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36
УДК 582.998(571.5/6)
P.G. Gorovoy П.Г. Горовой
THE GENUS RHAPONTICUM VAIL. (ASTERACEAE: CARDUEAE - CENTAUREINAE)
IN SIBERIA AND FAR EAST
Род RHApoNTIcuM VAIL. (ASTERACEAE: CARDUEAE - CENTAUREINAE)
в сибири и на дальнем востоке
Summary. A taxonomy of the Asteraceae in Asia the genus Rhaponticum Vail. (Leuzea DC., Stemmacantha Cass.) is still a taxon for discussions. The species Rh. carthamoides (Willd.) Iljin in East Siberia unites 3 subspecies: carthamoides, chamarensis (Peschkova) Zhirova and orientate (Serg.) Soskov; although they differ in morphology, they do not have isolated areas. In 1959 Yu.D. Soskov described Rhaponticum satzyperovii and pointed to its affinity to Rh. uniflorum (L.) DC. The plant height, stem indumentum degree, radical and stem leaf dissection degree to be diagnostic characters. Studies on living plants of Rh. satzyperovii and herbarium (LE, MHA, MW, VLA) examination have shown that diagnostic characters are not consistent. Yu.D. Soskov claims the species area to be an argument of Rh. satzyperovii being a distinct species, whose area covers the south of the Primorye Province in the Far East of Russia with some locations in the Jewish Autonomous Region and in North-East China. He reported a disjunction in the area of Rh. uniflora. In the course of specification of growth locations of Rh. uniflora in Russia, Mongolia, China and Korea, the area turned out to have no disjunction. The area continuity takes place due to the species’s growing in North-East China. The area of Rh. satzyperovii found itself «within» the area of St. uniflora. Thus, on the territory East Asia, Rh. uniflora occurs that is characterized with wide range of morphological variability and the area covering the territories of East Siberia, Mongolia, Central, North and North-East China, the Primorye Territory, the south of the Amur Region and Korean Peninsula. Rh. satzyperovii has been synonymized with Rh. uniflora.
Key words: Rhaponticum, Siberia, Far East, distribution.
Аннотация. Среди сложноцветных (Asteraceae) Азии, род Rhaponticum Vail. (Leuzea DC., Stemmacantha Cass.) остаётся одним из наиболее дискуссионных. Вид Rh. carthamoides (Willd.) Iljin в Восточной Сибири объединяет три подвида: carthamoides, chamarensis (Peschkova) Zhirova и orientate (Serg.) Soskov. Несмотряна некоторые морфологические различия, эти таксоны не являются полностью аллопатричными. В 1959 г. Ю.Д. Сосков описал Rh. satzyperovii, родственный Rh. uniflorum (L.) DC. и отличающийся от последнего размерами растений, интенсивностью опушения стебля, а также степенью рассечения прикорневых и стеблевых листьев. Изучение живых растений Rh. satzyperovii и гербарных коллекций (LE, MHA, MW, VLA) по данному таксону выявило непостоянство диагностических признаков. Одним из аргументов в пользу самостоятельности Rh. satzyperovii, по мнению Ю.Д. Соскова, является его ареал, охватывающий юг российского Дальнего Востока (юг Приморского края, несколько местонахождений также в Еврейской автономной области) и Северо-Восточный Китай, и не перекрывающийся с ареалом Rh. uniflorum. В процессе уточнения распространения Rh. uniflorum в России, Монголии, Китае и Корее, выяснилось, что ареал вида не является дизъюнктивным, а простирается сплошной полосой от Саян, Прибайкалья и Северной Монголии через Северо -Восточный Китай до Кореи и Дальнего Востока. Таким образом, ареал Rh. satzyperovii оказывается полностью включённым в область распространения St. uniflorum. Следовательно, на территории Восточной Азии произрастает один вид - Rh. uniflorum, характеризующая значительным диапазоном морфологической изменчивости и встречающийся в Восточной Сибири, Монголии, Центральном, Северном и Северо-Восточном Китае, в Приморье, на юге Амурской области и на п-ве Корея. Название Rh. satzyperovii следует отнести к синонимам Rh. uniflorum.
