FLORA OF THE USSR Volume XXVIII COMPOSITAE Tribes Cynareae and Mutisieae saslaitall bare oarrnayD rodinT ‘ FLORA OF THE USSR Initiated under the supervision and chief editorship of Academician V.L. Komarov VOLUME XXVIII COMPOSITAE Tribes Cynareae and Mutisieae Volume Editors E.G. BOBROV AND S.K. CZEREPANOV General Scientific Editors Stanwyn G. Shetler Galina N. Fet Ellen A. Unumb Translated from Russian Smithsonian Institution Libraries Washington, D.C. 2001 SMIN B 86-003 Flora SSSR. Tom XXVIII Akademiya Nauk SSSR Publishers Moscow-Leningrad, 1963 © 2001 Amerind Publishing Co. Pvt. Ltd., New Delhi Translator: Dr. A.K. Dhote General Editor: Dr. V.S. Kothekar Library of Congress Cataloging-in-Publication Data Flora SSSR. English Flora of the U.S.S.R. = Flora SSSR. Cover title: Flora U.S.S.R. Translation of: Flora SSSR. At head of title, v. -28: V.L. Komarov Botanical Institute. Academy of Sciences of the USSR. Reprint. Originally published: Leningrad: Izdatel’stvo Akademi nauk SSSR, 1934— Vols. Have imprint: Washington, D.C. : Smithsonian Institution Libraries and the National Science Foundation. Includes bibliographies and indexes. 1. Botany—Soviet Union—Classification. 2. Plants—Identification. I. Komarov, V.L. (Vladimir Leont’evich), 1869-1945. II. Botanichesku institut im. V.L. Komarova. III. Title: Flora SSSR. III. Title: Flora U.S.S.R. V. Title. K321.F69613 1985 85-904577 Translated and published under an agreement with the Smithsonian Institution Libraries, Washington, D.C., by Amerind Publishing Co. Pvt. Ltd., 66 Janpath, New Delhi 110001 Typeset at Radiant Printers, New Delhi 110 020 and Printed at Baba Barkha Nath Printers, 26/7 Najafgarh Road Industrial Area, New Delhi 110 015 INSTITUTUM BOTANICUM nomine V.L. KOMAROVII ACADEMIAE SCIENTIARUM URSS FLORA URSS (FLORA UNIONIS RERUMPUBLICARUM SOCIALISTICARUM SOVIETICARUM) AXVITT EDITIO ACADEMIAE SCIENTIARUM URSS MOSQUA 1963 LENINGRAD V.L. KOMAROV BOTANICAL INSTITUTE ACADEMY OF SCIENCES OF THE USSR Contributors A.G. Borissova, M.M. Iljin, M.V. Klokov, I.A. Linczevsky, E.G. Pobedimova, G.L. Semidel, Yu.D. Soskov, D.I. Sosnowsky, S.G. Tamamschian, A.L. Charadze, N.N. Tzvelev, S.K. Czerepanov, S.A. Shostakovsky SCIENTIFIC EDITORS’ PREFACE TO VOLUME XXVIII This is the fourth volume of the six (Vols. XXV—XXX) that treat the large family Compositae (Asteraceae). We have, as a practical matter, concentrated our editorial review of this volume on the keys, discussions, and the habitat and distribution statements. We are confident of the general accuracy of the translation but also recognize that there will be imperfec- tions. Throughout this volume, as in the other volumes on this family, the translators have used the Latin term “capitulum” (pl., capitula), instead of the more familiar English term, “head,” for the Russian term denoting the primary inflorescence of florets. “Head,” if it is used, describes a second- ary inflorescence, i.e., an arrangement of capitula. December 2000 STANWYN G. SHETLER Curator of Botany Emeritus National Museum of Natural History Smithsonian Institution Washington, DC 20560 Assisted by Galina N. Fet Vrmovulvi : ireini ri ANN Huntington, West Virginia = “\\ ob Ellen A. Unumb 1 135 2004 JAN Bethesda, Maryland LibKAME ¥ 4, a ¥ : 4 wal 4 Bow ~~ we piel te ea oe gasrane eet WY WMATE io tabhaloateni Mie : 00s ons A av 2eets pid as b. Husker, DA mie ‘ oy 4 SARE TETS) ' pas ae ape ET Charts, NN. ‘Crveley, SM Ceeropangy, |! wnotae 26W B.A Shiwakersky 19 Mi anileD yd betsiexA siaigiY teoW sotgoiinel dowel .A weila bnehosM aheodtsél Iv* PREFACE The twenty-eighth volume continues and concludes the treatment of materials on the tribe Cynareae. It also includes the small tribe Mutisieae. The volume covers 42 genera, which include 452 species. Centaurea is the largest genus; its 178 species are distributed in 22 subgenera. At present, the Editorial Board finds it more convenient, pend- ing a complete revision, to retain basically the traditional circumscription of the genus. This is because the subgenera considered in this volume are more or less equivalent and could be raised to the rank of separate genera. Cirsium is the second largest genus (111 species); a fresh look at the material of this genus allows us to describe several new species ay to establish a number of new subdivisions. A number of papers have been published in connection with the inves- tigations for the present volume; nevertheless, descriptions of 49 new spe- cies, sections, and subgenera have been included in the Addendum. Unlike in the earlier volumes, in the present volume the dates of actual publication of the different parts have been indicated when C.F. Ledebour’s Flora Rossica is cited. Editorial Board *Page number of the Russian original—General Editor. boa Lar | nad ¥ t he) cle Sb, Hep: j ; al | Deve ror ean te Cas » ¢d i i) " hs iG a yi : ad if Dra xP on a J ain ai wie: Obs i ahoieinudh seit linens sethweskitoni ote 1 sekmaay® wis ont m0. obnins Baie 2: forsee any gases EE ni hotinih ow eisogy HT wk eg tna et wos, pos eg lcleckinnones 4 ascii connen sets box adielans ea ( ara beat tec sind Hi hdanma rel Noctanal 6 ao : et. fpem eskangs inc: Sunivee adtvonate: ot cs eorotls apy silo Uateebante cs netpiery tvetece Wrpdinesprevnemire i , seoval dk tain enleencinaneh tailiiidee med wakemcpn Yount 4 , cay siae Ch ence Dadra saniies ace a ia ruieobbA of iti bobuloni ood oved procoydue brs enoioan yay \ lewis jo cxeh ode-sruloy inant of al asmulov raitues oft mi olla am 4 a HKrinbs fa eaumammaa arian mer chee ee Jbatio ai wales ae bino8 isinoibsl . ao RPO Lomb Mwnaveaie tenigive aaieaa® avi Yr ipa aged 654* S.K. Czerepanov S.G. Tamamschian M.M. Ijin and G.L. Semidel M.M. Ijin A.L. Charadze I.A. Linczevsky A.G. Borissova Yu.D. Soskov N.N. Tzvelev D.I. Sosnowsky CONTRIBUTORS Characteristics of the tribe and subtribes; Keys to the genera of the subtribe Carduinae and subtribe Centaureinae; Genera Notobasis, Zoegea, Russowia, Crupina, Plagiobasis, Chartolepis, Callicephalus, Acroptilon, Oligochaeta, Schischkinia, Cheiro- lepis, Tomanthea, Centaurea (Characteristics, Key to the subgenera, and subgenera Calcitrapa, Cyanus, Czerniakovskya, Jacea,_ Seridia, Solstitiaria, Sosnovskya, Tetramorphaea). Genera Carduus, Picnomon, Lamyra, Lamyropappus, Ancathia, Modestia, Cynara, Silybum, Onopordum. Genera Alfredia, Syreitschikovia. Genera Olgaea, Cnicus. Genus Cirsium. Genus Tricholepis. Genera Serratula, Synurus, Schumeria. Genus Rhaponticum. Genera Amberboa, Stizolopus, Hyalea, Centaurea (subgenera Amblyopogon, Centaurea, Hyalinella, Lopholoma, Microlophus, Odontolophopsis, Odontolophus, Pseudohyalea, Rhizocalathium, Xanthopsis), Phalacrachena. Genera Grossheimia, Aetheopappus, Centaurea (subgenus Psephellus). *Page number of the Russian original—General Editor. xl M.V. Klokov Genus Centaurea (subgenera Acrolophus, Heterolophus, Phalolepis). S.A. Shostakovsky Genus Carthamus. E.G. Pobedimova Key to the genera of the subtribe Mutisinae; gen- era Gerbera, Leibnitzia, Uechtritzia. Addenda XXVII—Diagnoses Plantarum Novarum in Tomo XXVIII Flore URSS Commemoratum, Additamentum. PLATES T.N. Shishlova Plates I-XII, XIV—-XVIII, XXII, XXV—XXVIII. O.P. Yakovleva Plates XIII, XIX, XXI, XXIII, XXIV. Z.V. Kobyletskaya Plate XX. CONTENTS SCIENTIFIC EDITORS’ PREFACE TO VOLUME XXVIII PREFACE CONTRIBUTORS SYSTEMATIC INDEX OF THE SPECIES OF THE TWENTY-EIGHTH VOLUME OF THE “FLORA OF THE USSR” KEY TO GENERA OF CARDUINAE Genus 1589. Genus 1590. Genus 1591. Genus 1592. Genus 1593. Genus 1594. Genus 1595. Genus 1596. Genus 1597. Genus 1598. Genus 1599. Genus 1600. Genus 1601. Subtribe 3. Centaureinae O. Hoffm. Genus 1602. Genus 1603. Genus 1604. Genus 1605. Genus 1606. Genus 1607. Genus 1608. Genus 1609. Genus 1610. Carduus L. emend. Gaertn. Alfredia Cass. Olgaea Ijin. Cirsium Mill. emend. Scop. Notobasis Cass. Picnomon Adans. Lamyra Cass. Lamyropappus Knorr. and Tamamsch. Ancathia DC. Modestia Charadze and Tamamsch. Cynara L. Silybum Adans. Onopordum L. Zoegea L. Russowia Wink. Crupina Cass. Plagiobasis Schrenk Tricholepis DC. Serratula L. Synurus Ijin Syreitschikovia Pavl. Schumeria jin X1V Genus 1611. Genus 1612. Genus 1613. Genus 1614. Genus 1615. Genus 1616. Genus 1617. Genus 1618. Genus 1619. Genus 1620. Genus 1621. Genus 1622. Genus 1623. Genus 1624. Genus 1625. Genus 1626. Genus 1627. Rhaponticum Adans. Amberboa (Pers.) Less. Grossheimia Sosn. and Takht. Chartolepis Cass. Callicephalus C.A.M. Acroptilon Cass. Oligochaeta C. Koch. Schischkinia (jin Aetheopappus Cass. Cheirolepis Boiss. Tomanthea DC. Stizolophus Cass. Hyalea (DC.) Jaub. and Spach Centaurea L. Phalacrachena Ijin Carthamus L. Cnicus L. TRIBE 13. Mutisieae Cass. Subtribe 1. Mutisinae O. Hoffm. Genus 1628. Genus 1629. Genus 1630. ADDENDA Gerbera Cass. Leibnitzia Cass. Uechtritzia Freyn INDEX ALPHABETICUS 305 320 330 332 337) 340 344 349 350 354 ao7 362 365 368 S07) 580 586 587 588 588 590 594 598 625 vy SYSTEMATIC INDEX OF THE SPECIES OF THE TWENTY- EIGHTH VOLUME OF THE “FLORA OF THE USSR”* Tribe 12. Cynareae Less. Subtribe 2. Carduinae O. Hoffm. Genus 1589. Carduus L. emend. Gaertn. Section 1. Carduus 15688. 1. C. nutans L. Ae 10 2. C. songoricus Tamamsch. A 11 15690 3. C. schischkinii Tamamsch. ne 2 4. C. coloratus Tamamsch. she 12 5. C. acanthocephalus C.A.M. of 13 6. C. hystrix C.A.M. se 14 7. C. poliochrus Trautv. a3 16 8. C. hajastanicus Tamamsch. ee 16 9. C. furiosus Tamamsch. Se 17 10. C. onopordioides Fisch. ex MB. a 18 11. C. atropatanicus Sosn. ex Grossh. a 20 Section 2. Carduastrum Tamamsch. 12. C. pseudocollinus (Schmalh.) Klok. ee 21 15700 13. C. transcaspicus Gand. be 21 14. C. hamulosus Ehrh. ee 22 1S. €. erispusilt ee 23 16. C. acanthoides L. pe 24 17. C. navaschinii Bordz. a, 25 18. C. uncinatus MB. ue 26 19. C. stenocephalus Tamamsch. oe D7. *Reproduced from the Russian original. Russian page numbers appear in the left-hand margin of the text—General Editor. VI XV1 15710 15720 15730 bay rat: Seowene) ae SS GO GS GG © Genus 1592. Cirsium Mill. emend. Scop. personata (L.) Jacq. glaucus Baumg. adpressus C.A.M. multijugus C. Koch | seminudus MB. . kerneri Simk. Section 3. Stenocephalus Rouy . pycnocephalus L. . beckerianus Tamamsch. . cinereus MB. . arabicus Jacq. . nikitinii Tamamsch. . nervosus C. Koch Genus 1590. Alfredia Cass. Section 1. Alfredia . cernua (L.) Cass. Section 2. Acantholepis Kar. and Kir. . acantholepis Kar. and Kir. . nivea Kar. and Kir. Genus 1591. Olgaea Ijin . eriocephala (Winkl.) Ijin . nivea (Winkl.) Ijin vvedenskyi Tin . longifolia (Winkl.) jin . lanipes (Winkl.) Ijin . nidulans (Rupr.) Ijin . Spinifera Tin . pectinata Ijin . baldshuanica (Winkl.) Tin Section 1. Caucasigena Charadze 1. C. aggregatum Ldb. 2. C. gagnidzei Charadze 40 41 42 44 45 46 47 47 48 49 50 50 69 71 3. C. longiflorum Charadze hs 71 4. C. czerkessicum Charadze 3: 72 5. C. imereticum Boiss. ans 74 6. C. kemulariae Charadze £. 