение T. chinensis в Монголии (горный массив Хархираа).
Ключевые слова: Trollius, T. chinensis, синонимика, ареал, гибридизация.
Trollius L. is a genus of ca. 30 (Kadota, 1985, 2016), 31 (Doroszewska, 1974) to 35 species (Luferov et al., 2018) of perennial herbs distributed entirely in the temperate/extratropical zones of the North Hemisphere, often (even mainly) in mountain areas — within forest-steppe, forest, subalpine, alpine — up to tundra/goltsy — altitudinal belts. On flat terrain Trollius species occur in wetlands (including wet forests and meadows), often near water streams and on flood plains.
Main reasons of uncertainty in the number of species within the genus Trollius are:
- still incomplete understanding of distribution ranges of Trollius species occurring in Asia;
— still unsatisfactory knowledge of variation range within groups of critical species;
Поступила в редакцию | Submitted: 03.09.2019
— unlimited capability of almost all the species to hybridize (also in natural plant communities);
- treating of T. lilacinus Bunge and sometimes T. chartosepalus Schipcz. (Khokhrjakov, 1977) within a separate genus Hegemone Bunge;
— absence of currently accepted monograph of the genus: the effort by Alina Doroszewska (1974) is obsolete and full of well-known discrepancies.
Thus, I must admit that V. N. Siplivinsky's eloquent phrase: "The genus Trollius belongs to those genera of our flora for which accumulation of material is far ahead of taxonomic development" (Siplivinsky, 1972: 163) — is still valid.
History of taxonomic study of the genus Trollius in Asian part of Russia dates back about a century. Primary accumulation of material was connected with big
Принята к публикации | Accepted: 02.12.2019
names of Russian botany: K. J. Maximowicz, V. L. Ko-marov, E. L. Regel, A. A. Bunge and others. The first monograph of the genus and treatment of Trollius for Flora of USSR published by N. V. Schipczinsky (1924, 1937) became obsolete already up to beginning of 1960s; the works by V. V. Reverdatto (1943) and L. I. Malyschev (1965) contributed a few new taxa but did not contain in-depth taxonomic considerations. At the same time the genus Trollius in Japan and Korea was studied by Japanese botanists T. Nakai, K. Miyabe and others. Their efforts were summarized by Miyabe (1943). Siplivinsky (1972) published a paper "Genus Trollius L. in Asia boreali et orientali" which became a long-term extremely important source for the Asian species of the genus. Two years later another monograph of Trollius was published by Doroszewska (1974). It contained an original intrageneric classification (the genus was subdivided into 7 new sections) which is still in use by taxonomists. However, this monograph has not become a fundamental basis for further research, since it contains a number of serious drawbacks: the author did not study type material of almost all the species (clearly confirmed that in the monograph), did not critically revise species in the Asian part of Russia and did not compose satisfactory identification key to the species. Until Trollius treatment for Flora Sibiriae (Friesen, 1993) — which cannot be considered a value-add to the taxonomic knowledge of the genus — only a few articles were published, particularly those containing descriptions of new species: T. bargusinensis Sipliv. (Siplivinsky, 1973) and T. aldanensis Volot. (Voloto-vsky, 1990). N. V. Stepanov (1994) described a new species from West Sayan — T. vitalii Stepanov (alongwith several infraspecific taxa), and another one — T. kolo^k Stepanov — from Kuznestk Alatau (Stepanov, 2018).
After a number of contributions (Kadota, 1985, 1987) Y. Kadota (2016) published a revision of the genus Trollius in Japan — a very important research, in which — besides a description of three new species (T. soyaensis Kadota, T. teshioensis Kadota and T. re-bunensis Kadota) and one new section (Yezoinsulicola Kadota) — substantiated a broad concept of T. riede-rianus Fisch. et C. A. Mey. and commented on usage of petals (nectaries) morphology as diagnostic (not just morphometric) characters. Unfortunately, in a revision of the genus Trollius in Far East of Russia (Luferov et al., 2018; nine species recognized for the region) the Kadota's paper was completely ignored. Another recent contribution on the genus Trollius in the flora of the Altai Mountain Country (Erst et al., 2018) comprises only three species — T. asiaticus L., T. altaicus C. A. Mey. and T. dschungaricus Regel. Despite the author's statement on a geographical delimitation of the
researched region (comprising a whole Altai Mountain Country, incl. Russian, Kazakh, Chinese and Mongolian territories) absence of T. chinensis Bunge on the list of species is doubtful.