Ключевые слова: Rhaponticum, Сибирь, Дальний Восток, распространение.
INTRODUCTION literature covering the studies on chemical composi-
The names Rhaponticum Vail. (Rhaponticum tion of the plant species of this genus.
Hill, Rhaponticum Ludw.), Leuzea DC. and Stem- The authorship of the name Rhaponticum is
macantha Cass. are used in floristic reports and in ascribed to M. Adanson (1763). However, this name
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Тихоокеанский институт биоорганической химии ДВО РАН (ТИБОХ ДВО РАН), пр-т 100 лет Владивостоку, 159; 690022, Владивосток, Россия; e-mail: gorovoy@piboc.ru
Pacific Institute of Bioorganic Chemistry, Far East Branch, Russian Academy of Sciences (PIBOC FEB RAS) 159 Pr-t 100 let Vladivostoku; 690022, Vladivostok, Russia
Поступило в редакцию 18.10.2010 г.
Submitted 18.10.2010
was used as early as H. Bock (1572), A. Prosperus (1612), G. Bauchin (1623) and other authors (Ditrich, 1973; Holub , 1973).
Many taxonomists developed the system of the genus Rhaponticum. In 1718, M. Vaillant used this name for a group of plants including species of the genera Leuzea and Acroptilon Cass., according to the contemporary conceptions. In 1742, A. Hauler used the name Rhaponticum (Holub, 1973) for the group of four species of the Asteraceae. Ludwig (1747) further expanded the concept of Rhaponticum (9 species) and later he (Ludwig, 1757) validated this generic name. K. Linneus (1763) developed the system of Rhaponticum, but he put many species of Centaurea L. into this genus. In 1779, J.B. Lamarck excluded C. rhapontica L. from the genus Centaurea and transferred it into the genus Rhaponticum as Rh. scariosum Lam. Later on, a number of authors clarified and changed the size of the genus Rhaponticum (Jussieu, 1789; De Candolle, 1837; Ledebour, 1845; Bentham, Hooker, 1876; Engler, Prantl, 1893), they put species of the genera Cnicus L., Serratula L., Cirsium Mill., and Centaurea into this genus. The genus name Rhapon-tica was suggested by J. Hill (1762) and the IAPT (International Association of Plant Taxonomy) resolved to consider this genus name to be the latest homonym of Rhaponticum (Dittrich, 1984). M.M. Ilyin (1933), Yu.D. Soskov (1959), M. Dittrich (1968, 1973, 1984), J. Holub (1973, 1974) studied the characters distinguishing Rhaponticum from closely related Leuzea and found out that these taxa differ in structure of achene surface, pappus structure and inflorescence form.
The genus Stemmacantha was described by
H. Cassini in 1817 and Serratula cynaroides DC. was chosen to be the type species. H. Cassini considered special structure of calyx bracts to be the distinguishing character of the genus Stemmacan-tha. Later, A.P. De Candolle (1837) turned the genus Stemmacantha into the section of Rhaponti-cum. After the genera Stemmacantha and Leuzea were recognized as separate species in 1978, the IAPT made a decision to move species and subspecies from the Rhaponticum into the genus Stemma-cantha (Dittrich, 1984). The system of the genus Stemmacantha proposed by M. Dittrich (1984) was adopted by S.K. Czerepanov in the check-list «Vascular plants of Russia and adjacent states» (1995) and it includes 20 species, 10 subspecies and 2 varieties. In the book «Systematics, evolution, and biogeography of Compositae» (Funk et el., 2009), the genus Stemmacantha is absent.
