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33Systematic notes..., 2018, 118: 64-76 DOI: 10.17223/20764103.118.5
>Ь-Щя^рЧ* ii
УДК 582.675.1+581.9
The genus Trollius L. (Ranunculaceae) in the flora of the Altai Mountain Country
A.S. Erst1, 2, A.N. Luferov3, V.I. Troschkina1, K. Xiang4,5, W. Wang4,5
institution of Russian Academy of Sciences, The Central Siberian Botanical Garden, The Siberian Branch of the RF Academy of Sciences, Zolotodolinskaya str., 101; 630090, Novosibirsk, Russia. E-mail: [email protected]
2Tomsk State University, Laboratory of structural and molecular analysis of plants of the Biological Institute of TSU, Lenin str., 36; 634050, Tomsk, Russia 3I.M. Sechenov First Moscow State Medical University, Pharmaceutical Natural Science Department, Izmailovsky Boulevard, 8, 105043, Moscow, Russia; [email protected] 4State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, 100093, Beijing, China 5University of Chinese Academy of Sciences, Beijing 100049, China
Abstract. Information on new characters, that were not previously used for the Trollius species identification is presented: consistence and degree of dissection of blade, type of margin, shape of teeth, number of petal veins, shape of nectarostigma, and morphological characters of fruits - type of the pericarp surface. The quantitative and qualitative morphological characters of the species are indicated: number of sepals and petals, distance from the petal base to the nectarostigma, length of the follicle and persistent style, as well as its shape and location. An identification key is proposed, which has been developed taking into account the suggested characters. A sinopsis of the genus Trollius (Ranunculaceae) of the Altai Mountain Country, including 3 species belonging to 2 sections is given. The synopsis includes nomenclature quotations, information on type specimens, distribution in the region, and general distribution for each species.
Key words: Trollius, Ranunculaceae, diagnostic characters, Altai Mountain Country.
The genus Trollius L. includes a 35 species (excluding Hegemone Bunge ex Ledeb.) distributed in the extratropical regions of the Northern Hemisphere. Hegemone is treated as a distinct genus (Schipczinsky, 1937). The common name of some species is "globeflower" (globe flower) or "Trollblume" (Steam, 1992). The genus is characterized by conspicuous orange- or yellow-colored flowers, similarly colored petals, subscapose habit, and ternate or deeply 3-lobed leaves (Kadota, 1987). Trollius is characterized by an unusual floral structure with petals divided into the blade, pit (nectarostigma) and claw (Wang et al., 2010). The length ratio of nectaries to stamens was considered to be one of the most important morphological characters for species delimitation (Schipczinsky, 1937; Siplivinsky, 1972).
A. Doroszewska (1974) divided the genus into seven sections. Sections are mostly natural, but the gaps among them seem to be too small to assume them as distinct ones (Tamura, 1995). In Russia, about 19-20 species are recognized (Luferov et al., 2018). The greatest species diversity is observed in Siberia (12 species: our data). Eleven species of Trollius were previously indicated for that region (Friesen, 2003). Many taxa described recently from Russia (Malyshev, 1965, Siplivinsky, 1972, 1973; Stepanov, 1994, 2018) require more detailed investigation and evidence of species independence (Luferov et al., 2018).
The Altai Mountains, sometimes referred to the Altai Mountain Country (AMC), are located in the borderland between Russia, Kazakhstan, China, and Mongolia (Kamelin, 2005). The AMC is situated in the western part of the Altai-Sayan ecoregion and stretches from approximately of N 45° to 54°, and E 80° to 100° as outlined by R.V. Kamelin. This region crosses not only administrative boundaries, but also floristic boundaries, and the region's varied topography, coupled with the intersection of floristic boundaries, has led to a degree of endemism in the Altai flora (Orme et al., 2005).
The aim of this work was to study the taxonomy and characters of the Trollius species in the flora of the Altai Mountain Country. Firstly we identified diagnostic morphological characters of the Trollius species of the Altai Mountain Country, searched for new charactes for the identification of taxa, compiled the key for species identification, and analyzed the distribution of Trollius species in the AMC.
Material and methods
Morphological and geographical methods were used. The specimens deposited in the following Herbaria: V.L. Komarov Botanical Institute (LE), the Main Botanical Garden (MHA), M.V. Lomonosov Moscow State University (MW), Central Siberian Botanical Garden (NS, NSK), Altai State University (ALTB), and Tomsk State University (TK), were examined. Meanwhile, living plants were also collected during 20 expeditions (2000-2018) in the Russian and Chinese Altai (2014, 2016). The photographs in the field were taken with a Nikon D90 camera. The morphological characters were measured using AxioVision 4.8. The flowering and fruiting periods and habitats are given as cited on the collector's labels. Data on chromosome numbers are presented by the Chromosome Counts Database, CCDB (Rice et al., 2015).