75 7. C. chlorocomos Somm. and Lev. a 76 8. C. caucasicum (Adams) Petrak Qs 78 9. C. adjaricum Somm. and Lev. as 79 15740 10. C. erythrolepis C. Koch id 81 11. C. tricholoma Fisch. and Mey. c 82 12. C. osseticum (Adams) Petrak f 83 13. C. buschianum Charadze pf 84 14. C. sosnowskyi Charadze a 85 15. C. abkhasicum (Petrak) Grossh. MM 86 16. C. fominii Petrak se 87 17. C. caput-medusae Somm. and Lev. = 88 Section 2. Epitrachys DC. Subsection 1. Congesta Petrak 18. C. congestum Fisch. and Mey. ex DC. eft 90 19. C. sorocephalum Fisch. and Mey. ua 91 VII Subsection 2. Bracteosa Petrak 15750 20. C. bracteosum DC. i 93 21. C. strigosum (MB.) MB. Bee 94 22. C. rigidum DC. se 96 23. C. aduncum Fisch. and Mey. ex DC. ie 97 Subsection 3. Phyllocephala (Petrak) Charadze 24. C. bornmiilleri Sint. ex Bornm. ss 99 Subsection 4. Armata Charadze 25. C. lappaceum (MB.) MB. om 101 26. C. anatolicum (Petrak) Grossh. Be 102 27. C. megricum Charadze bas 104 28. C. schelkownikowii Petrak bet 105 29. C. macrobotrys (C. Koch) Boiss. ie 106 15760 30. C. cosmelii (Adams) Fisch. ex Hohen. ee 107 31. C. arachnoideum (MB.) MB. ae 110 32. C. euxinum Charadze on 110 33. C. laniflorum (MB.) MB. oe 111 Vill XVIil 15770. 15780. 15790. 47. 48. 49. 50. Sal. a2: Gs 5 CE ANS: BG, . balkharicum Charadze | tomentosum C.A.M. . daghestanicum Charadze . ketzkhovelii Charadze G, OVO) Ol OTe eere BESO OTOTS ie @y (1 sublaniflorum Sojak tauricum Sojak cephalotes Boiss. pugnax Somm. and Lev. tindaicum Charadze macrocephalum C.A.M. argillosum V. Petr. ex Charadze turkestanicum (Rgl.) Petrak pseudolappaceum Charadze Subsection 5. Eriophora Petrak polonicum (Petrak) Iljin Subsection 6. Ciliata Petrak . serrulatum (MB.) MB. . ukranicum Bess. . ciliatum (Murr.) Moench . Szowitsii (C. Koch) Boiss. Subsection 7. Lanceolata Petrak vulgare (Savi) Ten. Section 3. Lamyropsis Charadze sinuatum (Trautv.) Boiss. lamyroides Tamamsch. Section 4. Echenais (Cass.) Petrak . semenovii Rgl. and Schmalh. . Sairamense (Winkl.) O. and B. Fedtsch. . glabrifolium (Winkl.) O. and B. Fedtsch. . badakhschanicum Charadze . alberti Rgl. and Schmalh. . Sieversii (Fisch. and Mey.) Petrak . echinus (MB.) Hand.-Mazz. 112 114 114 116 117 118 119 120 121 122 123 124 125 127 128 129 130 132 135 Wes7/ 138 140 142 143 144 145 146 147 15800. 15810. 62. 63. 64. 65. 66. 67. 68. 69. 70. Pl: M2: 73: 74. 75. 76. ULE 78. 1D: 80. 81. 82. 83. 84. 85. 86. Section 5. Cirsium Subsection 1. Montana Charadze kusnetzowianum Somm. and Lev. sychnosanthum Petrak . oblongifolium C. Koch . waldsteinii Rouy hypoleucum DC. CHS) 1 @) Subsection 2. Sinocirsium Kitam. C. maackii Maxim. C. schantarense Trautv. and Mey. Subsection 3. Apalocentron (Cass.) Charadze C. oleraceum (L.) Scop. C. rivulare (Jacq.) All. C. erisithales (Jacq.) Scop. Subsection 4. Cirsium C. heterophyllum (L.) Hill C. helenioides (L.) Hill C. dealbatum MB. C. canum (L.) All. C. biebersteinii Charadze C. pannonicum (L. fil.) Link Subsection 5. Spanioptilon (Less.) Kitam. C. vlassovianum Fisch. C. coryletorum Kom. C. serratuloides (L.) Hill C. schischkinii Serg. C. komarovii Schischk. Subsection 6. Borealicola Kitam. C. kamtschaticum Ldb. ex DC. C. weyrichii Maxim. C. pectinellum A. Gray C. palustre (L.) Scop. X1X 150 152 153 153 154 157 158 161 162 163 165 166 168 169 170 171 173 174 176 LY 178 WS) 181 182 183 XX 15820. 15830. 15840. 107. Subsection 7. Orthocentrum (Cass.) Charadze . depilatum Boiss. and Bal. hygrophiloides Charadze hygrophilum Boiss. elbrusense Somm. and Lev. albowianum Somm. and Lev. apiculatum DC. elodes MB. alatum (S.G. Gmel.) Bobr. svaneticum Somm. and Lev. uliginosum (MB.) MB. subinerme Fisch. and Mey. rhabdotolepis Petrak GQ FOS OO OOOO S Subsection 8. Aciculata Charadze obvallatum (MB.) MB. . simplex C.A.M. . armenum DC. GB) QO) Subsection 9. Acaulia Petrak . C. esculentum (Sievers) C.A.M. . C. rhizocephalum C.A.M. . C. frickii Fisch. and Mey. . C. brevipapposum Tschern. . C. acaule (L.) Scop. Section 6. Pseudo-eriolepis (Nakai) Kitam. C. pendulum Fisch. Section 7. Cephalonoplos (Neck.) DC. . setosum (Willd.) MB. . incanum (S.G. Gmel.) Fisch. ex MB. . ochrolepideum Juz. . arvense (L.) Scop. Gy) GO) © Genus 1593. Notobasis Cass. . N. syriaca (L.) Cass. 185 186 187 188 189 190 191 192 193 194 195 197 198 199 200 201 203 204 204 205 208 210 211 212 213 216 je E Genus 1594. Picnomon Adans. P. acarna (L.) Cass. Genus 1595. Lamyra Cass. L. echinocephala (Willd.) Tamamsch. Genus 1596. Lamyropappus Knorr. and Tamamsch. iN ie — 15850. 1. hWN L. schakaptaricus (B. Fedtsch.) Knorr. and Tamamsch. Genus 1597. Ancathia DC. A. igniaria (Spreng.) DC. Genus 1598. Modestia Charadze and Tamamsch. . M. darwasica (Winkl.) Charadze and Tamamsch. ... . M. mira (Ijin) Charadze and Tamamsch. Genus 1599. Cynara L. . C. scolymus L. Genus 1600. Silybum Adans. S. marianum (L.) Gaertn. Genus 1601. Onopordum L. Section 1. Onopordum . O. acanthium L. . O. cinereum Grossh. . O. prjachinii Tamamsch. . O. tauricum Willd. Section 2. Erecta Rouy O. armenum Grossh. O. frickii Tamamsch. . O. leptolepis DC. O. candidum Nebelek . O. seravschanicum Tamamsch. O. heteracanthum C.A.M. XXi 218 220 221 222 224 225 226 227 231 232 233 233 234 236 237 238 238 239 XXil 15870. NO — = Subtribe 3. Centaureinae O. Hoffm. Genus 1602. Zoegea L. Section 1. Zoegea . baldshuanica Winkl. Section 2. Subulatae Czer. . Z. purpurea Fresen. Genus 1603. Russowia Winkl. . Sogdiana (Bge.) B. Fedtsch. Genus 1604. Crupina Cass. . crupinastrum (Moris) Vis. . vulgaris Cass. Genus 1605. Plagiobasis Schrenk . centauroides Schrenk Genus 1606. Tricholepis DC. . trichocephala Lincz. Genus 1607. Serratula L. Section 1. Serratula . inermis Gilib. Section 2. Mastrucium (Cass.) DC. . coronata L. Section 3. Piptochaete Boiss. . erucifolia (L.) Boiss. . dissecta Ldb. . angulata Kar. and Kir. Section 4. Klasea (Cass.) DC. . centauroides L. 248 249 252 253 254 256 258 265 268 270 271 212 273 XII 15880. 15890. 15900. a2. 33: AHRNRNHNNANNANNRANNANNANUNUNNY . komarovii Yjin . radiata (Waldst. and Kit.) MB. . biebersteiniana (Iljin ex Grossh.) Takht. gmelinii Tausch . tanaitica P. Smirn. . heterophylla (L.) Desf. cardunculus (Pall.) Schischk. marginata Tausch algida Ijin kirghisorum Ijin dshungarica Ijin transcaucasica (Bornm.) Sosn. ex Grossh. chartacea Winkl. procumbens Rgl. . suffruticosa Schrenk aphyllopoda (jin sogdiana Bge. alatavica C.A.M. . lancifolia Zak. . hastifolia Kult. and Korov. . coriacea Fisch. and Mey. quinquefolia MB. Section 5. Iljinia Boiss. lyratifolia Schrenk . tianschanica Sap. and Nik. . modesti Boriss. Section 6. Leuzeopsis Boriss. . serratuloides (Fisch. and Mey.) Takht. Section 7. Demetria Boriss. . caucasica Boiss. Genus 1608. Synurus Ijin deltoides (Ait.) Nakai Genus 1609. Syreitschikovia Pavl. tenuifolia (Bong.) Pavl. XXill 273 274 275 276 278 219 280 281 283 283 284 285 286 287 288 289 289 290 291 292 293 294 296 207 298 300 301 302 304 XXIV 2. S. spinulosa (Franch.) Pavl. a: S. Genus 1610. Schumeria jin Section Serratulopsis Ijin latifolia (Boiss.) Ijin 2. S. litwinowii (Ijin) Thin 15910. 5. XH 15920. Fe oe Se Seren ee AR aA aaa DR = ih: 2 RG: Rh. ellis Genus 1611. Rhaponticum Adans. Section 1. Fornicium (Cass.) Soskov carthamoides (Willd.) Ijin serratuloides (Georgi) Bobr. lyratum Winkl. ex Iljin Section 2. Eleutherochaetum Soskov integrifolium Winkl. Section 3. Rhaponticum . pulchrum Fisch. and Mey. . Satzyperovii Soskov . uniflorum (L.) DC. Section 4. Rhaponticellum Soskov . nitidum Fisch. . aulieatense (jin . karatavicum Rgl. and Schmalh. . namanganicum Ijin . nanum Lipsky Genus 1612. Amberboa (Pers.) Less. . moschata (L.) DC. . glauca (Willd.) Grossh. . sosnovskyi Ijin . nana (Boiss.) Ijin . turanica (jin . amberboi (L.) Tzvel. . bucharica Wyjin 305 307 308 3h S13 314 315 316 319 318 319 320 321 321 322 325 326 327 328 330 331 332 15930. 15940. 1. —" WN e Genus 1613. Grossheimia Sosn. and Takht. G. macrocephala (Muss.-Puschk.): Sosn. and Takht. . G. ossica (C. Koch) Sosn. and Takht. Genus 1614. Chartolepis Cass. . Ch. glastifolia (L.) Cass. . Ch. intermedia Boiss. . Ch. pterocaula (Trautv.) Czer. Ch. biebersteinii Jaub. and Spach Genus 1615. Callicephalus C.A.M. C. nitens (MB.) C.A.M. Genus 1616. Acroptilon Cass. . A. australe Ijin A. repens (L.) DC. Genus 1617. Oligochaeta C. Koch. . divaricata (Fisch. and Mey.) C. Koch. . tomentosa Czer. . minima (Boiss.) Briq. @) 1S) %) Genus 1618. Schischkinia Ijin . Sch. albispina (Bge.) Ijin Genus 1619. Aetheopappus Cass. . A. vwvedenskii (Sosn.) Sosn. . A. caucasicus Sosn. A. pulcherrimus (Willd.) Cass. Genus 1620. Cheirolepis Boiss. . Ch. persica Boiss. Genus 1621. Tomanthea DC. . T. spectabilis (Fisch. and Mey.) Takht. . T. aucheri DC. . T. phaeopappa (DC.) Takht. ex Czer. T. daralaghezica (Fom.) Takht. XXV 333 334 336 337 338 339 342 344 345 347 348 349 352 354 355 355 358 361 362 363 364 XIV XXV1 15950. 15960. We 2. i. Zi Subsection 1. Montanae (Hayek) Stef. and Georg. 14. 15: 16. La: 18. 19. Genus 1622. Stizolophus Cass. S. balsamita (Lam.) Cass. ex Takht. S. coronopifolius (Lam.) Cass. Genus 1623. Hyalea (DC.) Jaub. and Spach H. tadshicorum (Tzvel.) Sojak H. pulchella (Ldb.) C. Koch Genus 1624. Centaurea L. Subgenus 1. Centaurea . ruthenica Lam. . modesti Fed. . alaica Ijin gontscharovii Ijin androssovii Ijin hajastana Tzvel. kasakorum jin . kultiassovii Wjin . phyllopoda yin lasiopoda M. Pop. and Kult. . Razdorskyi Karjag. . talievii Kleop. . turkestanica Franch. SIGEOENES OS) SEO @) QEaS Subgenus 2. Cyanus (Juss.) Hayek Section 1. Protocyanus Dobrocz. C. maramarosiensis (Javorka) Czer. C. mollis Waldst. and Kit. C. nigrifimbria (C. Koch) Sosn. Subsection 2. Fischerianae Czer. C. fischeri Willd. C. willdenowii Czer. C. cheiranthifolia Willd. 366 366 368 369 380 381 382 382 382 383 383 384 385 385 386 386 387 392 393 394 396 398 399 XV 15970. 15980. 15990. 20. 2. 22. 23 24 2S. 26. 27, 28. 29. 30. 31. 32. 33. 34. 315), 36. Bi: 38. 39: 40. 41. 