Recently A. S. Erst et al. (2019) described a new species from South Siberia — T. austrosibiricus Erst et Luferov. The authors compared it with T. chinensis and T. macropetalus (here considered as comb. inval.), i. e. with T. chinensis. A "Key to identification species of the genus Trollius from Russia" comprising 16 species was given (again, the recent revision of the genus Trollius in Japan by Kadota (2016) is ignored).
Trollius chinensis was chosen as a focus of the present paper, since the current status of knowledge of taxonomy and geography of this species appeared to be inadequate, although its distribution range in the Asian part of Russia is vast, morphological variation range is significant and the number of contact zones with other Trollius species is high.
Materials and methods
I have been studying taxonomy and geography of the genus Trollius since 2013. All the authentic specimens of Trollius in LE (alongwith a several type specimens from P and K) have been thoroughly studied, as well as important herbarium collections (LE, MW, MHA, MWG and others). Field material was collected in 2019 in Tunkinskii District, Buryatia (S macroslope of East Sayan Mountains) and N macroslope of Khamar-Daban Mountain Ridge, Baikal State Nature Reserve, Kabansk District, Buryatia. Field observations of previous years (1981-1992) in different regions of Siberia (Altai, Kuznetsk Alatau, East Sayan, Khamar-Daban, etc.) were analyzed. All the collected herbarium specimens are deposited in MHA, living plants are cultivated in Tsitsin Main Botanical Garden of RAS. Molecular genetic studies of species (currently T. chinensis and T. asiaticus) and interspecific hybrids (plants from hybrid populations T. chinensis x T. asiaticus) of the genus Trollius have been started.
Habitat data are recorded including:
— inventory and mapping of Trollius species populations;
— geobotanic descriptions of the plant communities in which species of the genus Trollius occur.
Another important source of valuable information is represented by photographs from natural habitats in different parts of the species distribution range.
Taxonomic treatment
Trollius chinensis Bunge, 1833, Enum. Pl. China Bor.: 3; id. 1833, Mem. Pres. Acad. Imp. Sci. St. Petersb. 2 [marked "1835" on front page], 1-2: 77; Luferov,
Fig. 1. Syntype of ^ollius ledebourii var. macropetalus (P00194367,
https://science.mnhn.fr/institution/mnhn/collection/p/item/p00194367?listIndex=235&listCount=1193).
P00194367
2004, Turczaninowia, 7 (1): 8; Luferov et al. 2018, Turczaninowia, 21 (2): 112. = T. asiaticus L. var. chinensis (Bunge) Maxim. 1889, Enum. Pl. Mongolia, 1: 25; Finet et Gagnep. 1904, Bull. Soc. Bot. France, 4: 396; Craib, 1914, Curtis's Bot. Mag. 140: tab. 8565; Schipcz. 1924, Izv. Glavn. Bot. Sada RSFSR, 23: 69; id. 1937, Fl. URSS, 7: 47; Sipliv. 1972, No-vosti Sist. Vyssh. Rast. 9: 169-170; Dorosz. 1974, Monogr. Bot. 41: 74.
Holotype ("lectotypus", Grabovskaya-Boro-dina, 2010: 372): "China, 1831, d[ed]. Bunge", fl. fragm. (LE: LE 01013585!).
Protologue: "Tr. sepalis 10-12 patulis, peta-lis 20 ligulatis sepala aequantibus, stamina plus duplo superantibus. Hab. in Chinae borealis provincia Schan-ssi, ubi flores in usum medicinalem colligun-tur. Flores tantum exsiccatos vidimus".
= Т. ledebourii Rchb. [var.] y macropetalus Regel, 1861, Mém. Acad. Imp. Sci. Pétersb., sér. 7, 4, 4 (Tent. Fl. Ussur.): 8. = T. chinensis subsp. macropetalus (Regel) Luferov, 1991, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 96, 5: 74; id. 2004, Turczaninowia, 7 (1): 8; Luferov et al. 2018, Turczaninowia, 21 (2): 113. — T. macropetalus (Regel) F. Schmidt, 1868, Mém. Acad. Imp. Sci. Pétersb., sér. 7, 12, 2 (Reisen Amur-Lande): 88, comb. inval.; Vorosch. 1982, Opred. Rast. Sovet. Dalnego Vostoka: 260.