MATERIALS AND METHODS
The paper is based on the studies on living plants and herbarium specimens on Rh. uniflorum (L.) DC. and Rh. satzyperovii Sosk. collected during expeditions in nature populations in the Irkutskaya, Chitinskaya, Amurskaya and Jewish Autonomous Regions, Khabarovsky and Primorsky Territories; herbarium material (LE, MHA, MW, VLA) and Pacific Institute of Bioorganic Chemistry (Vladivostok) the Far-Eastern Branch of the Russian Academy of Sciences (FEB RAS), the Botanic Garden-Institute FEB RAS (Vladivostok), the Institute of aquatic and ecological problems FEB RAS (Khabarovsk), the Institute of Complex Analysis of Regional Problems FEB RAS (Birobidzhan), the Zabaikalsky State Pedagogical University ofN.G. Chernyshevsky (Chita), the Blagoveschensk State Pedagogical University (Blagoveschensk) and literature.
RESULTS AND DISCUSSION
In the literature on flora of Siberia, the Russian Far East (RFE), Mongolia, Korea and China and in publications on taxonomy of the Asteraceae there is no agreement regarding the limits of the genus Rhaponticum (Stemmacantha). In the check-list of S.K. Czerepanov (1995) there are three species of Stemmacantha - St. carthamoides (Willd.) Dittrich, St. uniflora (L.) Dittrich and St. satzyperovii (Sosk.) Czer. reported for East Asia. Rhaponticum satzyperovii described in 1959 by Yu.D. Soskov based on the collections of 1913 from Primorsky Territory, M. Dittrich (1984) includes into Rhaponticum as a subspecies St. uniflora subsp. satzyperovii (Sosk.) Dittrich.
Rhaponticum carthamoides (Willd.) Iljin is distributed on subalpine meadows of Central and East Siberia and outside Russia it grows only in two locations in Mongolia (the Mongolian Altai) (Zhirova, 1997; Doronkin, 2003, Grubov, 1982) and is represented by the subspecies: carthamoides, orientale (Serg.) Soskov, chamarensis (Peschkova) Zhirova, that differ in morphological characters (shape of appendages of outer involucre leaflets), but do not have isolated areas. No more taxa of Rh. carthamoides affinity were described from Siberia. In the southeastern part of Siberia the area of Rh. carthamoides reaches the area of the southern lakeside of Baikal (Zhirova, 1997). Rhaponticum carthamoides is used in Russia as a medicinal plant (Leuzea the strong root) and as a source for ecdysteroid production. Today mass harvesting of these species accounts for working out measures on its conservation. The cited literature and synonyms are provided below.
Rhaponticum carthamoides (Willd.) Iljin, 1933, Act. Inst. Bot. Acad. Sci. USSR, ser. 1, 1 :
204; Soskov, 1964, Fl. URSS, 28: 311; Zemlinsky,
1958, Med. plants USSR: 359; Polozhyi, Postni-cov, Surov, 1980, Atlas areals a. resources med. pl. USSR : 185 (text, ic.), 103 (map); Zhirova, 1997, Fl. Sibiria, 13 : 229, maps 162-164. - Cnicus cartham-oides Willd. 1803, Sp. Pl. 3, 3 : 1685. - Stemma-cantha carthamoides (Willd.) M. Dittrich, 1984, Candollea, 39, 1 : 46; Sh. Chu, 1987, Fl. Reipubl. Popul. Sinicae, 78, 1 : 185; Czer. 1995, Sosud. rast. Rossii i sopr. gosud.: 195; Doronkin, 2003, Fl. Sibiria, 14 : 95. - C. centauroides Willd. 1803, l. c.,p. p. quoad pl. Sibir. - C. uniformis Sievers, 1796, in Pall. Neue nord. Beitr. 3 : 346, nomen nudum. -Serratula carthamoides Poir. 1804, in Lam. Encycl. meth. 6 : 561. - S. cynarifolia Poir. 1804, 1. c., p. p. quoad pl. Sibir. - Leuzea carthamoides DC. 1810, Ann. Mus. Paris, 16 : 205; id., 1837, Prodr. 6 : 666; Ledeb. 1845-1846, Fl. Ross. 2, 2 : 753; Turcz. 1856, Fl. Baic.-Dahur. 2, 1 : 135; Kryl. 1949, Fl. West Sibiria, 11 : 2943. - L. altaica Fisch. ex Schauer, 1834, Cat. Sem. hort. Vratisl.; id. 1835, Linnaea, 10 : 117, non Link, 1822. - Cirsium carthamoides Link, 1822, Enum. Pl. Hort. Berol. 2 : 303. - Halocharis carthamoides M. Bieb. ex DC. 1837, Prodr. 6 : 666, pro syn. - Centaurea carthamoides Benth. 1873, in Benth. et Hook. f., Gen. pl. 2 : 479.