In this study, we consider the Altai Mountain Country within its botanical -geographic borders, and follow R.V. Kamelin (2005) in dividing the AMC into floristic provinces and regions. These are: Altai province (regions: A1 - Northern Altai, A2 - North-Eastern Altai, A3 - Central Altai, A4 - Tchulyshman, A5 -Abakan-Dzhebash, A6 - Khemchik), Altai-Dzungarian province (regions: KAD1 -North-Westrn Altai, KAD2 - Kalbinsky, KAD3 - Tarbagatai, KAD4 - Saur, KAD5 - Zaissan, KAD6 - Bukhtarma, KAD7 - Markakol-Kanas, KAD8 - Kara-Irtysh, KAD9 - Altai-Dzungarian), and Tuvinian-Mongolian province (regions: ZM1 - Chuya-Khobdo, ZM2 - Tsagan-Gol, ZM3 - Khobdo-Tonkhil; UM -South-Mongolian).
Results and discussions
We recognize three species of the genus Trollius within the AMC: T. asiaticus L., T. altaicus C.A. Mey, and T. dschungaricus Regel. Some important characters can be used to distinguish all three species (Figure 1, 2; Table 1). The margin of blades and the shape of teeth are significant characters for T. altaicus and T. dschungaricus. Trollius altaicus is characterized by short teeth, arcuate-convex along the edge, and acute or slightly obtuse at the apex. In contrast to T. altaicus, T. dschungaricus is distinguished by small pointed teeth. An important character crucial for species identification is sepal dimorphism (the shape of the sepal can differ in one and the same flower). Trollius dschungaricus and T. altaicus exhibit this character, whereas T. asiaticus has identical sepals. The most significant character for identification of the taxa is the length of the persistent style. Trollius altaicus has the longest persistent style, which distinguishes it from two other species.
Figure 1. Species of the genus Trollius L. from Altai Mountain Country
A, B - flower of T. asiaticus L.; C, D - flower of T. altaicus C.A. Mey.; E - follicles of T. asiaticus L.; F - basal leaf of T. asiaticus L.; G - follicles of T. altaicus C.A. Mey.; H -basal leaf of T. altaicus C.A. Mey.; I, J - flower of T. dschungaricus Regel; K - follicles of T. dschungaricus Regel; L - basal leaf of T. dschungaricus Regel.
Photos: A, B - by V. Troshkina; C, D - by V. Saveliev & A. Raspopov; E - by V. Remorov; F - by P. Kosachev; G, H - by T. Rib; I - photo by A. Ebel; J, L - by
B. Kryzhatyuk, K - by A. Hamitov.
Trollius asiaticus grows in the wide range of habitats, and thus exhibits the morphological variability. Despite this, species is characterized by the constant features, such as thin, not fleshy consistence of blade of the leaf, rhombic shape of the basal leaf segments, spaced lobes of segments, oblong-triangular acute at the apex teeth with straight or slightly arcuate edges, sepals equal to petals, and very short persistent styles of follicles (in two other species, persistent styles are much longer).
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Figure 2. Leaf blade segments, sepals, stamens, petals and follicles of Trollius species:
A-E - Trollius altaicus C.A. Mey; F-J - T. dschungaricus Regel; K-O - T. asiaticus L.
(Figure by N. Prydak)
Interestingly, there are plants that combine diagnostic characters of T. altaicus and T. asiaticus leaves: lobes and teeth at the margin can sometimes be both straight and arcuate-convex on lateral sides, and the apex is pointed or acute. For T. dschungaricus, yellowish styles and stigmas are most characteristic, but sometimes they are dark-purple.
Table 1
Taxonomically significant characters of Trollius species from the AMC
Characters T. altaicus T. dschungaricus T. asiaticus
Number of basal leaf 5 3-5 5
segments (lobes in
T. dschungaricus)
Shape of the leaf blade Broadly rhombic Broadly obovate, Rhombic
segments (lobes in reniform
T. dschungaricus)
Leaf blade margins Lobes of the leaf Blade margin slightly Lobes of the leaf
segments close or with incised into short segments fan-like with
unequally serrate- crenate-dentate lobes: clearly visible grooves
dentate margins teeth small, acute or between them, margins
overlapped. Teeth rounded unequally serrate-
usually short, with dentate. Teeth oblong-
arcuate-convex edges, triangular with straight
acute at the apex, less or slightly arcuate
often slightly obtuse edges, pointed at the apex, less often acute
Leaf blade consistence Thin, not fleshy Thick, slightly fleshy Thin, not fleshy
Length of the basal 4-25(35) 4-10(15) 4-35(45)
leaf petiole, cm
Flower diam, cm 2.5-4 3.5-6 3-5
Number of sepals 10-15 5-20 10-20
Sepal shape Outer sepals broadly Outer sepals broadly Broadly elliptical or
obovate or reniform; obovate or reniform; obovate
inner ones - inner ones -
rhombiform or rhombiform or
narrowly obovate, narrowly obovate,
with cuneate base and with cuneate base and
tapering to the apex tapering to the apex
Sepal length, mm 12-19 17-29 14-24