Syntypes: "Coast of Manchuria, lat. 44-45 N, 1859, C. Wilford" (K, P00194367!). — Fig. 1.
Protologue: "floribus circiter 6-sepalis, se-palis suborbicularibus quam petala brevioribus. P. [corr.: T.] chinensis Max. prim. pag. 22 et Wilf. pl. exsice. [corr.: exsicc.] Am Tatarischen Golf (Kusne-tzoff). An der Küste der Mandschurei (Wilford)".
= T. vitalii Stepanov, 1994, Florogenet. Analiz, 1: 101.
Holotype: [Krasnoyarsk Territory] Ерма-ковский р-н, Западный Саян, хр. Кулумыс, окр. Ойского озера, субальпийский луг [Yermakovs-koye District, West Sayan, Kulumys Ridge, Ois-koye Lake vicinity, subalpine meadow], 11 VI 1990, N. Stepanov (NS0000407!, labelled "Isotype"); isotypes — LE: LE 01038901!, KRAS (not seen).
Protologue: "Typus: Sajan Occidental[is], trajectus Bujbinskij, vallis fl. Bolschaja Oja, pratum subalpinum, 11 VI 1990, N. Stepanov (NS, isotypi — LE, KRAS)... Тип: Западный Саян, хр. Кулумыс, окр. Ойского оз., субальпийский луг [West Sayan, Kulumys Ridge, Oiskoye Lake vicinity, subalpine meadow], 11 VI 1990, Н. В. Степанов [N. V. Stepanov] (NS, изотипы [isotypes] — LE, KRAS)".
— T. ledebourii auct. non Rchb.: Kom. 1903, Acta Horti Petropol. 22, 1 (Fl. Manch. 2, 1): 230.
Perennial herbs with simple or branched (up from the middle part) erect stems (20)30-120 cm tall, with one or several (usually 2) flowers; stems smooth, striate, enclosed with fibrous remains of the previous year's petioles at the base. Rhizome short, with brownish black thread-like roots. Basal leaves long-petioled, (1-2)3-6(10), 3-5-divided into more or less rhombic lobes; those dissected, margins serrate-dentate, with ovate to ovate-elliptic teeth shortly apiculate at the apex. Petioles smooth, up to half of the stem length or slightly longer, without pronounced geniculum in upper part. Leaf blade thin coriaceous (to coriaceous in the plants growing at highest altitudes); upper surface bright to dark green, velvety, with deeply impressed veins, sometimes with purple spot at the centre; lower surface contrastively silvery green, glossy, veins prominent, pubescent with appressed hairs (the contrast pattern persists in herbarium). Cauline leaves 3-7, lower ones on petioles up to 13(15) cm long, similar to the basal in shape, upper sessile, smaller (reduced); peduncles 5-15 cm long, getting longer at frutescence (to 20 cm long). Flowers 4-6 cm in diam., shallow bowl-shaped to saucer-shaped. Sepals of different tones of orange, usually bright orange (often turning yellow in herbarium specimens), slightly concave, broadly ovate to broadly elliptic, sometimes erose at the apex (sepals of the lower circle sometimes with a few teeth of irregular shape at the apex, tinged green), (5)6-12(14); petals (nectaries) about 20, getting (significantly) longer in course of anthesis, (subequal or shorter than sepals at the early flowering stage) longer (up to 2 times) than sepals, linear, more or less long aculeate or acute towards the apex (in herbarium specimens — to sword-like), 1.7-3 mm wide, same colour or slightly darker than sepals (in herbarium specimens often brighter than sepals), throughout the length fleshy but not thickened; nectary pit in the shape of a flat pocket, 1.5-3 mm from the base. Stamens two times shorter than petals, 13-20 mm long, incl. the anther length (2.5-4 mm), pale yellow to yellowish. Anthers with short acumen at the apex. Ovary green, more or less arcuate or curved, 3-3.5 mm long, style shorter than ovary (or subequal), stigma yellow. Follicles 12-40, in a head 1-2 cm diam.; follicle up to 14 mm long, with a beak up to 4.5(5) cm long; beaks straight, sometimes bent outwards at the basis, slightly arcuate (in herbarium specimens more often), (1.5)2-3.5(4) mm long, purplish/tinged purple as well as the area around their bases. Seeds 3-7 per follicle, dark brown, matt, oval. Flowers in June — July; second flowering (mid — late August) recorded in Amur Region and Primorye Territory. 2n = 16 (Doroszewska, 1974). — Figs. 2-4.