42. Subsection 3. Stricticaules Czer. C. ternopoliensis Dobrocz. C. stricta Waldst. and Kit. C. tanaitica Klok. C. czerkessica Dobrocz. and Kotov . C. angelescui (Prod.) Czer. Subsection 4. Variegatae Stef. and Georg. C. fuscimarginata (C. Koch) Juz. C. huetii Boiss. C. acmophylla Boiss. C. schelkovnikovii Sosn. C. woronowii Bornm. ex Sosn. Section 2. Cyanus Subsection 1. Depressae Stef. and Georg. C. depressa MB. Subsection 2. Cyaneae Stef. and Georg. C. cyanus L. Subgenus 3. Czerniakovskaya Czer. C. iljinii Czerniak. C. kopetdaghensis jin Subgenus 4. Psephellus (Cass.) Schmalh. Section 1. Hypoleucae (Sosn.) Sosn. C. albovii Sosn. C. abchasica (Alb.) Sosn. C. circassica (Alb.) Sosn. C. holophylla Socz. and Lipat. C. barbeyi (Alb.) Sosn. C. pauciloba Trautv. C. troitzkyi (Sosn.) Sosn. C. ciscaucasica Sosn. Section 2. Leucophyllae (Sosn.) Sosn. C. maleevii (Sosn.) Sosn. XXVIi 401 402 404 405 406 408 410 411 412 413 415 416 419 420 424 425 425 426 426 427 427 428 428 XXVIil 16000. 16010. 16020. QOS) OO Ge ao DONC STOr ee ere OV ee . leucophylla MB. . kolakovskyi Sosn. . declinata MB. maris-nigri Sosn. tuapsensis Sosn. . Salviifolia (Boiss.) Sosn. edmondi Sosn. hymenolepis Trautv. schistosa Sosn. Section 3. Psephellus . dealbata Willd. . exsurgens Sosn. . zuvandica (Sosn.) Sosn. taochia (Sosn.) Sosn. zangezuri (Sosn.) Sosn. sevanensis Sosn. karabaghensis (Sosn.) Sons. pambakensis (Sosn.) Sosn. somchetica (Sosn.) Sosn. colchica (Sosn.) Sosn. carthalinica (Sosn.) Sosn. meskhetica Sosn. Subgenus 5. Jacea (Juss.) Hayek Section 1. Jacea C. jacea L. mech é CV OlG) @ oO @ pannonica (Heuff.) Hayek substituta Czer. Section 2. Lepteranthus (Neck.) DC. Phrygia L. . carpatica (Porc.) Porc. . nigriceps Dobrocz. . abbreviata (C. Koch) Hand.-Mazz. . alutacea Dobrocz. . abnormis Czer. b ¢. oi Gin Oe aa OF pseudophrygia C.A.M. stenolepis Kern. salicifolia MB. hyrcanica Bornm. 429 429 430 431 431 432 432 433 433 434 435 435 435 436 436 437 437 438 438 439 439 444 446 448 449 451 452 453 454 455 456 458 458 460 XVII 16030. 16040. 77 84. 96 97 . C. trichocephala MB. Subgenus 6. Heterolophus (Cass.) Dobrocz. . carbonata Klok. . marschalliana Spreng. . sumensis Kalen. . sibirica L. . turgaica Klok. . sergii Klok. (©) Cre) rere Subgenus 7. Amblyopogon (DC.) Tzvel. C. ‘meyeriana Tzvel. Subgenus 8. Xanthopsis (DC.) Tzvel. . erivanensis (Lipsky) Bordz. . xanthocephaloides Tzvel. . xanthocephala (DC.) Sosn. SE O@R Subgenus 9. Sosnovskya (Takht.) Czer. . amblyolepis Ldb. . daghestanica (Lipsky) Czer. . ruprechtii (Boiss.) Czer. . arpensis (Czer.) Czer. Sr Bee) Subgenus 10. Odontolophus (Cass.) Hayek . C. trinervia Steph. . C. kobstanica Tzvel. . C. avarica Tzvel. Subgenus 11. Odontolophopsis Tzvel. . C. phaeopappoides Bordz. Subgenus 12. Pseudohyalea (Tzvel.) Tzvel. . C. leuzeoides (Jaub. and Spach) Walp. Subgenus 13. Hyalinella (Tzvel.) Tzvel. Section 1. Hyalinella . C. bella Trautv. XXiX 462 465 466 467 469 470 471 473 475 476 476 479 479 480 481 483 483 484 486 487 489 XVIII XXXK 16050. 16060. 16070. 98. 99. 100. 101. 102. 103. 104. 105. 106. 107. 108. 109. 110. WI 2? 113. 114. Wes 116. 117. 118. 119. 120. iP) 122. 123. C. nathadzeae Sosn. C. bagadensis Woron. C. simplicicaulis Boiss. and Huet C. dimitriewiae Sosn. C. adjarica Alb. Section 2. Albovia Tzvel. C. pecho Alb. Subgenus 14. Lopholoma (Cass.) Dobrocz. Section 1. Acrocentron (Cass.) DC. salonitana Vis. rubriflora (lar. reflexa Lam. sosnovskyi Grossh. Era a Section 2. Orientales (Hayek) Tzvel. C. orientalis L. C. kotschyana Heuff. Section 3. Lopholoma scabiosa L. integrifolia Tausch. ossethica Sosn. . apiculata Ldb. . adpressa Ldb. C. pseudoscabiosa Boiss. and Buhse C. glehnii Trautv. C. carduiformis DC. C. sintenisiana Gand. Wer eerie Section 4. Stereophyllae (Tzvel.) Tzvel. C. stereophylla Bess. Subgenus 15. Acrolophus (Cass.) Dobrocz. C. rhenana Boreau C. pseudomaculosa Dobrocz. C. kubanica Klok. C. micranthos Gmel. 490 490 491 491 492 493 498 499 499 500 501 502 503 504 506 507 507 508 509 509 510 511 517 518 518 519 XIX 16080. 16090. 16100. 124. 125: 126. Walle 128. 129. 130. Bile 132? 1337 134. 135. 136. lea 138. 139; 140. 141. 142. 143. 144, 145. 146. 147. 148. 149. 150. 151; 152; 38% 154. L535: 156. IS 158. 159. Subgenus 16. Phalolepis (Cass.) Dobrocz. (GE C. CQ’) © SON @HOFeNO ao OO) OOOO) Oia) Sie) @ Gee CVO eres majorovii Dumb. sophiae Klok. arenaria MB. odessana Prod. . borysthenica Gruner | savranica Klok. . besseriana DC. | lavrenkoana Klok. . steveniana Klok. . koktebelica Klok. vicina Lipsky ovina Pall. gulissashvilii Dumb. diffusa Lam. aemulans Klok. caspia Grossh. . squarrosa Willd. . albida C. Koch . aggregata Fisch. and Mey. Section 1. Pseudophalolepis Klok. pseudoleucolepis Kleop. proto-margaritacea Klok. margaritacea Ten. margarita-alba Klok. konkae Klok. appendicata Klok. proto-gerberi Klok. doneizica Klok. pineticola jin breviceps Ijin paczoskii Kotov ex Klok. gerberi Stev. dubjanskii Ijin Section 2. Phalolepis . semijusta Juz. . vankovii Klok. . sterilis Stev. . sarandinakiae Illar. XXX1 520 521 522 523 Bye) 524 526 526 527 528 530 530 531 532 533 534 535) 536 337 542 543 543 544 545 545 546 547 548 548 549 550 550 551 552 53 555 XXXli 16110. 16120. 160. 161. 162. 163. 164. 165. 166. 167. 168. 169. 170. Wk 172. We: 174. LS 176. Laer: 178. C. novorossica Klok. C. georgica Klok. C. latiloba Klok. C. transcaucasica Sosn. ex Grossh. C. alexandri Bordz. Subgenus 17. Microlophus (Cass.) Hayek Section 1. Microlophus C. behen L. Section 2. Polypodiifoliae Tzvel. C. szovitsiana Boiss. C. polypodiifolia Boiss. Subgenus 18. Seridia (Juss.) Czer. C. stevenii MB. Subgenus 19. Rhizocalathium (Tzvel.) Tzvel. C. rhizantha C.A.M. C. grossheimii Sosn. C. rhizanthoides Tzvel. C. armena Boiss. C. oltensis Sosn. Subgenus 20. Solstitiaria (Hill) Dobrocz. C. solstitialis L. C. adamii Willd. Subgenus 21. Calcitrapa (Adans.) Hayek C. iberica Trev. C. calcitrapa L. Subgenus 22. Tetramorphaea (DC.) Czer. C. belangeriana (DC.) Stapf Genus 1625. Phalacrachena Ijin . Ph. inuloides (Fisch. ex Schmalh.) Ijin . Ph. calva (Ldb.) Tin 556 557 557 558 559 561 562 563 564 567 568 568 569 570 Sai 573 574 575 578 580 581 16130. 16140. Genus 1626. Carthamus L. Section 1. Kentrophyllum (Neck.) Fiori and Paol. 1. C. lanatus L. 2. C. glaucus MB. Section 2. Carthamus 3. C. oxyacanthus MB. 4. C. gypsicola Ijin 5. C. tinctoricus L. Genus 1627. Cnicus L. 1. C. benedictus L. Tribe 13. Mutisieae Cass. Subtribe 1. Mutisinae O. Hoffm. Genus 1628. Gerbera Cass. 1. G. jamesonii Bolus Genus 1629. Leibnitzia Cass. 1. L. anandria (L.) Turcz. . L. knorringiana (B. Fedtsch.) Pobed. i) Genus 1630. Uechtritzia Freyn 1. U. armena Freyn 2. U. kokanica (Rgl. and Schmalh.) Pobed. XXXIl1 583 584 584 585 586 587 590 593 594 596 O97 # oat (Peck kp St Hie: ails tei) Dela 4 re ia | #. +N JB oS KEY TO GENERA OF CARDUINAE Stamen filaments hairy or densely barbate..................:eseseseeseeeeseeeees 2. Stamen filaments glabrous, rarely glandular or slightly tuberculate, uit meyer Wainy’ tise vis ris. syed mee metic ches one ke seuel meee ull a ut 8. Bl AMtS OUSPUIY, 86 lh cs Xaa nla cai cas panna astra ANE De oasis 3. MANNS SPOMIVY cB aces aan Oenaes pected dt aautaos nn ERA WA NO A 8 4. Receptacle flat, alveolate, with pits surrounded by small scales; achene lacking corona, with obtuse edges; plants acaulescent or with short (up to 30 cm high), usually simple stem (Caucasus). paler AMIR fo cccesasieisntcasecetdadenas 1585. Jurinella Jaub. and Spach Receptacle almost cylindrical, convex, covered with long scales; achene with small apical corona; stems much taller (50-100 cm high), strongly branched from base, numerous (Central Kopetdae):a .elictulas.sxmneloavscaae welauve ators abe 1588. Nikitinia Ijin Pappus hhairs/scabrotisi): srs. beneenes ional. pieeeet ere ee... 5. Pappus hairs plumose isis... nieces... cere, «luni, 8 EON, cose 6. Stamen filaments free above, fused below because of dense short, mucilaginous hairs covering them; pappus surrounding corona of very short, thin, smooth hairs along upper edge of ringlet joining base of hairs; leaves with large white spots.......1600. Silybum Adans. Stamen filaments free throughout their length; pappus lacking co- rona of short hairs; leaves lacking spots............. 1589. Carduus L. Achenes with thick pericarp, becoming hard with maturity, woody, ovate-hemispherical or somewhat pyriform........ 1595. Lamyra Cass. Achenes with less thick pericarp, of different shape..................... Tf! ~ Receptacle covered with scales; capitula homogamous, with bi- Sexudlifertile floretsiit swe. se bemted.. meneeints nieve Re. .o Me doo osetye' Behram! NO Pichi oniinninn 1592. Cirsium (Mill.) (cf. also couplet 26). Receptacle covered with narrow linear scales; ray florets externally similar to bisexual fertile disk florets, but with undeveloped ovary and empty (lacking pollen) anthers, as a result capitula considered hetero gamolsrs:. 22us...emcrereen cdl seucweb., se 1593. Notobasis Cass. Receptacle deeply pitted; pits with membranous, unevenly incised- toothed margin; capitula large (4—7 cm in dia, smaller only in O. frickii Tamamsch.); plants highly spiny.......... 1601. Onopordum L. Receptacle covered with bristles or scales, very rarely (in some species of Saussurea DC.) glabrous. .........:.c.c.ccccscccesscesssesenseeseceees 9: Pappus bristles scabrous or serrulate, rarely achenes lacking pappus (im Some:species Of Cousinia Cassie sc iswsssrcseccuscedoesbesnoendsadss 10. 14. lie 18. All or only inner pappus bristles plumose...................::seseeeeee 18. Anthers lacking basal appendages; achene (mature) usually one in each capitulum (Nuratau Mountains in Soviet Central V2.1 (21) | ee en emcee eet Coa Sone 2h 1577. Anura (Juz.) Tschern. Anthers with basal appendages; achenes numerous in each capitu- DOWN yo cus canis nccexesvcsh ac endeeeesee Savevaees eeepc eateencee nes eaten cols Beaee ee easeaeaee 11 Pappus bristles connate at base forming ring, readily falling off tO BETS 0. 20 N.S EE soa 12 Pappus bristles not connate at base and not forming ring, falling off SUNG seas weveasocdsrnseraeremneusdaorndsiessonore eee Oa ee me sbeces 15 LeAVes NOt SPI ves csasseaevecoseocaveresst ees sesed TIA AER AO tae de 13: Leaves and involucral bracts Spimy...............sssesseesesseeeeeeeseeeeeeeeesees 14. Receptacle covered with few scales; achenes oblong-obconical, with inconspicuous ribs, lacking apical corona (Central Kopetdag ii} inte SBI d. sectencasssnsnrarasseroandees 1586. Perplexia Iljin Receptacle usually densely covered with scales; achene terete, 4- angled, ribbed, less often somewhat compressed, nearly oblong, with apical corona............ 1584. Jurinea Cass. (cf. also couplet 21). Involucral bracts with scaly appendages; corolla tube always shorter than limb; basal appendages short (1.0—2.5, less often to 4 mm long), broad; inner pappus bristles slightly flattened at fp ln..19), QRARARR IIE. RRA. ROE, SA, BE 1590. Alfredia Cass. Involucral bracts lacking appendages; corolla tube longer than limb, less often as long; basal appendages long and narrow; inner pappus bristles not flattened at tip but much longer and closely toothedced. Mawes adh Menno. 