Rhaponticum uniflorum (L.) DC. is reported in «Key-book of plants of Primorye and Pria-murye» (Vorobyev et al., 1966) and Rh. satzyperovii is the synonym of this species. V.N. Woroschilow (1966) reported two species of Rhaponticum in the Russian Far East (FE): Rh. uniflorum, Rh. satzyperovii and noted that in the Primorye Territory there are plants both similar to typical specimens of Rh. uni-florum and with characters transitional between Rh. satzyperovii and Rh. uniflorum. Later, V.N. Woroschilow (1982, 1985) treatyed Rh. satzyperovii as Rh. uniflorum subsp. satzyperovii (Sosk.) Worosch. In the survey «Vascular plants of the Soviet Far East» (Barkalov, 1992) there are two independent species: Rh. uniflorum and Rh. satzyperovii.
In the floristic surveys on Siberia, M.G. Popov (1959), G.A. Peshkova (1979) and O.S. Zhirova (1997) report Rh. carthamoides and Rh. uniflorum for the territory of Sibiria. In the «Key-book on vascular plants of Mongolia» there is only Leuzea uniflora (L.) Holub (Grubov, 1982). M. Kitagawa (1979) reports Rh. uniflorum for Siberia, Mongolia, Amursky and Ussuriysky Regions, Korea and China. In the surveys on flora of Korea (T. Lee, 1993; Y. Lee, 1996) there is only Rh. uniflorum as well. Rh. carthamoides and Rh. uniflorum are reported
Fig. 1. Distribution of Rhaponticum carthamoides.
Fig. 2. Distribution of Rhaponticum uniflorum: • - Rh. uniflorum, m - Rh. satzyperovii (synonym).
(Chu, 1987) for China, however Rh. satzyperovii is treated as synonym of Rh. uniflorum.
There is a special research in order to ascertain whether Rh. satzyperovii is an independent species. The species description (Soskov, 1959) in «The Flora of the USSR» (Soskov, 1963) says, that Rh. satzyperovii is close to Rh. uniflorum, but «... it has isolated area and is well distinguished by its robust stem up to 1 m high, lyrate radical leaves and large superficially partite, often pinnatilobate, stem leaves and a number of other characters...».
The studies on living plants of Rh. satzypero-vii, as well as observation of herbarium materials have shown that the diagnostic characters, indicated by Yu.D. Soskov to be distinguishing, are not consistent. Having observed more than 300 plants of Rh. satzyperovii in «locus classicus» we were able to collect some specimens that corresponded to Rh. uniflorum in their morphological characters. There is high variability in plants of both species. The most variable are the shape and dissection of radical and lower stem leaves, i. e. the characters that were considered to be distinguishing while describing Rh. satzyperovii. Within a single population of Rh. satzyperovii there are plants with lyrate radical leaves with a large terminal lobe and slightly partite
lobed stem leaves, as well as individuals with pin-natipartite or pinnatifid leaf blades without a large terminal lobe. Often in the same leaf rosette there are leaves of two forms with all transitions between them. Such pattern is typical to both Rh. satzyperovii from «locus classicus» and the plants in the northernmost location of the area (by Soskov Yu.D.) of Rh. satzyperovii in the Jewish Autonomous Territory in the vicinities of Bidzhan Settlement. Such pattern in leaf variability is observed in plants of Rh. uniflorum. The species characters describing Rh. satzyperovii are easily fit into a number of character variability of Rh. uniflorum. There are not any differences between Rh. uniflorum and Rh. satzyperovii in other morphological characters.