Fig. 2. Trollius chinensis at anthesis.
1-3 — Primorye Territory, Ussuriysk Municipal Region, Monakino vicinity, 2 VII 2008, photos by L. V. Kraynik; 4 — Kunashir Island, Kurilskii State Nature Reserve, Filatov's cordon vicinity, VII 2002, photo by Yu. V. Ovchinnikov; 5 — Khamar-Daban Mountain Range, Snezhnaya River valley, 25 VII 2016 (plant of hybrid origin — T. chinensis x T. asiaticus), photo by Yu. V. Ovchinnikov; 6 — plant from old horticultural collection at Botanical Garden of Irkutsk State University, photo by S. S. Kalyuzhny.
Fig. 3. Trollius chinensis at fruiting.
1, 2 — fruiting plant: Republic of Buryatia, Tunkinskii District, Tubota River valley, study plot 2019-Tubota_main, 24 VII 2019, photos by K. P. Savov; 3, 4 — fruiting heads: Primorye Territory, Shkotovskii District, Sukhodol River basin, Novonezhino vicinity, 6 VII 2014, photos by L. V. Kraynik.
Table. Key characters to differentiate Trollius chinensis and T. ledebourii
Diagnostic character Trollius chinensis Trollius ledebourii
Petals per flower About 20 Up to 10-12
Petals/stamens length ratio 2 1.2-1.4
Petal apex Long aculeate/acute Rounded, often incised
Anther length and shape Up to 4 mm, more or less cylindric Up to 8 mm, elongate and flattened
Teeth of leaf blade margin More or less equal in size, ovate to ovate-elliptic, shortly apiculate at the apex Teeth of irregular size and shape, acute or aculeate at the apex
Leaf blade texture Coriaceous, upper surface velvety Thin to thin coriaceous (rarely and mostly in hybrid plants), upper surface smooth
Beak of the follicle, length Up to 5 mm Up to 2 mm
Comments to morphological description. All the leaves to illustrate intrapopulation variability of the basal leaf blades (Fig. 4) were collected within the study plot 2019-Tubota_main (3.4 x 7.5 m2 ; 51.872618° N 102.802318° E; h = 1039 m a. s. l.).
Flowers with number of sepals around 14 could be observed mostly in cultivated plants, often of hybrid origin (Doroszewska, 1974). Forming follicles of T. chinensis x T. asiaticus are often characterized by rather big purple spots and red to purple sutures.
According to our observations, the beaks of mature follicles remain straight up to opening, while in herbarium specimens of the same plants the beaks could turn arcuate in course of pressing and drying.
Affinity
Morphologically, T. ledebourii should be considered the closest species to T. chinensis. Both species have shallow bowl-shaped or saucer-shaped flowers (almost obligatory flat saucer-shaped in T. ledebourii) and long linear and not thickened petals always longer than stamens. At a superficial glance the plants with small number of sepals from T. chinensis populations are particularly close to T. ledebourii. Some important diagnostic characters to differentiate these species are presented in the Table.
In the contact zones of T. chinensis and T. ledebourii distribution ranges (e. g., in Amur Region) these species often form spontaneous hybrid populations, that causes identification difficulties. Many horticultural hybrids of these species have been originated long ago and still are widely cultivated.
Taxonomic and nomenclatural notes
The history of Bunge's publication "Enumeratio plantarum, quas in China boreali collegit" (Bunge, 1833) which caused so many errors in nomenclature citations of the species, described by the author from his collections in China (1830-1831), is analyzed in detail by A. E. Borodina-Grabovskaya (2007). It appears, though, it is 1833 that should be considered a year of
the actual publication of issues 1-2 of the second volume of "Mémoirs de l'Académie Impériale des Sciences de St.-Pétersbourg... Tome second, 1re & 2me livraison". Besides that, I am supporting and following her decision to cite the both preprint and the publication of 1833 at all those taxa.