Sepal width, mm 12-25 (outer sepals), 7-18 (inner sepals) 12-32 12-28
Sepal color Yellow or yellow- Yellow or pale- Orange or reddish-
orange yellow; occasionally slightly reddish on the outside orange
Number of petals 8-16(19) 8-14 18-30(40)
Petal shape Linear Obovate or spatulate Oblong-elliptic
Petal length, mm 9-18 5-8 12-24
Petal width, mm 0.8-1.2 1.5-2 2.5-4(6)
Shape of the petal Rounded, less often Rounded or truncated, Rounded, less often
apex truncated or with a small notch less often with a small notch acute
Number of petal veins 3-5 3 (below nectarostigma) - 5 (above nectarostigma) (3)7-17
Characters
Colour of petals
Length ratio of petal to sepal
Length ratio of petal to stamen
Nectarostigma shape
Distance from the
petal base to nectarostigma, mm
Colour of styles and stigmas
Follicle surface
Follicle length, mm
Persistent style length, mm
Persistent style shape and its location
T. altaicus
Orange or yellow-orange
Petals 2-3 times shorter than sepals
Petals 1-4 mm longer than stamens, less often almost equal to them
Narrowly-elliptical 1-1.5
Black-purple or reddish-violet
Transverse veins sharply protruding
12-17
2-3
Thin, erect or curved outwards at the base, curved to the fruit center at the apex
T. dschungaricus
Yellow
Petals 4-7 times shorter than sepals
Petals equal or 1 -3 mm shorter than stamens
Narrowly-elliptical 0.8-1
Yellowish or dark purple
Almost smooth with hardly visible veins 9-12 1.8-2.2
Erect or arcuate curved outwards
Table 1. Continued T. asiaticus
Orange or orange-red
Petals somewhat shorter or almost equal to sepals
Petals 2-3 times longer than stamens
Broadly-elliptical or obovate
2-3
Yellowish
Veins slightly protruding 8-11 0.5-1
Arcuate curved inwards or erect, deflected outwards
Interestingly, there are plants that combine diagnostic characters of T. altaicus and T. asiaticus leaves: lobes and teeth at the margin can sometimes be both straight and arcuate-convex on lateral sides, and the apex is pointed or acute. For T. dschungaricus, yellowish styles and stigmas are most characteristic, but sometimes they are dark-purple. It can be assumed that this combination of characters indicates possible hybridization between different species in contact zones. It is important to note that the length of follicles and persistent styles are very different at the different stages of development due to the features of fruit morphogenesis. Three species exhibit elongated persistent styles at the initial stage of development, but during fruiting the length of the follicle increases more than the length of the persistent style (except for T. altaicus). The identification of plants collected at different phenological phases can be erroneous if this is not taken into account.
In three species studied, the flower buds and initial stages of the flower blooming are often characterized by various colors of sepals and petals: from pale yellow, dark yellow, and greenish to yellow-orange with tints of pink, reddish, and sometimes violet. These morphological characters are of an important diagnostic value and are used in this study to compile the identification key for the species studied.
The considered combinations of structural characters typical of different species are important for species identification. The causes of the morphological variability can only be clarified by further studies using different methods.
According to A. Doroszewska (1974), species of the genus Trollius from the Altai Mountain Country belong to two sections: Trollius (T. altaicus C.A. Mey., T. dschungaricus Regel), and Longipetala Dorosz. (T. asiaticus L.). The former is characterized by petals elongated, convex, thickened, shorter than sepals (Doroszewska, 1974), whereas the latter is distinguished by petals linear, flat, thin, longer than sepals. Trollius altaicus and T. dschungaricus belong to the same group, but exhibit a number of important differences. Trollius altaicus is mainly characterized by 5 segments of basal leaves (T. dschungaricus is distinguished by 3-5-lobed leaf blade), thin, not fleshy leaf blade consistence, often with the glossy surface (T. dschungaricus is distinguished by the thick, slightly fleshy blade, often with the glossy surface).
Molecular-phylogenetic studies do not cover all groups and most species (Despres et al., 2003; Wang et al., 2010). L. Despres et al. (2003) considered T. altaicus and T. asiaticus as belonging to the same clade and different sub-clades of the molecular-phylogenetic tree. W. Wang et al. (2010) analyzed T. asiaticus only; T. dschungaricus and T. altaicus were not investigated using molecular-phylogenetic methods. Since all three species were poorly investigated by molecular methods, it is difficult to consider their taxonomic relationship and affiliation with certain groups.