Within the species one can find plants with xeromorphic pattern: dwarfish, with small dissected (often as deep as to the central costa) leaf blades with up to 12 pairs of narrow-lanceolate lobules. Usually those are plants of open southern steppe or stony hill slopes. Xeromesophytic plants are tall (up to 1 m high), with large, pinnatilobate or slightly partite into 3-8 pairs of wide ovate-oblong lobules, lyrate leaves - they often occur on elevated sites of river valleys, sea shores, sometimes under forest canopy. Plants with xeromorphous pattern grow in the
north-western part of the area in the regions with continental, arid climate (the Tuva and Buryatiya Republics, the Irkutskaya, Chitinskaya and the north of Amurskaya Regions, Mongolia, North and Central China). Plants with xeromesophytic pattern restricted to the regions with humid, warm, monsoon climate (the southern areas of the Amurskaya Region, the Jewish Autonomous Region, Primorsky Territory, North-East China and Korea Peninsula). The area of Rh. satzyperovii (Soskov, 1956) covers the territory of south Primorsky Territory of the RFE, the Jewish Autonomous Region (Stolbovoye Village and Bidzhan Settlement) and North-East China (near Sochintsy, Matsyaokhe Station). In the area of Rh. uniflorum, Yu.D. Soskov reports a significant disjunction from the northern part of Zeya-Bureya floristic region (in the Amurskaya Reion) and «...as an introduced plant on the Muraviov-Amursky Peninsula, Putyatin Island and in the vicinities of Ussuriysk city ...». While specifying the growth locations of Rh. uniflorum in Russia, Mongolia, China and Korea we ascertained that this plant does not occur in the Russian Far East and Siberia as an introduced plant and the area of the species turned out to be continuous (with no disjunction). The area continuity of Rh. uniflorum provided by this species growing in North-East China (Manshuria). The area of Rh. satzyperovii found itself «within» the area of Rh. uniflorum.
Morphometric studies on carpological character variability in Rh. satzyperovii and Rh. uni-florum (Basargin, Vorobyeva, 2004), morphology and anatomy of achenes, as well as stomatographic research did not reveal any specific difference (Za-rembo, 2000).
In East Siberia, Mongolia, Central, North and North-East China, in Primorsky and the south of Amursky Terrytories, in Korea Peninsula there is Rh. uniflorum that is characterized with a wide range of morphological variability, and therefore Rh. satzyperovii schould be reduced to the synonyms of Rh. uniflorum. Detailed nomenclature of Rh. uniflo-rum is given below.
Rhaponticum uniflorum (L.) DC. 1810, Ann. Mus. Paris, 16 : 189; id., 1837, Prodr. 6 : 664; Ledeb. 1845-1846, Fl. Ross. 2 : 751; Turcz. 1856, Fl. Baicdahur. 2 : 133; Franch. 1883, Nouv. Arch. Mus. Hist. Nat. Paris, 6 : 62; id., 1884, Pl. David. 1 : 183; Popov,
1959, Fl. Mid. Sib. 2 : 866; Soskov, 1963, Fl. SSSR, 28 : 318; Woroschilow, 1966, Fl. Sov. Far East : 436; Vorobyev, 1966, Key-book of plants Prim. and Pria-mur. : 428; S.Y. Hu, 1967, 1. c. 20, 3-4 : 310; Pesh-kova, 1979, Fl. Central Sib., 2 : 890; Kitag. 1979, Neo-Lineam. Fl. Mansh. : 666; Czer. 1981, Vascular