The only known to the date original specimen of T. chinensis, cited as the lectotype by Grabovskaya-Boro-dina (2010), contains 8 flowers in different safe state, stored in an envelope. No other authentic specimens have been found so far (P, TU). Bunge notices in the protologue: "Flores tantum exsiccatos vidimus [I have seen dried flowers only]" (Bunge, 1833: 3), therefore the cited specimen is most probably the only element of the original material, i. e. holotype (Turland et al., 2018: Art. 9.1(b)). According to Bunge (1833) flowers of T. chinensis were used in traditional local medicine.
The history of the name T. macropetalus (Regel) F. Schmidt is curious and paradoxical. Authors of many taxonomic papers referred to the work by F. Schmidt (1868: 88) as a place of valid publication of this combination. As a matter of fact, one can find on that page the only mention of the name T. macropetalus in a more than 220-pages paper. It says: "An Kräutern in dieser Region wüsste ich im Süden nur wenig Neues zu nennen, so Trollius macropetalus und Lilium Glehni [I have not known a lot about herbs in south of this region, such as Trollius macropetalus and Lilium glehnii]". In contrary, neither of the researchers referred to the nomenclature citation, diagnosis and taxonomic comment for T. ledebourii var. macropetalus given by Schmidt on pages 105-106 of the publication (within a Synopsis — section Specieller Theil):
"18) Trollius Ledebourii var. macropetala Rgl. et Maak fl. ussur. n. 24. T. chinensis Max. n. 38.
T. 1-2 pedalis, foliis supra glauscescentibus, sepalis 5-7 rotundatis patulis, petalis c. 20-25 erectis lineari-lanceolatis aurantiacis sepala superantibus, quam stamina duplo longioribus stylis rectis 1/4 carpellorum non superantibus.
Fig. 4. Intra-population variability of basal leaves in Trollius chinensis: Republic of Buryatia, Tunkinskii District, Tubota River valley, study plot 2019-Tubota_main, 24 VII 2019, photo by K. P. Savov.
Differt a. T. Ledebourii genuino petalis longioribus stylisque rectis brevioribus; a. T. chinensi Bge. sepalis paucioribus patulis.
T. japonicus Miq. l. c. III, p. 6 petalis stamina aequan-tibus nec illa sub duplo superantibus differe videtur".
V. N. Siplivinsky (1972) followed by A. N. Luferov (1991) additionally referred to page 218 of the Schmidt's publication where one can find the following paragraph: "Ich bin mehrfach in das bekannte Dilemma von gutten und schlechten Arten gerathen, und es finden sich noch jetzt unter meinen neuen Species einige, die vielleicht mit mehr Recht als Varietäten bei schon bekannten Arten untergebracht werden könnten, währeud manche Varietäten wiederum von Anderen als gute Species hingestellt werden moechten. Ich will hier gleich auf diejenigen Species hinweisen, deren Artrechte ich für zweifelhaft halte, wie auch diejenigen Varitäten, die vielleicht wieder zu Species erhoben werden duerften: Trollius Ledebourii var. macropetala, die ich früher als eigene Species hingestellt hatte, kann doch vielleicht eine gute Art sein. [I have repeatedly fallen into the well-known dilemma of good and bad species, and among my new species there are still some
which perhaps may be better treated as good varieties, and some varieties, in turn, considered by others to be good species I would rather like to discard. I will immediately point out the species which I consider doubtful, as well as the varieties which perhaps may be raised to species level: Trollius Ledebourii var. macropetala which I had considered earlier to be a separate species may appear to be a good species.]" (Schmidt, 1868: 218).
Trollius ledebourii var. macropetala opens a list of doubtful taxa with no final taxonomic decision taken, so it is the only accepted name for the taxon in this publication. Occasional usage of combination T. macropeta-lus on p. 88 of the publication is most probably a lapse and/or trace of the author's reminiscences about evaluating the status of this taxon. The construction "I had considered earlier" on p. 218 refers to the time when F. Schimdt had started to evaluate this taxon, not to the previous episode in the publication. Page 105 should be the only correct reference in the respective nomenclature citation.