2a) Mae 1591. Olgaea Iljin Pappus bristles slightly unequal in length but usually of same thickness. Ae ORG, BAO, TERT, SN Se, A 16 Pappus bristles distinctly unequal in length.....0...0.. eee eee 17; Involucral bracts coriaceous, spinescent, usually not uncinate; receptacular bristles longer than achene or almost equal, sometimes 1/2-2/3 as long; pappus bristles readily falling off................... SRR RPG. AOD LUIS DURA REE A eccrine 1578. Cousinia Cass. Involucral bracts scaly, with very small apical spine; receptacular bristles 1/5—1/4 as long as achene; pappus bristles persistent, fall- ine Off later LNB. 8.28. ASS LOG Be 1579. Lipskyella Juz. Involucral bracts strongly spinescent, not uncinate; receptacular scales serrulate-scabrous; plants perennial; leaves spiny (Tien Shian) 2 tate, AAAS, AR a 1580. Hypacanthium Juz. Involucral bracts with thin uncinate spiny; receptacular bristles sca- brous; plants biennial; leaves not spiny................. 1576. Arctium L. Receptacle covered with scales, occasionally with papillae (recep- tacle glabrous in some species of Saussured DC.).........::esccecceseeeeee 19. 20. 20 22. 23: Die Receptacle covered: withibristlesen 2 AR Ree 22} Achenes pubescent with scattered long hairs, recurved below, squar- rose above; bristles of inner row of pappus connate at base and fused to tube at different heights, tube appressed-hairy outside (3.0- 5-5 mm Jong) s2teretiveen ene e eed & 1583. Polytaxis Bge. Achenes glabrous; pappus bristles not fused to tube... 20. Pappus many-rowed; its bristles unequal, connate at base in ring or firmly fused with achene and then ring absent............. cesses 21. Pappus one- or two-rowed; bristles of outer row of pappus short- ened, toothed or finely plumose, readily falling off singly, those of inner row equal, always long-plumose, connate at base in ring, PEISISIEHE A. AISA. WAR ea 1582. Saussurea DC. Anthers covered with long, thin, sinuate, fragile, readily deciduous Pappus bristles not connate at base in ring, readily falling off sin- gly, short-plumose. Alpine and subalpine plants with twice pin- nately cut leaves (Tien Shan)............... 1581. Schmalhausenia Winkl. Pappus bristles connate at base in ring, falling off together.......... 23: Capitula surrounded by apical leaves; outer involucral bracts with plumose and spiny appendages; annual or biennial...................... POERSTE NIN BE ES IEEE AE CE NE, 1594. Picnomon Adans. Capitula not surrounded by apical leaves; spines of involucral bracts not pluniose; perennial ee. yGeNt. ROO AGRA OA RARL Be 24. Receptacle fleshy; leaves very large, broad, twice pinnately cut, clustered in rosette. Cultivated plants................... 1599. Cynara L. Receptacle not fleshy; leaves almost entire, undivided or pinnately lobed ih G20 AOL, ARGS, UB DA Te es 25 Achenes slightly compressed from both sides, not ribbed............ 26. Achenes not compressed, ribbed-angular.............eeeeeeeseeeeneeeees 2 Achenes obliquely truncated at apex; pappus many-rowed....... FEE Dis GON iN RIN, 1592. Cirsium Mill. (cf. also couplet 7). Achenes apically flat, with saucer-shaped depression; pappus two- rowed, of bright golden bristles, outer ones at first somewhat thick- ened brushlike at top, then acuminate (Soviet Central Asia)...... MORALE, STIOL RRM, 1596. Lamyropappus Knorr. and Tamamsch. Stem winged; leaves oblong-elliptical or lanceolate; achenes white, pale yellow or brownish; pappus many-rowed, of short-plumose bristles, outer pappus shorter (Pamiro-Alai Region).............. fewest, Ae ES ae 1598. Modestia Charadze and Tamamsch. Stem not winged; leaves linear or linear-lanceolate; achenes dark brown or almost black; pappus more or less two-rowed, of long- plumose, almost equal bristles................ccccceeeee 1597. Ancathia DC. GENUS 1589. Carduus L. emend. Gaertn.'? L. Sp. pl. (1753) 820 p. p.; Gaertn. De. fruct. II (1791) 378. Capitula homogamous, homochromous, with few (10-12) or nu- merous (100) bisexual flowers; usually all fertile. Involucre ovate, cylindrical, more or less oblong, campanulate, obconical, globose or flat-globose; involucral bracts many-rowed (8—10 rows), imbricate, ap- pressed or divergent, their tips projecting in same or different direc- tions, appressed or recurved at apex, sometimes geniculate, constricted in middle; outer bracts ovate, deltoid-ovate, lanceolate or linear with short or long cusp; middle longer, sometimes abruptly narrowed up- ward from much wider base; innermost bracts usually longest, some- times colored in upper half, membranous, acute or obtuse, usually not spiny, fimbriate-toothed or irregularly incised on upper margin; all bracts more or less flat or convex outside, with prominent or incon- spicuous midrib, glabrous or variously pubescent with unicellular hairs usually on margin or with vesicular greenish or yellowish pseudo- glandular hairs (usually in annual bristle-thistles having a close resemblance with C. pycnocephalus), or with clavate hairs in lower part on outside or margin, or with long multicellular arachnoid hairs. Receptacle flat or slightly convex, densely covered with long bristles. Corolla tubular-cyathiform, often irregularly incised; corolla tube equal to, shorter or longer than limb including throat, usually straight or geniculate in ray florets such that ray florets form a circle from sur- face, wider than receptacle and involucre; limb lobes narrowly lan- ceolate, bent hood-like at tip. Stamen filaments pilose in middle, with long and curly hairs; anthers with slightly incised basal appendages. Style branches short, usually approximate, spirally recurved in some species at end of flowering, with ring of short erect hairs at base; style with thin articulation at base where it breaks. Achenes 3-6 mm long, oblong, ovate, obcuneate or more or less terete, slightly compressed laterally, weakly truncate at base, not ribbed, smooth, with oily lustre, olive, grayish, bright- or dark brown, with or without very thin longi- tudinal striations and with darker undulating transverse striations; after falling of purple, orange, pink, bluish-pink, rarely white or yellowish corolla and pappus there remains in upper portion of achene a nectary- 'Treatment by S.G. Tamamschian. From the ancient Latin name used for spiny plants. 5 like tubercle of uncertain morphology, which is variously called in the literature (umbonus, mamelon, regma, epiregma, elaiosome, nectary); lower portion of hilum scarcely visible, almost straight or somewhat truncate, narrow; pappus of many-rowed bristles often unequal in length and thickness, connate at base in ring, entirely falling off, equal to, longer or considerably shorter than inner involucral bracts or corolla; each bristle consists of relatively thick central axis in contrast with bristles of Cirsium formed by short and broad cells, usually tapering upward, along sides covered with short trichomes not longer than the diameter of axial part of pappus bristles. Annual, biennial or, less often, perennial, usually spiny herbs with alternate, usually large, un- divided, pinnately lobate, pinnately parted or pinnately cut leaves, spiny on margin and at tip. Type of genus: C. nutans L. Note. The genus Carduus [plumeless or bristle-thistles] includes about 120 species, of which more than 30 are found in our country. Despite numerous reviews of the genus for different parts of the world (it is found mainly in Europe, Northern Africa, and West Asia, and is introduced in America and Australia), there is no monograph of the genus. The systematics of Carduus poses tremendous difficulties be- cause of the confusion of polymorphic species aggregates with very many hybrid forms. Many hybrids have been described as new species in Western Europe. Economic Importance. Almost all bristle-thistles are good honey- producing plants. Some species are highly ornamental, but they are weeds nonetheless. Oil is contained in their cotyledons. 1. Involucre globose, oblong or hemispherical or broadly ovoid, bien- nial or perennial, quite large (approximately 40 to 100 cm high) PMRRRES ERE UE Hi tat A ays eh keh NRO OS OIA lo oF + Involucre oblong, cylindrical, narrowly campanulate or truncate- OBEONICA LEGA Sian I Le can ean, se OUR | ee, 26. 2. Capitula nodding, involucral bracts somewhat broad, herbaceous, slightly constricted and geniculate near middle, often recurved in TEpPPe Tea het aU OD RE, PTE RR. ON a + Capitula on straight peduncles or very slightly nodding at end of peduncles just below capitulum; involucral bracts never genicu- Fetter TEETER, NU) IOs» idee Cel es ANNE AN ER Soy 6. 3. Plants 50-100 cm high; stems usually many, branched; capitula large, globose, (3) 4—6 (7) cm in dia; leaves glabrous or pubescent pencath along Vems file tik ee) alee 1. C. nutans L. + Plants to 40-50 cm high; stem usually solitary, less often 2-3, simple; capitula oblong-globose..............:cccsssssessessssssseesssesecseessseees 4. 11. Plants hardy, with highly spiny-winged stem and spiny leaves; ca- pitula in racemes at stem apex; corolla usually white.................00. na sap» Gather hala septa at Sea anh a eb dake 2. C. songoricus Tamamsch. Weaker and less spiny plants with peduncles not winged; capitula usually ico litaiyy is ae trse sh eaceton haat epsitiye. bd depp and te Peas a) Involucral bracts 1.5—-2.0 mm wide, green, all uniformly colored; slender, few-leaved plants................. 3. C. schischkinii Tamamsch. Involucral bracts 2-3 mm wide; outermost bracts yellowish-green at base, greenish-pink in middle and more or less violet at tip; capitula oblong-globose, large, 2.5—-3.5 cm long; corolla pink or Straw colores: cisteatee otebeses go tun tact uence 4. C. coloratus Tamamsch. Involucral bracts herbaceous, but narrow, stiff, sometimes woody, projecting upwards, slightly or sharply recurved, with long cusps or distinctly, orandistinethy uncinate!srced. bots ted eee a Te Involucral bracts not woody, usually with short spines or cusps, imbricate or indistinctly imbricate below, more or less recurved in upper half, widely divergent or uncinate..............cccceeesseeseeeeseeseeee 16. Involucral bracts tapering from more or less broad base to more or less long, cuneate cusp, erect or divergent, recurved or in various directions, sometimes almost horizontal or curved saber-like.......... 8. Involucral bracts narrow, grooved inside, distinctly (and then ca- pitula globose) or indistinctly (and then capitula more oblong) UNnCINAteit 6). nrceewiehentes h.. Be. eet. ape te hee ae eg 14. Plants glabrous or slightly pubescent, light-or yellowish-green or moreson less, bluish-darks ora yi chisserielnvsnscctepeeeestt. verte. heaven, Sees 11. Plants more densely pubescent, with grayish-arachnoid-hairy pu- bescence, white-tomentose or dark QreeM............ eee eseeeeeeseeeneeeeeees 9. Whole plant white-tomentose; involucral bracts narrow, curved saber-like, in upper half narrowed to sharp, slightly curved spine, below tightly appressed................esseesseseeeeeeeeees 6. C. hystrix C.A.M. Plants dark green with fine, more or less arachnoid-hairy pubes- GON GErt5, oer)... seule ace roa... caren ah eas ole creel de at 10 Capitula ovate-globose, to 2—3 cm in dia; involucre subglabrous or slightly pubescent; involucral bracts subulately spinescent from narrowly lanceolate base colored.................. 7. C. poliochrus Trautv. Capitula globose, 34 cm in dia, almost sessile; involucral bracts narrowed from ovate base, to long straight, more or less distinctly triangular spinesis.c.. cases ceed et 5. C. acanthocephalus C.A.M. Plant dark bluish-green; stem tall (70-80 cm), divaricately branched; capitula solitary, on slightly nodding peduncles; corolla dark purple.. met BN eal aa a eee ek a ee 8. C. hajastanicus Tamamsch. Plant bright green, glabrous or slightly pubescent; stem generally solitary, unbranched; capitula solitary or aggregated on short erect 12. 13. 14. iS: 16. 1h peduncles; corolla light-pink or slightly purple............0........0 12. Capitula oblong or broadly obconical; involucral bracts with short spines, smaller and more or less herbaceou.................:ccsccecceee 13. Capitula globose, involucral bracts, stiff, abruptly narrowed from more or less broad base to subulate tip, terminating in almost horizontal stiff spine; bracts variously directed... .eeeeeeeees Deep cant alan alive dene. ww cveroann cxnsibeth ssunnst es seh 9. C. furiosus Tamamsch. Involucral bracts almost equal in length; outermost usually shorter than middle and inner ones, all acuminate-spinescent from broad base to straight, small, erect spine, with prominently raised midrib, more or less distinctly triangular.....................000 sa Suemparenendlivven, sobeaatenls, cca uuarhye, tag 10. C. onopordioides Fisch. ex MB. Involucral bracts distinctly unequal in length, outer 1/2 as long as erect inner ones, recurved, 1/2 as broad as in preceding species, lacking distinct midrib, more flat and less stiff.........000000.000... ieOsted wee kinetic, wa orsnlee Jaa 11. C. atropatanicus Sosn. ex Grossh. Capitula oblong, medium-sized, 1.5—2.0 cm in dia; 2.5—3.0 cm long; involucral bracts indistinctly uncinate, more or less distinctly imbri- cate; belowewtslemaniuedt 12. C. pseudocollinus (Schmalh.) Klok. Capitula flat-globose, large, 3.5 cm in dia and long, sometimes Plant large, often to 1 m high; peduncles 18-20 cm long, usually not winged or narrowly and discontinuously winged only above, with small leaves; capitula large (3.5) 4.04.5 cm in dia; involucral bracts very distinctly and uniformly uncinate; corolla tube includ- ing throat 16-18 mm long, limb 8-9 mm long....................... eR anh eeanth. sepencied: cline: beetle ee seen aah dl 13. C. transcaspicus Gand. Plant shorter, 50-70 cm high; peduncles shorter, almost always winged; capitula globose or slightly oblong; involucral bracts not so distinctly and uniformly uncinate; corolla tube including throat 16-18 mm long, limb 9-10 mm long......... 14. C. hamulosus Ehrh. Involucral bracts appressed, distinctly imbricate, outer short, ovate, abruptly narrowed to short spinule........... 22. C. adpressus C.A.M. Involucral bracts not appressed, indistinctly imbricate, recurved to vanyingwdesreesyinaipper halfiene) .2eetdemel Av rn... 17. Involucral bracts more pliant, generally grayish-pubescent, sharply uncinate-recurved; capitula oblong or depressed-globose............ 18. Involucral bracts more stiff, squarrose, recurved in upper half, but not uncinate, usually glabrous or slightly pubescent either on mid- Fb OG GN: MAH MM, seeks eal meetsincryen Gti). eecl ale Maerua 19. Capitula large, 3.5—4.0 (5.0) cm in dia, broader than long, de- pressed-globose; involucre often compressed in middle, grayish- Fe LPLCILIUONEO ace. tres: epoca tesa os. eee eee eee 18. C. uncinatus MB. Ble 22) 23: 24. 25: Capitula smaller, oblong, 2.5—3.0 cm long, 1.5 cm in dia; involucre oblong, not compressed in middle; involucral bracts almost glabrous, slightly arachnoid pubescent only in upper part........... BEL ers, Rar ee ere a UE 19. C. stenocephalus Tamamsch. Peduncles short, usually winged right up to capitula or not winged only below capitula, but covered with spinules..............0...0.2 20. Peduncles long, wingless, lacking spines, but with white or grayish PUBESCENE EES AER NCO TP RAN Se AG 23: Corolla white; peduncles sometimes not winged just below capitu- WAS EEE LAE. PA A 17. C. navaschinii Bordz. Corolla light or dark purple; peduncles spiny-winged or simply spiny, rarely not winged, almost always densely pubescent........... Pale Leaves pliant, long-petiolate, ciliate-spiny on margin, grayish be- neath from dense pubescence, dark green above, sometimes slightly pubescent, large (to 20 cm long); middle cauline leaves elliptical, slightly incised or entire, sometimes almost orbicular; upper leaves always entire, lanceolate..................0 20. C. personata (L.) Jacq. Leaves stiffish, often more or less coriaceous, smaller, if large then short-petiolate, lanceolate, broadly lanceolate or oblong, but dis- tinctly lobed, with lobes of different SizeS................::ccessseceeseeeeees 22 Plants dark green; capitula much smaller, in clusters of 3—4(5) at tip of peduncles, occasionally fewer; peduncles sometimes not winged just below capitulum; involucral bracts less stiff, upper part recurved at obtuse angle, linear; lower leaves sometimes slightly lobed Mlargen eee ee ee Aa ee 15. C. cripus L. Plants bluish-gray; capitula large, usually solitary, on spiny- winged peduncles; involucral bracts stiffer, with longer upper half obliquely divergent and recurved at wide angle, with shorter, yellowish Spine ee A RO ae 16. C. acanthoides L. Plants dull green with bluish tinge; capitula somewhat drooping at MOWER ES 2. E.R 21. C. glaucus Baumg. Plants green or grayish- -green; capitula on erect, very long pe- dunclesmee i Sahn RR aL 24. Involucral bracts and pappus bristles considerably shorter than corolla; involucral bracts distinctly imbricate in lower part........ 25! Involucral bracts and pappus more than 1/2 as long as corolla; outer and middle bracts linear, almost equal, recurved; stem often glabrous, 0.8 cm in dia, ribbed-sulcate.......... eee esse eseeeeneeeeeees A SR A Oe ee A OE Bre 23. C. multijugus C. Koch. Peduncles equal to or more than half the length of entire plant, irregularly floccose-white tomentose; leaves lanceolate, narrow, 2— 3 cm wide, thick, or broader, to 5 cm, but then thinner, slightly Sinuatestoothed Rei. UA 8 24. C. seminudus MB. 9 + Peduncles not so long, pinkish white-tomentose; leaves oblong, thick, sometimes even more or less coriaceous, pinnately parted or very symmetrically and deeply pinnately lobed........... 25. C. kerneri Simk. 26. Upper part and triangular spine of involucral bracts in mature and young capitula recurved. Corolla 16.0-16.5 mm long............ NRE os tele Manat Gres tecr ec teeeeene cae tee actanes tee 26. C. pycnocephalus L. + Involucral bracts appressed, with erect, not recurved, spine or with slightly recurved spine only in outermost bracts...............:000 Die 27. Outer involucral bracts lanceolate, ovate, or ovate-lanceolate, to- gether with middle bracts in upper half, with dark purple spot, more or less thickened and hard, abruptly ending in small spine....... bea eh RE, A RR 27. C. beckerianus Tamamsch. + Outer and middle involucral bracts at upper end not thickened, lacking dark spot, imperceptible narrowing to a spine................ 28. 28. Involucral bracts strongly pubescent, not lustrous after flowering, besides pseudo-glandular hairs, densely covered with multi-cellu- lar, often entangled hairs; plants grayish- or white-pubescent.......... 29: + Involucral bracts subglabrous or weakly pubescent; capitula often becoming somewhat lustrous green toward end of flowering; bracts narrowly scarious along margin, yellowish only at base from pseudo- lang lary havi Se. Se eee eee 30. 29. Plant entirely light gray; capitula medium-sized, on unwinged con- spicuous peduncles, usually 1-2; achenes 3-4 mm long, if up to 5 mm, then peduncles also longer and capitula larger and soli- tei SR, TR OLE De. eae ree 28. C. cinereus MB. + Plant weakly pubescent or white-pubescent; capitula sessile, in clus- ters of 3-4, axillary and terminal on usually winged and spiny- winged peduncles; usually one middle capitulum much larger and more densely set, rest readily falling off already in predehiscent Condition: 224N9..conle wel ake, Bere fee Ee, 29. C. arabicus Jacq. 30. Plants 5—18 cm high, with 1—2 small, almost sessile capitula 10-11 mm long; capitula in leaf axils and at end of peduncles; leaves Gera VASIa ee Me Ma, LON Pinkie, Saree 30. C. nikitinii Tamamsch. + Plants 30-40 cm high, with capitula 15-18 mm long at stem tips in clusters of 3—6 cm long, thick (1.5—2.0 mm in dia) peduncles.......... Ne Ree A a aie heals 31. C. nervosus C. Koch. Section 1. Carduus.—Sect. Carduus subsect. Pachycephali auct. non-null. p. p.—Sect. Platycephali Rouy, Fl. Fr. 1X (1905) 75 p. p.— Capitula large, 3-6(7) cm in dia, globose, flat-globose, broadly obo- vate or somewhat oblong; involucral bracts with long spines. Biennials or perennials. Type of section: type of genus. 10 Series 1. Nutantes Tamamsch.—Capitula nodding; involu- cral bracts broad, herbaceous, slightly constricted in middle and geniculate. 1. C. nutans L. Sp. pl. (1753) 821; DC. Prodr. VI, 621; Ldb. FI. Ross. II, 718; Boiss. Fl. or. III, 515; Hegi, Ill. Fl. VI, 2, 848; Grossh. Fl. Kavk. IV, 174; Shishk. in Kryl. Fl. Zap. Sib., XI, 2878; Klok. in Vizn. Rosl. URSR, 566; Tamamsch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XV, 392.—C. nutans B. thoermeri Schmalh. F1. II (1897) 97.—C. armenus Boiss. FI. or III (1875) 516, pro syn.—C. thoermeri Weinm. in Bull. Soc. Nat. Mosc. X (1837) 69; Iljin in Izv. Bot. Sada Akad. Nauk SSSR, XXX, 3-4, 349; Grossh. Opred. Rast. Kavk. 479; Klok. loc. cit.; Kapeller in Fl. Gruzii, VIII, 465.