Key to the Trollius species
1. Petals 18-40, 1.5-2 times longer than stamens and almost equal to or slightly shorter
than sepals. Nectarostigma broadly-ovate or obovate, 2-3 mm from the petal base. Petal veins (3)7-17. Sepals are monomorphic, broadly elliptic or obovate. Persistent style 0.5-1 mm long. Leaf lobes fan-shaped, with clearly visible grooves between them, margins unequally serrate-dentate. Teeth oblong-triangular with straight or
slightly arcuate edges, pointed at the apex, less often acute..........................T. asiaticus
+ Petals 8-16(19), about the same length as stamens or slightly longer than stamens and more than 2 times shorter than sepals. Nectarostigma narrowly ovate, 0.8-1.5 mm from the petal base. Petal veins 3-5. Sepals dimorphic: outer - broadly obovate or reniform; inner- rhombiform or narrowly ovate, with a cuneiform base, narrowed to the apex. Persistent styles longer. Leaf blades of a different structure.......................................2
2. Leaf blades thin, not fleshy, dissected into 5 broadly rhombic segments with oblong-
lanceolate, close and often overlapping lobes, their margins unequally serrate-dentate. Teeth usually short, with arcuate convex edge, acute at the apex, less often slightly obtuse. Petals 9-18 mm long, linear, longer than pistils or equal to them. Styles and stigmas black and purple, violet or reddish. Follicles 12-17 mm long. Persistent styles
2-3 mm long...................................................................................................T. altaicus
+ Leaf blades thick, slightly fleshy, triquinate; lobes broadly obovate or reniform, slightly incised along the edge into short crenate-dentate segments: teeth with small pointed tips, less often rounded. Petals 5-8 mm long, obovate or spatulate, with a rounded apex and slightly curved inwards edges, usually shorter than pistils. Stiles and stigmas yellowish or dark-purple. Follicles 9-12 mm long. Persistent styles 1.8-2.2 mm long ..............................................................................................................T. dschungaricus
Sinopsis for the Trollius genus of the AMC
Subgen. 1. Trollius - Flowering shoots apical. The inflorescence cymose or flowers solitary. Basal leaves (or just a leaf) arranged in rosettes. Stem leaves sessile or petiolate. Rhizomes branching and ascending.
Type: T. europaeus L.
Sect. Trollius, 1974, Monogr. Bot., 41: 159. - Sect. Eutrollius Prantl in Engler u. Prantl, 1891, Nat. Pflanzenf., 3(2): 56, p.p. - Petals elongated, convex, thickened, shorter than sepals (Doroszewska, 1974).
Type: T. europeus L.
1. Trollius altaicus C.A. Mey., 1831, Verz. Pfl. Cauc.: 200; Krylov, 1931, Fl. Zap. Sib., 5: 1122; Schipcz, 1937 in Fl. USSR, 7: 45; Gamajun., 1961, in Fl. Kazakhst., 4: 14; id., 1969, 1ll. opred. rast. Kazakhst.: 365; Pachom., 1972, in Opred. rast. Sred. Azii, 3: 144; Revushkin, 1988, Vys. fl. Alt.: 64; Grubov, 1982, in Opred. sosud. rast. Mongolii: 108; Friesen, 1993, in Fl. Sib., 6: 104; Gubanov, 1996, Konsp. fl. Vneshn. Mong. (sosud. rast.): 51; Liangqian, Tamura, 2001, in Fl. China, 6: 140; Borodina-Grabovskaja, 2001, in Pl. Centr. Asia, 12: 18. -T. caucasicus C.A. Mey. non Steven, 1830, in Ledeb. Fl. Alt., 2: 301.
Lectotype (Siplivinsky, 1972: 168): "Altai, in pratis subalpinis legit Dr. Bunge" (LE!). A.E. Grabovskaya-Borodina (2018: 233) published the following syntypes: Western Siberia, Russia: "In pratis subalpinis, Altai [fl.], Bunge, Herb. Meyer" (LE); Kazakhstan: "In pratis subalpinis, Altai [fl.], Bunge, Herb. Meyer" (LE); "R[iddersk], 1 V [1826], [fl.], № 57, Herb. Ledebour" (LE); "R[iddersk], 18 V [1826], [fl.], № 206, Herb. Ledebour" (LE); "ad Schilgan fluv. 12 VI [1826], [fr.], № 390, Herb. Ledebour" (LE).
Description: Perennial herbs 15-60 cm high, to 80-100 cm at fruiting. Stem straight, simple or branched, smooth. Basal leaves with petioles 4-25(35) cm. Leaf blade ones or less dissected. Lower cauline leaves petiolate, upper ones sessile, or all cauline leaves sessile. Pedicle 1-12 cm, elongating to 25 cm at fruiting. Flowers 2.5-4 cm in diam., open (Figure 1C, D). Sepals orange 10-15, 11-18 mm long, outer: broadly ovate or cordate, oval and tapering to the center (Figure 2B); sometimes finely dentate and greenish at the apex. Petals 10-16(19), orange or yellow-orange, 9-18 mm long, 0.8-1.2 mm wide, linear, rounded at the apex, less often truncated or with 3.5-6 x 6.5-11 cm, base cordate, palmately 5-sect, segments broadly rhombic, 3-fid; terminal segments with unequally serrate-dentate margins (Figure 1H). Teeth usually short, with arcuate convex edge, acute at the apex, rarely slightly obtuse (Figure 2A). Cauline leaves 2-4, similar to the basal a small notch, 2-3 times shorter than the sepals (Figure 2D). Stamens, including anthers, 815 mm long. Anthers 2.5-3.5 mm long (Figure 2C). Nectarostigma narrowly ovate, 0.81.5 mm from the petal base. Styles and stigmas black and purple, reddish and violet. Fruit a globose head of numerous follicles, each 12-17 mm long. Persistent style thin, erect or curved outwards at the base, curved inward at the apex, 2-3 mm long (Figures 1G, 2E). Seeds ellipsoid-globose, black, slightly glossy.