Thus, the combination "Trollius macropetalus" was not explicitly accepted by Schmidt, therefore it was not
validly published (Turland et al., 2018: Art. 36.1) and should be withdrawn from usage.
Treating of plants with a small number of strongly deflexed sepals (6 according to the protologue; 5-7 according to Schmidt (1868)) and petals up to 35 mm long gradually tapering at the top as a separate subspecies T. chinensis subsp. macropetalus (Regel) Luferov (Luferov, 1991, 1995) appears to be an unsatisfactory decision due to the following reasons:
— number of sepals within a single population varies significantly; the plants at the first anthesis are often characterized by a small number (5-7) of sepals;
— a long-lasting growth of petals often results in changing the shape of apex (gradual tapering/sharp narrowing) during anthesis;
— sepals deflection angle also changes significantly in course of anthesis.
Luferov (1991) suggested that there are graceful transitions between type subspecies and T. chinen-sis subsp. macropetalus in a form of clinal characters change (from West to East): e. g., number of sepals reduction, petal length increase, petal shape change. That is not corroborated by our data from different parts of T. chinensis distribution range: e. g., in the Far East populations plants with 7-10 sepals per flower were recorded many times while within the western sector of the distribution range plants with 5-7 sepals per flower are not rare.
Siplivinsky (1972) considered T. macropetalus as a synonym of T. chinensis. He commented on intraspe-cific variability of the species as follows: "Growing in such a vast mountain territory T. chinensis does not remain stable in its morphological characters. E. g., in the South, in Changbai Mountains, the most robust plants with big flowers and leaves, petals 1.5-2 times longer than sepals (5-8 per flower) could be observed. In the North and East (e. g. Primorye) — in contrary, plants with 5 sepals and shorter petals predominate. In the West number of sepals varies from 8 to 12 and petals are only slightly longer than sepals or subequal to them" (Siplivinsky, 1972: 169-170, translated from Russian). Taking into consideration lack of data from a significant part of T. chinensis distribution range, a big number of wrongly identified herbarium specimens and lack of in-depth knowledge of variation within hybrid populations of the species I consider it premature and inaccurate to argue the clinal characters change in T. chinensis.
A broad concept of T. chinensis is adequate and instrumental. However, a correct identification of herbarium specimens imposes a simultaneous presence of petals, basal leaves and anthers on the voucher, since the labels almost never contain detailed information on phenology/population patterns in the locality. For ex-
ample, plants of hybrid origin (T. chinensis x T. ledebourii) are often characterized by 5-6 sepals and very long petals gradually tapering to the apex, but petal shape (rounded, often incised at the apex) and long anthers on short and weak filaments) indicate "blood" of T. ledebourii in the plant.
N. V. Stepanov (1994) published one new species, two its varieties and one form from West Sayan: T. vi-talii Stepanov, T. vitalii var. forficuloides Stepanov, T. vi-talii var. nadezhdae Stepanov, and T. vitalii f. asiaticifo-lius Stepanov. As diagnostic characters of T. vitalii the author mentioned long straight beaks of follicles (up to 3-3.5 mm long), petals shorter than sepals, curved ovaries, style as long as ovary. All those characters agree with T. chinensis diagnosis and variation range, which is completely corroborated by authentic specimens. It is worth mentioning that the author stated in diagnosis/ description of T. vitalii that the number of its sepals varies from 16 to 25. The number of sepals in the original specimens (KRAS, LE and NSK) does not, however, exceed 12. Comparatively short petals in flowers of the original specimens indicate early flowering phase.
It is possible that the author did not consider T. chinensis to be (at least) an affinity of T. vitalii, since none of the available bibliographic sources contained information on T. chinensis distribution in Siberia.
On the website of the Plantarium project (https:// www.plantarium.ru/), there is a gallery of T. vitalii images (Trollius vitalii Stepanov, 2007-2019), where all the depicted plants are identified by N. V. Stepanov and several ones are published by him. One of them (Stepa-nov, 2009) is taken in locus classicus of T. vitalii, and illustrates a plant with a flower of 14 sepals. Some other photographs (e. g., Dragan, 2016; Pool, 2018) represent plants of a hybrid origin (T. chinensis x T. asiaticus).