—C. thoermeri ssp. sibiricus Ijin, loc. cit.; Schischk. in Kryl. Fl. Zap. Sib. XI (1949) 2878. Biennial. Plant 50-100 cm high. Stem usually branched, less often simple, cylindrical, weakly sulcate, arachnoid-hairy or subglabrous, winged, wings sinuate-toothed, on margin covered with long spines. Lower leaves short-petiolate; middle and upper leaves decurrent, 15— 40 cm long, 5—10 cm wide, uniformly bright green on both sides, with prominent midrib, glabrous or pubescent beneath along midrib, some- times lateral veins with long, multi-cellular crisped hairs; sometimes entire surface somewhat arachnoid-hairy-pubescent, pinnate and lo- bate, with 2-5 ovate, shallow lobes, sharply spinose on margin. Ca- pitula nodding, 4—7 cm in dia, sometimes with as many as 100 florets, solitary on white-tomentose wingless peduncles. Involucre globose; involucral bracts unequal, outer and middle geniculate and somewhat constricted usually below middle, sublanceolate above, broadly del- toid or ovate, flat or somewhat grooved inside, divergent or recurved, sometimes recurved at an acute angle, less often erect, terminating into sharp spine. Corolla purple or pink, to 15 mm long; limb including throat 5 mm long, quite irregularly incised. Achenes 4—5 mm long, light gray or light olive, with fine longitudinal striations and transverse fine zigzags; pappus of white scabrous hairs, considerably shorter than corolla. Flowering June to September. Borders, roadsides, weedy places, glades, pastures—European Part: Almost all regions except Arctic; Western Siberia: Upper Tobol, Ob Region, Irtysh, Altai (in the last two places rarely); Caucasus: All regions. Soviet Central Asia: Aralo-Caspian Region, Lake Balkhash Region, Kyzyl-Kum (rarely). General distribution: Almost whole of Western Europe, Armenia and Kurdistan, Iran, North Africa. Described from Western Europe. Type in London. 11 Economic Importance. It is considered a good honey-producing plant; it yields abundant nectar, and the honey obtained is light-col- ored, very aromatic, and with a pleasant taste. The seeds contain up to 20-30% rich oil in their cotyledons. It can be recommended as an ornamental plant. 2. C. songoricus Tamamsch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XV (1953) 391.—C. nutans var. songoricus Winkl. ex O. and B. Fedtsch. Perech. Rast. Turk. IV (1911) 277, nomen.—Ic.: Tamamsh. op. cit. Fig. 8 on p. 387. Perennial. Stem thick, sulcate, straight, simple, winged, densely leafy. Basal leaves on short, spiny winged petioles, densely pubescent, particularly on midrib with long crisped hairs, oblong or oblong-obo- vate, pinnately lobed, lobes 5S—7 toothed, acute, spiny; middle and upper leaves much narrower and longer, decurrent, pinnately lobed, lobes 3—5 toothed with prominent veins, extended into stiff sharp spine, on both sides, particularly on midrib, with very sparse, long hairs. Capitula at stem tips in clusters of 3—5, on short, drooping, white- tomentose peduncles, winged almost to capitulum, globose, to 4-6 cm in dia. Involucral bracts geniculate, ovate in lower half, lanceolate in upper half, slightly recurved, ventrally grooved, dorsally with promi- nent midrib, terminating in acute spine, with 2—4 thin, distinct, lateral veins. Corolla mostly white, sometimes pink; limb irregularly incised; pappus bristles whitish, slightly scabrous, more than a half as long as corolla. Achenes 6 mm long. Flowering July to August. Stony steppes, dry mountain slopes.—Soviet Central Asia: Dzhungaria-Tarbagatai, Pamiro-Alai Region. General distribution: Dzhungaria-Kashgaria. Described from Dzhungarian Alatau. Type in Leningrad. 3. C. schischkinii Tamamsch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk. SSSR, XV (1953) 389.—C. onopordioides auct. non Fisch. ex MB.: O. and B. Fedtsch. Perech. Rast. Turk. IV (1911) 277 and auct. al. fl. turkest.—Ic.: Tamamsh. op. cit. Fig. d, e on p. 387. Annual or biennial. Plant to 50-60 cm high. Stem simple, mostly single, slightly angular, sulcate, slightly contorted, narrowly winged, wings on margin finely ciliate-spiny. Leaves decurrent, of average size, 10-12 cm long, 2.0—2.5 cm wide, oblong-ovate, slightly lyrate or sinu- ate; lobes 3—S-toothed with thin spines; lamina bright green on both sides, glabrous above, slightly hairy beneath at base on midrib with long wavy hairs; peduncles long, not winged, white-tomentose, with solitary capitula. Capitula almost cylindrical; florets almost twice as long as involucre. Involucral bracts narrow, 1.5—2.5(3.0) mm wide; 12 outer bracts more or less appressed; middle and inner slightly recurved, geniculate, with oblique spine saber-like curved; innermost involucral bracts colored at tips. Corolla light-purple, irregularly incised. Achenes immature, large, gray, oblong; pappus hairs distinctly shorter than corolla. Flowering June to August. Rocky, stony mountain slopes.—Soviet Central Asia: Tien Shan, Dzhungaria-Tarbagatai. Endemic. Described from banks of Ketmen River. Type in Leningrad. Note. Among the collections of Aitchison from Afghanistan (Kuroma Valley) one specimen greatly resembles this species. Owing to the limited material we are refraining from referring this specimen to C. schischkinii. 4. C. coloratus Tamamsch. in Bot..Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XV (1953) 390.—C. nutans var. coloratus Winkl. ex O. and B. Fedtsch. Perech. Rast. Turk. IV (1911) 277, nomen. Perennial. Stem branched almost from base, 80-100 cm high, ribbed, slightly sulcate between ribs, covered with very sparse, long, wavy hairs, broadly and discontinuously winged at base, wing lobes rectangular, toothed, spinescent. Leaves quite narrow, oblong, sinuate- toothed; lobes deltoid, with small and large spines; lamina thin, decur- rent, green on both sides, covered with long, very sparse, undulate hairs mainly on veins. Peduncles to 5—7 cm long, white-tomentose, not winged, with solitary capitulum. Capitula nodding, quite large, 3.54.0 cm in dia. Involucre more or less broadly oblong; outer bracts in young capitula horizontal, inner ones more or less erect, appressed; bracts in opened capitula oblong-lanceolate, geniculate in middle, sharply sulcate inside in upper part, with long recurved cusp, yellow- ish in lower part, green in middle, bright purple above, particularly innermost puberulent outside, finely ciliate; narrow spinulose bracts below capitulum at base of involucre. Corolla stramineous or pink; corolla limb irregularly incised; pappus of white silky hairs with fine teeth, distinctly shorter than corolla. Flowering July to August. Foothills, on dry slopes.—Soviet Central Asia: Tien Shan, Pamiro- Alai Region, Kyzyl-Kum, Syr-Darya. General distribution: Dzhungaria- Kashgaria. Described from Alai Mountain spurs, upper reaches of Shakhimardan River. Type in Leningrad. Series 2. Acanthocephali Tamamsch.—Grayish or densely white- tomentose plants; involucral bracts narrow, spiny, erect, divergent or curved saber-like, lacking constriction in middle, not geniculate. Cau- casian species, of which one species occurs in Transcaucasia and northern Iran. 13 5. C. acanthocephalus C.A.M. Verzeichn. (1831) 71; DC. Prodr. VI, 622; Ldb. FI. Ross. II, 718; Boiss. Fl. or. III, 516; Grossh. FI. Kavk. IV, 174; Isaev in Fl. Azerb. VIII, 381. Perennial. Stem branched, sulcate, arachnoid-hairy, broadly winged; deltoid wing lobes unevenly sinuate on margin, with yellowish, stiff, unequal spines, middle lobe with distinct vein terminating in long, to 1.0-1.5 cm long, yellow spine; sometimes wings narrowed below capitulum or abruptly narrowed into stiff, long, acute cusp. Leaves oblong, decurrent, densely hairy beneath; pinnately lobed; lobes large, distant, irregularly toothed, with long stiff spines. Peduncles very short with solitary capitulum, winged and spiny. Capitula large, 3.0—4.5 cm long, subglobose. Involucral bracts densely grayish-arachnoid-hairy, subulate; outer and middle bracts more or less recurved, somewhat flat in lower half, grooved inside in upper half, deltoid in transection, hard, spinescent, with prominent midrib and 2 distinct lateral veins, with pseudo-glandular hairs in lower part; inner bracts not spiny, slightly pinkish, linear; corolla purple, 20-25 mm long, irregularly incised, distinctly exceeding inner involucral bracts. Achenes oblong, yellow- ish-brown, with darker and quite distinct longitudinal and weaker trans- verse lines, tubercle indistinctly lobed; pappus of white fragile hairs, considerably shorter than corolla. Flowering July to August. Alpine zone, in meadows.—Caucasus: Ciscaucasia, Eastern Caucasus. Endemic. Described from Tfandag Mountains. Type in Leningrad. 6. C. hystrix C.A.M. Verzeichn. (1831) 71; DC. Prodr. VI, 622; Ldb. Fl. Ross. II, 719; Boiss. Fl. or. III, 517; Grossh. Fl. Kavk. IV, 175; Isaev in Fl. Azerb. VIII, 382. Biennial or perennial. Whole plant light-colored, almost whitish from dense pubescence. Stem 80-100 cm high, branching from middle, deeply sulcate, wingless below, arachnoid-hairy-pubescent, white-to- mentose above including peduncles, narrowly winged up to capitulum, lobes of stem wings unevenly incised, with sharp unequal spines. Leaves slightly pubescent above, white-tomentose beneath; lamina of lower leaves oblong-ovate, 10—12 cm long, 4—5 cm wide, pinnately parted; lobes broadly deltoid, sinuate, with short, thin spine along margin; middle leaves narrower, decurrent, pinnately lobed, spinose-toothed and ciliate, narrowed toward apex, terminal lobe lanceolate, spinescent. Capitula pedunculate, solitary, erect, quite large, to 3.0-3.5 cm in dia, cylindrical before flowering; peduncles wingless below capitulum. Involucre only up to opening arachnoid-lanate; outermost bracts some- what shorter than others, appressed; other bracts to 2-3 cm long, saber-like recurved, more or less subulate, broad at base, spinescent, 14 with prominent, hard midrib beneath, stiff; innermost bracts linear, not stiff, not spinose, with 3 distinct veins beneath, colored above. Corolla light purple, to 38-40 mm long. Pappus white, pinkish-golden, more or less silky, distinctly longer than corolla tube. Achenes 4-5 mm long, lustrous, grayish-olive, with distinct longitudinal brownish lines and weak transverse striation. Flowering August to September (Plate II, Fig. 3). Dry high-mountain meadows, up to 2,000-—3,000 m.—Caucasus: Eastern Transcaucasia, Talysh. General distribution: Iran. Described from Talysh. Type in Leningrad. 7. C. poliochrus Trautv. in Tr. Peterb. Bot. Sada, III (1875) 272; Boiss. Fl. or. Suppl. 