Chromosome number: 2n=16 (out of the AMC) (Doroczewska, 1967; Rostovseva et al., 1981).
Habitat: Alpine and subalpine meadows, the upper part of the forest belt; up to 2200 m.
Phenology: Flowering May - June; fruiting June - July.
Distribution: A1, A2, A4-A7, KAD1, KAD2 (Ubinsky District), KAD3, KAD4, KAD5, ZM2, ZM3.
General distribution: China (Nei Mongol, Xinjiang), Kazakhstan (Almaty Region, East Kazakhstan Region, Jambyl Region), Kyrgyzstan (Chuy Region, Issyk-Kul
Region, Talas Region), Mongolia (Bayan-Olgii Province), Russia (Altai Republic, Altai Krai, Tuva Republic) (Krylov, 1931; Siplivinskiy, 1972; Borodina-Grabovskaya, 2001; Liangqian, Tamura, 2001).
2. T. dschungaricus Regel, 1880, Acta Horti Petropol., 7: 383; Karamysheva, 1992, Fl. Tarb.: 54; Liangqian, Tamura, 2001, in Fl. China, 6: 138; Borodina-Grabovskaja, 2001, in Pl. Centr. Asia, 12: 19. - T. europaeus L. var. songoricus Regel, 1870, in Bull. Nat. Mosc., 43, 1: 243.
Lectotype (Siplivinsky, 1972: 169): [Kazakhstan] "Am 10 Mai auf den Gebirgen um das Fort Wernoje an den Quellen des Grossen Almaty-Flusses (Sewerzow)" (LE !).
Description: Perennial herbs 5-40 cm high, to 60-70 cm at fruiting. Stem simple or slightly ascending, or branched, with 1-3 flowers, smooth. Basal leaves with petioles 4-10(15) cm long. Leaf blade 1.5-4.5 x 2-7.5 cm, base cordate, palmately 3-5-sect, slightly incised into short crenate-dentate lobes at margins (Figure 1L). Teeth acute or rounded (Figure 2F). Cauline leaves 2-4, similar to the basal ones or more dissected and smaller in size. Lower cauline leaves petiolate, upper ones sessile. Pedicle 1-15 cm, elongating to 20 cm at fruiting. Flowers 3.5-6 cm in diam., open (Figure 1I, J). Sepals yellow or pale yellow, sometimes slightly reddish on the outside, 17-29 mm long, 12-32 mm wide, 520 mm wide, broadly obovate or elliptic (Figure 2G). Petals 8-14, yellow, 5-8 mm long, 1.5-2 mm wide, ovate or reniform, shorter than the sepals, equal to or shorter than the stamens (Figure 2H, I). Stamens, including anthers, 8-12 mm long. Anthers 2.5-3 mm long. Nectarostigma narrowly elliptic, 0.8-1 mm from the petal base. Styles and stigmas yellowish or dark purple. Fruit a globose head of numerous follicles, each 9-12 mm long (Figure 1K). Persistent style erect or arcuate curved outwards, 1.8-2.2 mm long (Figure 2J). Seeds round, smooth-angular, brown-black, slightly glossy.
Chromosome number: 2n = 16 (out of the AMC) (Doroczewska, 1967; Zhukova, 1967).
Habitat: Wet meadows, sedge bogs, wetted habitats in a strip of spruce forest and among juniper, along mountain streams, subalpine meadows, on peaty soil, near glacial moraines and near snow spots in the middle and upper mountain belt; up to 2500 m.
Phenology: Flowering July - August; fruiting August - September.
Distribution: KAD5 (Tarbagatay Ridge).
General distribution: Kazakhstan (Almaty Region, East Kazakhstan Region), Kyrgyzstan (Issyk-Kul Region, Chuy Region, Osh Region), Uzbekistan (Andijan Region, Fergana Region, Namangan Region, Tashkent Region), China (Xinjiang), Mongolia (Bayan-Olgii Province, Govi-Altai Province) (Siplivinskiy, 1972; Borodina-Grabovskaya, 2001; Liangqian, Tamura, 2001).
Sect. 2. Longipetala Dorosz., 1974, Monogr. Bot. (Warszawa), 41: 159. - Sect. Eutrollius Prantl, in Engler u. Prantl, 1891, Nat. Pflanzenf., 3(2): 56, p.p. - Petals linear, flat, thin, longer than sepals or nearly equal to them (Doroszewska, 1974; Luferov, 2004).