plants USSR : 92; Woroschilow, 1982, Key-book of plants Sov. Far East : 583; H.Ch. Fu, 1982, Fl. Intra-mong. 6 : 211; Woroschilow, 1985, Florist. issled. v raznykh raionakh SSSR : 198; Barkalov, 1992, Vascular plants Sov. Far East, 6 : 314; Y.N. Lee, 1996, Fl. Korea : 858; Zhirova, 1997, Fl. Sibiria, 13 : 229. - Cnicus uniflorus L. 1771, Mant. Altera. : 572; Georgi, 1797, Reise, 3 : 1225; Willd. 1804, Car. Lin. Sp. Pl., 3 : 1685. - Centaurea monanthos Georgi, 1775, Reise, 1 : 231; Forbes et Hemsl. 1888, Journ. Linn. Soc. Bot. 23 : 470; Palib. 1898, Consp. Fl. Kor. : 120; Komarov, 1907, Fl. Manchuria, 3 : 761; Nakai, 1911, Fl. Korea, 2 : 48; id., 1923, Fl. Sylv. Koreana, 14 : 107; Komarov and Alisova, 1932, Key-book of plants of the Far-East. Province, 2 : 1086; Chen, 1934, Bull. Mem. Inst. Biol. Bot. 5 : 94; Ling, 1935, Contr. Inst. Bot. Nat. Acad. Peip. 3(4) : 178; S.Y. Hu, 1966, Quart. Journ. Taiwan Mus. 19, (1-2) : 21. - С grandiflora Pall. 1776, Reise, 3 : 237, 321; Kitag. 1936, Index Fl. Jehol. : 54. - С. membranacea Lam. 1783, Encycl. Meth. 1 : 666; S.Y. Hu, 1966, 1. c. : 20. - Serratula uniflora (L.) Spreng. 1826, Syst. Veg. 3 : 388. - Leuzea daurica Bunge, 1833, Enum. PI. Chin. : 37; id., 1835, Mem. Acad. Sc. St. Petersb. Sav. Etrag. 2 : 111. - Leuzea uniflorum (L.) Holub, 1973, Folia Geobot. Phylotax. (Praha), 8 : 392; Grubov, 1982, Key for vascular plants of Mongolia : 262. - Leuzea satzyperovii (Sosk.) Holub, 1973,
1. c. : 392. - Rhapontica uniflora DC. 1837, Diss. Соmр. : 33; Maxim. 1859, Prim. Fl. Amur. : 176; Korsh. 1892, Acta Hort. Petrop. 12 : 361; Diels in Engler, 1905, Bot. Jahrb. Beibl. : 108; Kitam. 1937, Mem. Coll. Sci. Kyoto Univ., ser. В. 13 : 30; Kitag. 1939, Lineam. Fl. Mansh. : 461; Nakai, 1952, Bull. Nat. Sc. Mus. Tokyo, 31 : 118; Т.В. Lee, 1993, Illust. Fl. Korea : 778. - Stemmacantha uniflora (L.) Dittrich, 1984, Candollea, 39, 1 : 49; Sh. Chu, 1987, Fl. Reipubl. Popul. Sinicae 78, 1 : 184; Czer. 1995, Sosud. rast. Rossii i sopr. gosud. : 195, Doronkin, 2003, Fl. Sibiria, 14 : 95. - Rhaponticum dahuri-cum (Bunge) Turcz. 1838, Bull. Soc. Nat. Mosc. 11 : 95. - Rh. monanthum (Georgi) Worosch. 1953, Seed list of Main Bot. Garden Acad. Sci. USSR : 8. - Rh. satzyperovii Sosk. 1959, Bot. mat. Gerb. Bot. In. AN SSSR, 1963, 19 : 400; l. c. : 317; Worosch. 1966, l. c. : 436; Czer. 1981, l. c. : 92; Barkalov, 1992, l. c. : 314. - Rh. uniflorum subsp. satzyperovii (Sosk.) Worosch. 1982, l. c. : 583; 1985, l. c. : 198. -Stemmacantha satzyperovii (Sosk.) Czer. 1995, l. c. : 195. - St. uniflora subsp. satzyperovii (Sosk.) Dittrich, 1984, l. c. : 49. - Icon.: Gmel. 1749, Fl. Sibirica, 2 : tab. 38.
ACKNOWLEDGEMENTS This research was supported by the inte-
Author thanks to A.N. Vorobyeva for ob- gration grant of Far Eastern Branch and Siberian
servation the herbarium specimens in Chita, Bla- Branch Russian Academy of Sciences (09-11-SD-
goveschensk and Birobidzhan cites. 06-004)
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