307; Grossh. Fl. Kavk. IV, 176 (sphalm. C. polyochrus). Biennial. Plant 25-35 cm high. Stem corymbosely branched, arach- noid-pubescent, narrowly winged. Leaves oblong, decurrent, more or less pinnately lobed, ovate-deltoid; lobes abruptly narrowed, spinescent, rarely sparsely arachnoid above, densely arachnoid white-tomentose beneath. Peduncles narrowly winged, usually bearing solitary capitu- lum; capitula 2-3 cm long. Involucre subglabrous or slightly pubes- cent; involucral bracts somewhat recurved till full bloom, subulately spinose from narrow lanceolate base, with prominent vein in mature capitula, and long but straight apical spines; outer bracts almost 1/2 as long as middle, hairs in lower part on dorsal surface pseudo-glandular; inner bracts subulately narrowed, but not spinose, with 3 prominent veins, finely ciliate, colored. Corolla purple, 12-13 mm long. Pappus longer than corolla tube excluding throat. Achenes 5.0—5.5 mm long, narrow, grayish or brownish, with few indistinct, darker, longitudinal striations, narrowed below and slightly oblique, with small 5-lobed tubercle above. Flowering July to August. High-mountain meadows.—Caucasus: Dagestan. Endemic. De- scribed from Akhta and Murukh. Type in Leningrad. Series 3. Qnopordioides Tamamsch.—Subglabrous, usually light green, rarely dark green plants with oblong or globose capitula; in- volucral bracts more or less herbaceous with variously curved spines, but not geniculate. Species of this series are found mainly in different regions of Transcaucasia. They are, however, also found in the northern part of Iran. 8. C. hajastanicus Tamamsch. in Addenda XXVII, 599. Perennial. Plant bluish-green, 40(50)-100 cm high. Stem divaricately branched from base; branches discontinuously winged, S VIZ ‘ BWGi WZ \\ ae —= = __. aN y eS \\ ZY fin Wy VY, WZ , AZ ty 2 Vi dj Gee WZ 4 Ww > i \Wiea Wes xs S he \\ NS MNS “A 15 Plate I. 1—Carduus furiosus Tamamsch.; 2—C. hajastanicus Tamamsch.; 3—C. onopordioides Fisch. ex MB. 16 spiny, more or less ribbed-sulcate like main stem, slightly arachnoid- pubescent, wingless in upper part of peduncles, with few prickly cilia and sparsely pubescent. Leaves narrow, oblanceolate, sessile, shallow and unevenly pinnately lobed, lobes deltoid with long, stiff, yellow spine to 8-9 mm-long, bluish-green, both surfaces with fine pseudo- glandular hairs and distinct veins, lighter than lamina. Capitula large, solitary, 3.0—3.5 cm long and 2.5—3.0 cm in dia, oblong-globose, pe- dunculate, slightly nodding just below capitula. Involucre subglobose, in lower part slightly appressed around peduncle; involucral bracts unequal, broad and appressed at base, subglabrous or pubescent only along margin and inside; outer bracts more or less brownish, almost horizontally deflected, cuspidate, on margin arachnoid-pubescent, dor- sally with pseudo-glandular hairs and prominent thick midrib; inner bracts narrow, purple, with white, pointed tip, ciliolate; florets longer than involucre, dark purple or reddish-purple. Corolla 19.0-19.3 mm long; corolla tube considerable shorter than limb including throat. Pappus 3/4 as long as corolla; pappus bristles distinctly scabrous. Achenes oblong, straight or somewhat curved, slightly narrowed be- low, including tubercle 6 mm long, light-brown, with lighter, thin, longitudinal lines and darker, transverse striations. Flowering May to August. (Plate I, Fig. 2.) Grassy, stony mountain slopes.—Caucasus: Southern Transcaucasia. Endemic. Described from shores of Lake Sevan. Type in Leningrad. 9. C. furiosus Tamamsch. in Addenda XXVII, 599.— C. onopordioides auct. cauc. nonnull. non. Fisch. ex MB. Biennial. Plant highly spiny, 30-35 cm high, light-green. Stem single, densely leafy, ribbed-sulcate, narrowly spiny-winged; wing lobes deltoid, with stiff, yellowish spines 5-15 mm long. Leaves slightly coriaceous, lighter above, with thick, raised midrib beneath, sparsely arachnoid-pubescent; lower and middle leaves almost equal, upper leaves somewhat reduced; all leaves decurrent, 7-12 cm long, 2.0—2.5 cm wide, lanceolate, pinnately shallow-lobed, with short deltoid spiny lobes. Peduncles below capitula usually spiny winged, erect, less often wingless but spiny. Capitula flat-globose, 3.0—3.5 cm long, 3—4 cm in dia; except upper undeveloped ones, capitula axillary in middle of stem. Involucre subglabrous or sometimes slightly arachnoid-pubes- cent; all bracts longer than pappus and corolla; outer and middle bracts stiff, becoming very hard toward end of flowering, abruptly narrowed from somewhat broad base into sulcate tip, with long, stiff, almost horizontal or recurved, divergent spines, latter 1.5—2 times as long as involucral bracts, yellow or purple. Corolla dark pink, 22—23 mm long; 17 corolla tube shorter than limb including throat. Achenes thick, fusi- form, including tubercle 6 mm long, dull gray, with many longitudinal darker striations; pappus 1.5—2 times as long as achene. Flowering June to July. (Plate I, Fig. 1.) Gravel beds, dry stony slopes.—Caucasus: Eastern Transcaucasia (southeastern part), Talysh. General distribution: Northern Iran, whence described (Badalan). Type in Leningrad. Note. The-diagram of “C. onopordioides Fisch.” in Plate XX XIX in Volume VIII of the Flora Azerbaidzhana [Flora of Azerbaidzhan] probably refers to this species. 10. C. onopordioides Fisch. ex MB. FI. taur.-cauc. III (1819) 552; DC. Prodr. VI, 622; Ldb. FI. Ross. II, 719; Boiss. Fl. or. III, 516; Grossh. Fl. Kavk. IV, 174; Kapeller in Fl. Gruzii, VIII, 465. Annual or biennial. Plant light colored or yellowish-green, 30—50 cm high. Stem simple below, weakly branched above, sulcate, subglabrous, unevenly winged; wing lobes deltoid, spinose on margin, with long, straight, less often curved spine in middle. Leaves thin, thick or slightly coriaceous, glabrous or with thin isolated hairs only on midrib, sessile, somewhat unevenly and pinnately lobed, oblong or oblong-lanceolate, strongly spinose, with stiff thin spines on margin of leaf lobes. Peduncles branciied at apex, short, usually with solitary capitula, wingless below capitulum, grayish-tomentose or with short, small, discontinuous wings or spines. Capitula oblong or ovate-cylin- drical, 2.5—3.0 cm long. Involucre subglabrous, slightly arachnoid pubescent only when young; all bracts more or less herbaceous, not very stiff, spinescent; outer bracts recurved, sulcate inside with lighter midrib, inner almost erect, as long as florets or somewhat shorter, membranous, acute but not spinescent, finely ciliate, with 3 or 5 dis- tinct veins in middle. Corolla bluish pink or darker, 20.0—-20.3 mm long; corolla tube excluding throat 9.0—9.3 mm long. Pappus of very thin silky hairs, up to 1/2 as long as corolla. Achenes grayish or light- brown, quite compressed, with indistinct, thin, longitudinal and many transverse striations, thick, 5-lobed tubercle somewhat narrowed be- low, along with tubercle 5 mm long. Flowering June to August. (Plate Larig: 3.) Dry stony places.—Caucasus: Eastern Transcaucasia (central part). Endemic. Described from vicinity of Tbilisi from Wilhelms’ collec- tions. Type in Leningrad. Note. This species does not grow in Soviet Central Asia. All her- barium specimens seen by me identified as C. onopordioides belong to C. schischkinii. This remark pertains also to literature references to the occurrence of C. onopordioides in Soviet Central Asia. In Volume 20 18 VIII of Flora Azerbaidzhana [Flora of Azerbaidzhan], Ya. Isaev in- cluded C. atropatanicus Sosn. in the synonymy of C. onopordioides. The history of the former species is as follows: D.I. Sosnowsky him- self did not publish the description of this species, but only gave it the name. The first description, by Grossheim, is in Flora Kavkaza [Flora of the Caucasus], Vol. IV (1934). However, to judge from herbarium specimens, Grossheim understood quite different things under the label “atropatanicus,” which only with great difficulty could be re- ferred to the given series or even the section. Sosnowsky himself, in his time (in litt.), found it difficult to determine the affinity of C. atropatanicus. Guided by the autotype with Sosnowsky’s signature, I came to the conclusion that C. atropatanicus undoubtedly is close to C. onopordioides but is a separate more southern race. 11. C. atropatanicus Sosn. ex Grossh. Fl. Kavk. IV (1934) 175 p. p.—C. onopordioides auct. p. p. non Fisch. ex MB.; Isaev in FI. Azerb. VIII (1960) 382. Annual or biennial. Plants to 30-35 cm high, subglabrous, light colored, or yellowish-green. Stems spiny-winged, slightly sulcate. Leaves oblong below, lanceolate above, pinnately lobed, with spinu- lose deltoid lobes and thick, brighter midrib more prominent dorsally; whole plant considerably less spiny than the preceding two species. Capitula small, obovate, 4 cm in dia above, 2.5 cm below. Involucre broadly obconical, somewhat foveate below; involucral bracts at tip subglabrous, sparsely arachnoid-tomentose below, light green; inner bracts bright purple at tip, slightly pubescent, weakly spinescent; outer bracts gradually narrowed from oblong base, spinescent, recurved. Corolla pale pink, sometimes almost white, scarcely exserted from inner involucral bracts and pappus, 17—18 mm long; corolla tube ex- cluding throat as long as corolla lobes including throat; achenes with tubercle 5.0—-5.2 mm long, 2 mm wide, inequilateral, truncate below, light olive-gray or yellowish with darker, thin, longitudinal and many transverse striations; tubercle sessile, indistinctly lobed. Flowering June. Middle mountain zone, in dry places.—Caucasus: Southern Transcaucasia. Endemic. Described from Nakhichevan. Type in Tbilisi; cotype in Leningrad. Section 2. Carduastrum Tamamsch. in Addenda XXVII, 600.— Sect. Platycephali Rouy, Fl. Fr. IX (1905) 75 p. p.—Sect. Carduus subsect. Pachycephali auct. europ. non-null. p. p.—Capitula large or of average size, depressed globose or more or less oblong; involucral bracts narrow, stiff hard or somewhat soft, distinctly or indistinctly imbricate, appressed or divergent. Plants with spiny-winged or wing- less long peduncles. \\ oie Uy : “ = = = Fae ies =\\ a a ' | VW Na AN Fl Wi | VE < oe HD, 19 Plate II. 1—Carduus uncinatus MB.; 2—C. personata (L.) Jacq. 3—C. hystrix C.A.M.; 4—C. transcaspicus Gand. 21 20 Type of section: C. crispus L. Series 1. Collini Tamamsch.—Involucral bracts narrow, hard, uncinate. Capitula oblong or depressed-globose. The species of this series, widely distributed in the eastern and southeastern part of Western Europe, number about 6—7, three of which we have. 12. C. pseudocollinus (Schmalh.) Klok. in Vizn. Rosl. URSR (1950) 566; Geidemann, Opred. Rast. Mold. SSR, 288.—C. collinus auct. non. Waldst. and Kit.: Ldb. Fl. Ross. II (1846) 721; Boiss. FI. or. III, 519.