Type: T. asiaticus L.
3. T. asiaticus L., 1753, Sp. Pl.: 557; DC., 1824, Prodr., 1: 46; C.A. Mey., 1830, in Ledeb. Fl. Alt., 1: 301; Ledeb., 1842, in Fl. Ross., 1: 73; Krylov, 1931, Fl. Zap. Sib., 5: 1121; Schipcz., 1937, in Fl. USSR, 7: 49; Gamajun., 1961, in Fl. Kazakhst., 4: 15; Pachom., 1972, in Opred. rast. Sred. Azii., 3: 144; Dorosz., 1974, Monogr. Bot., 41: 64; Grubov, 1982, in Opred. sosud. rast. Mongolii: 108; Lomon., 1984, in Opr. Rast. Tuv. ASSR: 34; Revushkin, 1988, Vys. fl. Alt.: 64; Friesen, 1993, in Fl. Sib., 6: 106; Gubanov, 1996, Konsp. fl. Vneshn. Mong.
(sosud. rast.): 51; Liangqian, Tamura, 2001, in Fl. China, 6: 141; Borodina-Grabovskaja, 2001, in Pl. Centr. Asia, 12: 19. - Trollius sertiflorus Salisb., 1807, Trans. Linn. Soc. London, 8: 303. - T. sajanensis (Malysch.) Sipliv., 1972, Novosti Sist. Vyssh. Rast., 9: 168. - T. bargusinensis Sipliv., 1973, Novosti Sist. Vyssh. Rast. ,10: 358. - T. vitalii Stepanov, 1994, Florogenet. Analiz, 1: 101, p.p. -T. kolonok Stepanov, 2018, Systematic notes, 117: 41, p.p.
Lectotype (Siplivinsky, 1972: 168): "In Sibiria" (LINN).
Description: Perennial herbs 5-90 cm high. Stem straight, simple or branched, with 1-3 flowers, grooved. Basal leaves with petioles 4-35(45) cm. Leaf blade 4.5 x 8.5 cm, base reniform, palmately 5-sect, segments rhombic, 3 -fid, fan-like lobes with clearly visible grooves between them, with unequally serrate-dentate margins (Figure 1F ). Teeth often oblong-triangular, with straight or slightly arcuate edges, pointed at the apex, less often acute (Figure 2K). Cauline leaves 2-6, similar to the basal ones, or smaller. Lower cauline leaves petiolate, upper ones sessile. Pedicle 1-10 cm, elongating to 15 cm at fruiting. Flowers 3-5 cm in diam., open (Fig. 1A, B). Sepals 10-20, orange or reddish-orange, 14-24 mm long, broadly elliptical or obovate (Figure 2L). Petals 18-40, orange or orange-red, 12-24 mm long, 2.5-4 (6) mm wide, oblong-elliptical, rounded at the apex, slightly shorter than or almost equal to the sepals (Figure 2M, N). Stamens, including anthers, 8-12 mm long. Anthers 2.5-3 mm long. Nectarostigma broadly elliptic or obovate, 2-3 mm from the petal base. Styles and stigmas yellowish. Fruit a dense globose head of numerous follicles, 8-11 mm long including the persistent style (Figures 1E, 2O). Persistent style arcuate curved inwards or erect, deflected outwards, 0.5-1 mm long. Seeds rounded, 3-edged, black, glossy.
Chromosome number: 2n = 16 (out of the AMC) (Doroczewska, 1967; Zhukova, 1967).
Hab it at: Wet meadows, forest glades, alpine and subalpine meadows in the upper and middle mountain belt., up to 2800 m.
Phenology: Flowering June - August; fruiting August - September.
Distribution: A1 (Baschelaksky Ridge), Anui River, A2 (Teletskoye Lake), A3-A6, KAD1, KAD3-KAD5, ZM1, ZM2, ZM3.
General distribution: China (Xinjiang), Kazakhstan (Almaty Region, East Kazakhstan Region), Kyrgyzstan (Chuy Region, Issyk-Kul Region, Talas Region), Mongolia (Bayan-Olgii Province, Govi-Altai Province, Zavkhan Province, Uvs Province, Khovsgol Province, Selenge Province), Russia (Altai Krai, Altai Republic, Buryatia Republic, Irkutsk Oblast, Kemerovo Oblast, Khakassia Republic, Krasnoyarsk Krai, Novosibirsk Oblast, Omsk Oblast, Tomsk Oblast, Tyumen Oblast, Tuva Republic) (Krylov, 1931; Igoshina, 1968; Siplivinskiy, 1972; Borodina-Grabovskaya, 2001; Liangqian, Tamura, 2001).