Hybrid genesis of the illustrated plants is corroborated by petals morphology, flower shape and other characters. T. chinensis x T. asiaticus hybrids differ from T. chinensis s. str. by broadly lanceolate multiveined petals (similar to T. asiaticus) rarely exceeding sepals, globose (not shallow bowl-shaped) flower of many sepals arranged in circles (similar to T. asiaticus).
In my opinion, T. vitalii should be placed in a synonymy of T. chinensis; the infraspecific taxa described by Stepanov should not be treated taxonomically but taken into consideration from the perspective of studying variation range, particularly in hybrid populations, since they are, in fact, interspecific hybrids T. chinen-sis x T. asiaticus.
Recently described Trollius austrosibiricus Erst et Luferov (Erst et al., 2019) is most likely an interspecific hybrid T. asiaticus x T. chinensis: morphological comparison between T. austrosibiricus and related species,
presented by the authors, does not contain diagnostic differences, e. g. one can find lots of plants with simple rhizomes within a single T. chinensis population; all the morphometric differences mentioned are not in fact diagnostic. However, the shape of petals is very typical of T. asiaticus x T. chinensis hybrids: it is very close to that of T. asiaticus (broad and multiveined), except for the apex which is not rounded but clearly acute. For the time being (and until a thorough study of all the authentic specimens of T. austrosibiricus — holotype and six paratypes are cited) I will keep considering this taxon as another T. asiaticus x T. chinensis hybrid. It is weird, that the authors do not comment on T. vitalii although one of T. austrosibiricus paratypes was collected near locus classicus of the former species.
Distribution
General distribution range. Asian part of Russia, 700-2100 m alt.; Mongolia (Uvs Province, Harhiraa), ca. 2700 m alt.; North and North-East China (Hebei, N Henan, W Jilin, W Liaoning, E Nei Mongol, Shanxi provinces), 1000-2200 m alt. ("on grassy slopes" — Li, Tamura, 2001); north of Korean Peninsula.
Distribution in Asian part of Russia. Krasnoyarsk Territory. Yermakovskoye District: West Sayan (Kulumys Ridge, Oiskoye Lake vicinity, Bol-shoi Kebezh River upstream). Republic of Buryatia. Tunkinskii and Okinskii Districts: East Sayan (Orlik vicinity, Mondy vicinity, Arshan vicinity, Tubota River
valley); Zakamensk District: Jida uplands (Tsakirka River upstream, Malyi Subutui River fountain); Ka-bansk District: Khamar-Daban Mountain Range (Sne-zhnaya River valley, 4 km upstream from Monomakh Hat peak; Baikal Nature Reserve); Barguzin District, near Bolshoi Chivyrkui River mouth; Bauntovskii Evenkiiskii District. Trans-Baikal Territory. Amur Region (south). Sakhalin Region. Sakhalin (south-west, Holmsk vicinity), Kunashir Island (Kurilskii State Nature Reserve, Filatov's cordon vicinity). Khabarovsk Territory (near Grossevichi Bay). Primorye Territory
(Fig. 5).
The herbarium specimen from Mongolia (Harhiraa Ridge, Tsagan-Dygl Mt., Ogureyeva (MWG!)) not only proves the Mongolian locality but also marks the highest elevation (2700 m alt.) T. chinensis reaches within its distribution range (Fig. 6). Occurrence of T. chinensis in Mongolian Altai seems to be highly probable.
Previous views on T. chinensis distribution range in the Asian part of Russia (Schipczinsky, 1937; Siplivin-sky, 1972; Voroschilov, 1982; Luferov, 2004; Luferov et al., 2018) should be radically changed. The material studied during this research project (herbarium specimens, living plants collected in the wild, photographs from different parts of distribution range, etc.) proves a vast distribution range of T. chinensis in the Asian part of Russia (from West Sayan in the west to Kunashir Island in the east). Moreover, it is the second-third in area within the genus after T. asiaticus and T. sibiricus Schipcz.
Fig. 5. Distribution of Trollius chinensis in the Asian part of Russia and Mongolia.
Fig. 6. Herbarium specimen of Trollius chinensis from Mongolia (MWG).
The distribution ranges of T. chinensis, T. asiaticus, T. sibiricus, T. ledebourii and T. riederianus s. l. within the Asian part of Russia overlap on a large area and extent, forming numerous contact zones and hybrid populations. Further studies should be focused on improving the knowledge of the distribution ranges, revealing and describing the hybrid populations and carrying out the necessary molecular genetic research.
Habitat and ecology
Forest and forest-steppe altitudinal zones: swamp coniferous forests — swamp larch open forest with dwarf bog birch (tree and shrub strata dominants: Larix gmelini (Rupr.) Kuzen., Betula pendula Roth, Salix beb-biana Sarg. — Betula fruticosa Pall.) on a flat summit of the coniform hill, h = 1090 m alt. (Selyutina, 2018, pers. comm.); wet meadows on fertile soils, forest edges, riversides and towpaths, open/thin forests at the upper forest boundaries. Subalpine altitudinal zone: subalpine meadows, often near water streams. Alpine (goltsy) al-titudinal zone: meadows near headstreams, subniveal small meadows, 1600 m a. s. l. (Khamar-Daban) — 2700 m a. s. l. (Harhiraa Ridge).
According to our observations in Tunkinskii District, Tubota River valley (July 2019) the populations of T. chinensis were recorded within the higher part of the forest altitudinal zone — in forest edges, riversides and roadsides. One population was described in details (results to be published elsewhere): Republic of Burya-tia, Tunkinskii District, Tubota River valley, study plot 2019-Tubota_main (3.4 x 7.5 m2); 51.872618° N 102.802318° E, h = 1039 m a. s. l. It comprised 47 plants: 15 fertile mature, 14 immature, 18 juvenile.
Cultivation
T. chinensis — one of the most attractive species of the genus Trollius — was introduced in European horticultural collections at the end of 19th century along with several other species of Globe Flower (Trollius). A mess with names of cultivated plants existing at that time resulted in another introduction (re-introduction) of this species in the beginning of 20th century in the UK by the company J. Veitch & Sons. The company distributed the plants of the clone collected in China by W. Purdom. Thus, T. chinensis gained a broad popularity and won the praise of publication (and colour plate) in Curtis's Botanical Magazine (Craib, 1914).
Practically from the very beginning of the initial introduction numerous horticultural cultivars were originated on the basis of T. chinensis and/or via its hybridization with other species. Few species were used for hybridization: T. europaeus L. and T. ledebourii among them (Doroszewska, 1965, 1970). As a result of the
both targeted and spontaneous hybridization in garden collections many forms carrying labels T. chinensis (being very different from the species morphologically) became widespread across Europe. One of the consequences of that tendency was a misconception of the species which is still existing among horticulturalists.
Selection of the horticultural forms was aimed to create tall, multi-flowered plants with big bright flowers (of many sepals) and maximally long petals (Fig. 2: 6). That is why horticultural forms with extremely long petals (often incised at the apex) are very popular.
Geographically representative selection of collections studied
Herbarium specimens: Appendix 1 (see the journal's website: www.binran.ru/journals/novitates/).
Photographs from natural habitats: Appendix 2 (see the journal's website: www.binran.ru/journals/ novitates/).
Acknowledgements
Keepers, curators and staff members of the following Herbaria are greatly acknowledged for assistance: LE, MHA, MW, MWG, P. I would like to thank K. P. Savov, I. N. Urbanavichene and G. M. Serebryanyi for participation in field research. I am very grateful to fellow botanists and plant amateurs, supplying me with well-documented and high-quality images, particularly to participants of the project Plantarium (www.plan-tarium.ru) for long-term fruitful collaboration. Informal group of friends, colleagues and fellow botanists is deeply acknowledged for providing photographs, observations and another important materials for study (in alphabetical order): N. S. Gamova, S. S. Kalyuzhny, L. V. Kraynik, Yu. V. Ovchinnikov, I. Yu. Selyutina, M. V. Skotnikova, N. N. Voropaj. Dr. I. V. Sokolova is sincerely acknowledged for reviewing and editing the paper, particularly for the discussion on attributing the date of the Bunge's publication. Dr. A. N. Sennikov is acknowledged for the discussion on Trollius macropeta-lus. I am also grateful to Prof. G. N. Ogureyeva and Dr. A. K. Sytin for valuable discussions. I am deeply grateful to my family for support.
Supplementary materials (Appendices 1, 2) to the article are available on the journal's website (www.binran.ru/journals/novitates/).
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