Conclusion
The study of morphological characters of the Trollius species growing in the Altai Mountain Country allowed us to propose new characters of the leaves and flowers that were not previously used for the species identification: consistence of blade and degree of its dissection, type of margin, shape of teeth, number of petal veins, shape of nectarostigma, venation of follicles, and shape and direction of persistent style. The quantitative differences of the species are specified: length of the basal leaf petiole, number of sepals and petals, distance from the petal base to the nectarostigma, and length of persistent style. The suggested key based on a combination of vegetative and generative morphological characters allows us to
identify species more accurately. As a result of the study, three species of Trollius were are identified for the Altai Mountain Country: T. altaicus, T. asiaticus, and T. dschungaricus; intermediate morphological forms, apparently of a hybrid origin, are detected.
ACKNOWLEDGEMENTS
We thank the staff of the Herbaria visited (LE, MHA, MW, NS, NSK, ALTB, TK) for opportunity to work with collections. We are grateful to V. Saveliev, A. Raspopov, V. Remorov, P. Kosachev, T. Rib, A. Ebel, A. Hamitov, and B. Kryzhatyuk for the photos of Trollius species.
Funding: The research was supported by the Scientific programme АААА-А17-117012610055-3 of the Central Siberian Botanical Garden of SB RAS, the Tomsk State University competitiveness improvement programme and the Russian Foundation for Basic Research, grant No. 18-04-00653 A.
REFERENCES
Borodina-Grabovskaya A.E. 2001. Trollius L. In: Rasteniya Tsentralnoi Asii [Plantae
Asiae Centralis]. SPb.: SPKHFA Publ., 12: 17-21 [In Russian]. Despres L., Gielly L., Redoutet B., Taberlet P. 2003. Using AFLP to resolve phylogenetic relationships in a morphologically diversified plant species complex when nuclear and chloroplast sequences fail to reveal variability. Mol. Phylogenet. Evol. 27(2): 185-196. Doroszewska A. 1967. Chromosomes of some Trollius species. Act. Polsk. Towarz. Bot. 36: 567-577.
Doroszewska A. 1974. The genus Trollius - A taxonomical study. Monogr. Bot. Pol. Akad. Nauk, 41: 1-167.
Friesen N.V. 2003. Trollius L. In: Flora of Siberia. Enfield (NH), USA / Plymouth, UK:
Science Publ. Inc. 6: 98-103. Grabovskaya-Borodina A.E. 2018. Trollius altaicus C.A. Mey. In: I.V. Sokolova, ed.-in-chief. Catalogue of the type specimens of the vascular plants from Siberia and Russian Far East kept in the Herbarium of the Komarov Botanical Institute (LE). Saint-Petersburg / Moscow: KMK Scientific Press, 2: 233 [In Russian]. Igoshina K.N. 1968. About the types of Trollius L. in the polar Urals. Botanicheskiy
zhurnal [Botanical Journal], 53(6): 779-794. [In Russian]. Kadota Y. 1987. Genus Trollius L. (Ranunculaceae). Japan. Bull. Natn. Sci. Mus. Ser. B. 13(3): 107-121.
Kamelin R. V. 2005. New flora of Altai (aims and the conception of new floristic revision).
In: Flora Altaya [Flora Altaica]. Barnaul: AzBuka, 1: 79. [In Russian]. KrylovP.N. 1931. TrolliusL. In: Flora Zapadnoi Sibiri [Flora of Western Siberia]. Tomsk.
5: 1120-1124. [In Russian]. Liangqian L., Tamura M. 2001. Trollius L. In: Flora of China. Beijing: Science Press, St.
Louis: Missouri Botanical Garden Press, 6: 137-142. Luferov A.N. 2004. A taxonomic synopsis of Ranunculaceae of the Far-East of Russia.
Turczaninowia, 7(1): 5-85 [In Russian]. Luferov A.N., Erst A.S., Luferov D.N., Shmakov A.I., Wang W. 2018. The genus Trollius
(Ranunculaceae) in the Russian Far East. Turczaninowia. 21(2): 110-116. Malyshev L.I. 1965.Vysokogornaya flora Vostochnogo Sayana [High-mountain flora of the
Eastern Sayan]. Moscow; Leningrad: Nauka Publ. 368 p. Orme C.D.L., Davies R.G., Burgess M., Eigenbrod F., Pickup N., Olson V.A., Webster A.J., Ding T.-S., Rasmussen P.C., Ridgely R.S., Stattersfield A.J., Bennett P.M.,
Blackburn T.M., Gaston K.J., Owens I.P.F. 2005. Global hotspots of species richness are not congruent with endemism or threat. Nature. 436: 1016-1019.
Rice A., Glick L., Abadi S., Einhorn M., Kopelman N.M., Salman-Minkov A., Mayzel J., Chay O., Mayrose J. 2015. The Chromosome Counts Database (CCDB) - a community resource of plant chromosome numbers. New Phytol. 206(1): 19-26.
Rostovtseva T.S. 1981. Chromosome numbers of some species of the family Ranunculaceae II. Botanicheskiy Zhurnal [Botanical Journal], 66(12): 1751-1755. [In Russian].
Schipczinsky N.V. 1937. Trollius L. In: Flora of the USSR. Moscow / Leningrad: Publishing House of the Academy of Sciences of the USSR, 7: 42-53 [In Russian].
Siplivinskyi V.N. 1972. Genus Trollius L. Asia boreali et orientali. Novosti sistematiki vysshikh rasteniy [Novitates Systematicae Plantarum Vascularum], 9: 163-182. [In Russian with Latin description].
Siplivinskyi V.N. 1973. Notula de flora Baicalensi, 1. Novosti sistematiki vysshikh rasteniy [Novitates Systematicae Plantarum Vascularum], 10: 345-361.
Stepanov N.V. 1994. Florogeneticheskiy analis (na primere severo-vostochnoi chasti Zapadnogo Sayana [Flogogenetic analysis (on the pattern of north-east part of Western Sayan)]. Krasnoyarsk: Krasnoyarsk State University. Vol. 1. 108 pp. [in Russian& Latin].
Stepanov N.V. 2018. New species of Trollius L. (Ranunculaceae) from Kuznetzk Alatau Mountains. Sistematicheskie zametki po materialam Gerbariya imeni P.N. Krylova Tomskogo gosudarstvennogo universiteta [Systematic notes on the materials of P.N. Krylov Herbarium of Tomsk State University], 117: 40-45.
Stearn W.T. 1992. Stern's dictionary of plant names for gardeners. London: Timber Press. 373 p.
Tamura M. 1995. Trollius L. In: Hiepko P. (Ed.) Die Natürlichen Pflanzenfamilien. Berlin: Duncker und Humblot. 17(4): 238-244.
Wang W., Hu H., Xiang X.-G., Yu S.-X., Chen Z.-D. 2010. Phylogenetic placements of Calathodes and Megaleranthis (Ranunculaceae): Evidence from molecular and morphological data. Taxon, 59(6): 1712-1720.
Zhukova P.G. 1967. Karyology of some plants, cultivated in the Arctic-Alpine Botanical Garden. In: N.A. Avrorin (ed.) Plantarum in Zonam Polarem Transportatio. Leningrad. 2: 130-149. [In Russian].
Received 25 November 2018 Accepted 20 December 2018
Citation: Erst A.S., Luferov A.N., Troschkina V.I., Xiang K., Wang W. The genus Trollius L.
(Ranunculaceae) in the flora of the Altai Mountain Country. Sistematicheskie zametki po materialam
Gerbariya imeni P.N. Krylova Tomskogo gosudarstvennogo universiteta [Systematic notes on the
materials of P.N. Krylov Herbarium of Tomsk State University]. No 118. P. 64-77. DOI:
10.17223/20764103.118.5
Систематические заметки ..., 2018. № 118. С. 64-76 DOI: 10.17223/20764103.118.5
Род Trollius L. (Ranunculaceae) во флоре Алтайской
горной страны
А.С. Эрст12, А.Н. Луферов3, В.И. Трошкина1, К. Ксианг4,5,
В. Ванг4,5
Центральный Сибирский ботанический сад СО РАН, Новосибирск, Россия; erst_andrew@yahoo. com
2Томский государственный университет, Томск, Россия
3Первый Московский государственный медицинский университет им. И.М. Сеченова, Москва, Россия; [email protected] 4Институт Ботаники Китайской академии наук, Пекин, Китай 5Университет Китайской академии наук, Пекин, Китай.
Аннотация. В статье представлены сведения о новых признаках, которые ранее не использовались для идентификации видов Trollius: текстура и степень рассечения пластинки, зубчатость и форма зубцов, число жилок лепестка, форма нектаростигмы, морфологические признаки плодов - тип поверхности околоплодника (наличие резко выступающих жилок у T. altaicus; слегка выступающие жилки у T. asiaticus, гладкая поверхность у T. dschungaricus). Указаны количественные морфологические характеристики видов: число чашелистиков и лепестков, расстояние от основания лепестка до нектаростигмы, длина листовок, их форма и расположение. Предложен идентификационный ключ, разработанный с учетом предложенных признаков, приведен конспект рода Trollius (Ranunculaceae) Алтайской горной страны, включающий 3 вида, относящиеся к 2 секциям. Конспект включает номенклатурные цитаты, информацию о типовых образцах, распространении в регионе и об общем распространении каждого вида.
Ключевые слова: Trollius, Ranunculaceae, диагностические признаки, Алтайская горная страна.
Funding: Исследования поддержаны научной программой АААА-А17-117012610055-3 ЦСБС СО РАН, Программой повышения конкурентоспособности Томского государственного университета и РФФИ (грант № 18-04-00653 A).
Поступила в редакцию 25.11.2018 Принята к публикации 20.12.2018
Цитирование: Erst A.S., Luferov A.N., Troschkina V.I., Xiang K., Wang W. The genus Trollius L. (Ranunculaceae) in the flora of the Altai Mountain Country // Систематические заметки по материалам Гербария им. П.Н. Крылова Томского государственного университета. 2018. № 118. С. 64-76. DOI: 10.17223/20764103.118.5