—C. collinus var. caucasicus C. Koch in Linnaea, XXIV (1851) 394.—C. hamulosus B. pseudocollinus Schmalh. Fl. Yugo-Zap. Ross. (1886) 317; Fl. II, 98; Grossh. Fl. Kavk. IV, 175; Stank. and Tal. Opred. Vyssh. Rast. (1949) 660. Biennial or perennial. Stem simple or branched from middle or above, discontinuously and unevenly winged; wings spinose. Leaves decurrent or with winged petioles; petioles sulcate above, carinate below; lamina sparsely pubescent above, but grayish beneath from denser pubescence; lower leaves almost linear, deeply pinnately lobed, lobes deltoid, 3—5-toothed, spinescent, finely ciliate; middle and upper leaves gradually reduced, lanceolate, spinose; peduncles below capitu- lum wingless, almost white from densely tomentose pubescence, with solitary capitula. Capitula of average size, 1.5—-2.0 cm long. Involucre globose-ovate, toward tip slightly constricted or hemispherical; involu- cral bracts more or less stiff, brownish or purple-violet, narrowly lin- ear, spinescent, in young capitula more or less appressed, in mature capitula uncinate, indistinctly grooved inside; middle bracts dorsally with somewhat prominent midrib, terminating in uncinate spine; inner bracts more or less appressed, with 3 distinct veins. Corolla purple; its limb irregularly incised, distinctly exceeding involucral bracts. Achenes small, about 3.0—3.5 mm long, light-brown with darker longitudinal striations and inconspicuous transverse zigzags between lines; pappus longer than involucral bracts. Flowering June to August. Dry meadows, steppes, on stony and calcareous, sunny slopes.— European Part: Black Sea Region, Upper Dnieper, Middle Dnieper, Upper Dniester; Bessarabia, Lower Don, Lower Volga, Crimea; Caucasus: Ciscaucasia, Southern Transcaucasia (occasional). General distribution: Central Europe. Described from southwest European Part of USSR. Type in Kiev. 13. C. transcaspicus Gand. in Bull. Soc. Bot. France, LXV (1918) 37; Tamamsch. in Fl. Turkm. VII, 237.—C. uncinatus var. transcaspicus Rech. f. in Ann. Nat. Mus. Wien, 14-15 (1943) 16. 22 Zl Perennial. Stem up to | m high, sometimes even taller; branched almost from base, cylindrical, slightly sulcate, subglabrous below, narrowly winged in middle. Lower leaves to 10—15 cm long, gradually tapering below, slightly decurrent, glabrous above or with isolated hairs, somewhat tomentose-pilose beneath, irregularly pinnately lobed with inconspicuous spinules on margin; middle and upper leaves gradu- ally reduced, narrowly lanceolate, slightly pilose above, white-tomen- tose beneath, more prickly than lower leaves. Peduncles to 18-20 cm long, narrowly winged below, slightly sulcate, white-tomentose, termi- nating in large, solitary capitula to 4 cm long. Involucre almost cylin- drical, strongly spinose, longer than florets in young capitula, more than 1/2 as long as corolla at flowering; outer and middle bracts stiff, narrow, uncinately recurved, somewhat carinate outside, grooved in- side, glabrous or sometimes weakly arachnoid-pubescent at base, with pseudo-glandular yellowish hairs outside, finely ciliate inside, spinescent, pinkish at base; inner bracts with gradually less stiff and less recurved spinule, colored; innermost bracts carinate outside like preceding ones, violet. Florets pink; corolla lobes irregularly incised. Pappus snow-white, with hairs longer than corolla tube. Achenes ob- long, light- or dark-brown, 1/2 as long as receptacular bristles. Flow- ering August. (Plate II, Fig. 4.) Slopes and in subalpine zone.—Soviet Central Asia: mountainous Turkmenia. Endemic. Described from Kyzyl-Arvat. Type in Paris. Economic Importance. It can be recommended as an ornamental plant because of the pink flowers in rather large capitula, nice color of the involucre, and bright white peduncles. 14. C. hamulosus Ehrh. Beitr. Naturk. VII (1792) 166; Waldst. and Kit. Descr. et icon. pl. rar. Hung. III (1812) 258; MB. FI. taur.- cauc. II, 268; Bess. Enum. pl. Volhyn. 31; DC. Prodr. VI, 623; C. Koch in Linnaea, XVII, 42; Ldb. Fl. Ross. II, 720; Boiss. Fl. or. III, $17; Grossh. Fl. Kavk. IV, 175; Klok. in Vizn. Rosl. URSR, 566; Kapeller in Fl. Gruzii, VIII, 466.—Ic.: Rechb. Ic. Fl. Germ. XV, tab. 875, II].—Exs.: Fl. exs. austr.-hung. No. 1782. Biennial or perennial. Plant to 60-70 cm high. Stems 2-3, branched, cylindrical, sulcate, winged to very tip, less often wings not reaching to capitula; wings narrow, interrupted, pubescent together with stem, with long articulate hairs. Lower leaves with winged petioles crowded at stem base; all leaves dull grayish-green from long hairs, particularly dense on veins and small pseudo-glandular hairs; upper leaves oblong- ovate with 5—7 pinnate lobes, spinulose-ciliate; peduncles often nar- rowly winged, spinose, white-tomentose. Capitula large, 3—4(5) cm in dia, subglobose or somewhat oblong. Involucre subglabrous or with small pseudo-glandular hairs; involucral bracts narrow, lanceolate, more 23 22 or less deltoid, very stiff; outermost bracts more or less appressed middle (particularly in unopened capitula) uncinately recurved in up- per half, dark inside, with prominent midrib, on outer side covered with fine pseudo-glandular hairs; innermost bracts subulately narrowed, membranous, green in lower part, purple above, somewhat pubescent, with 3 distinct veins, like middle uncinately recurved. Corolla purple, irregularly incised into lobes. Achenes lustrous, olive-reddish-brown, oblong, slightly compressed; at base indistinctly ribbed, with fine, trans- verse, pilose striations and quite large-lobed tubercle at tip; pappus longer than inner involucral bracts, to 2/3 as long as corolla. Flower- ing June to August. Dry, often weedy places, dry shrubby thickets, in lower and middle mountain zones.—European Part: Bessarabia, Black Sea Region, Crimea; Caucasus: almost all regions particularly the southern ones. General distribution: southern and Central Europe, Balkans-Asia Mi- nor. Described from southern Europe. Type in Berlin(?). Series 2. Crispi Tamamsch.—Involucral bracts stiff but not hard, divergent, sometimes clearly uncinately recurved or curved in middle at wide angle and obliquely upright. The species of this series are widely distributed throughout Eu- rope, Asia, and Africa. 15. C. crispus L. Sp. pl. (1753) 821; DC. Prodr. VI, 623; Ldb. FI. Ross. II, 721; Boiss. Fl. or. II, 520; Schmalh. FI. II, 97; Hegi, Ill. FI. VI, 2, 860; Grossh. Fl. Kavk. IV, 176; Klok. in Vizn. Rosl. URSR, 567; Kapller in Fl. Gruzii, VII, 471.—C. crispus var. albiflora Bordz. in Izv. Kievsk. Bot. Sada, XII—XIII (1931) 141.—C. crispus var. comosus M. Pop. Fl. Sredn. Sib. II (1959) 752, diagn. ross.—Ic.: Hegi, Ill. Fl. VI, 2, Taf. 270, fig. 4. Perennial. Plant 50-120 cm high, dark green. Stem cylindrical, sulcate, finely pubescent, winged to very apex and sharply spinulose, branched from middle, above or only at apex. Leaves ovate or oblong- ovate, sparsely pilose above, grayish-tomentose or more sparsely pu- bescent beneath; lower leaves to 20 cm long and 10-15 cm wide, with winged petiole, to 10-12 cm long, sometimes less pubescent than up- per leaves, unevenly and sinuately emarginate-lobed-toothed and pin- nately lobed; lobes deltoid or ovate, ciliate-spinulose; middle and upper leaves gradually reduced, oblong or lanceolate, ciliate-spinose with distinct veins and larger terminal lobe; peduncles axillary, usually 10-15 cm long, white-tomentose or less densely pubescent, mostly to very tip prickly-winged or simply spinose. Capitula 2-3 cm long, in terminal clusters of 3—4, usually densely clustered, very rarely 2 or 24 23 solitary and sessile in leaf axils, ovate or more or less globose. Involu- cre mostly broadly ovate, shorter than florets; bracts in young capitula straight, divergent, outer bracts linear-subulate at flowering, slightly recurved, more or less hard, appressed at bend, other bracts inequal, highly recurved, as a result involucre looking shaggy; inner bracts thin, almost membranous, appressed. Corolla usually purple, very rarely lighter or white. Achenes small, about 3 mm long, grayish, with thin, longitudinal darker striations and lacking stalk; tubercle slightly lobed; pappus shorter than corolla, scarcely exceeding involucre. Flowering July to September. Weed—in fields, among low shrubs, in forests in moist shady places.—European Part: almost all regions; Caucasus: all regions; Western and Eastern Siberia: almost all regions; Far East: all regions; Soviet Central Asia: rarely. General distribution: all of Europe, Arme- nia and Kurdistan, Iran, North America (introduced). Described from Northern Europe. Type in London. Note. A highly polymorphic species. The variation of the charac- ters is particularly striking in the vegetative parts: length of stem; form of lamina which sometimes is almost entire, other times deeply pin- nately lobed, with the upper surface almost glabrous or roughly to finely pilose; spines sometimes strong, other times weak; involucral bracts with veins distinct or indistinct. However, in Mongolia, Japan and China this species is so different in appearance that it can hardly be considered C. crispus; in the future, plants from these regions re- ferred to in the literature as C. crispus probably should be treated as separate races, which I have already done for Mongolia (see Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XV). Possibly, the form reported by M.G. Popov from Central Siberia under the name var. comosus, should be treated as a separate race. Economic Importance. It is nectariferous plant. 16. C. acanthoides L. Sp. pl. (1753) 821; DC. Prodr. VI, 623; Ldb. Fl. Ross. II, 719; Boiss. Fl. or. III, 518; Grossh. Fl. Kavk. IV, 175; Klok. in Vizn. Rosl. URSR, 567; Kapeller in Fl. Gruzu, VIII, 467.—C. acanthoides 6. thessalus Boiss. FI. or. III (1875) 518.—C. thessalus Boiss. and Heldr. in Boiss. Diagn. pl. or. ser. II, 3 (1856) 46; Grossh. Fl. Kavk. IV, 175.—Ic.: Rchb. Ic. Fl. Germ. XV, tab. 873. Perennial. Plant to 60-70 cm high, light green. Stem cylindrical, finely sulcate, narrowly spiny winged to very tip, simple or branched at apex, sparsely pubescent with long articulate hairs. Lower leaves early deciduous, middle and upper decurrent, oblong or oblong-ovate, 15-20 cm long, with very sparse hairs on both surfaces, pinnately lobed, in turn with unequal 3 or 5, quite remote lobes bearing sharp 2 Nn 24 yellow spines, usually with larger, lanceolate, shallow emarginate ter- minal lobe. Peduncles short, usually narrowly winged to capitula, spi- nose, with solitary capitula, very rarely capitula few. Capitula average sized, 3 cm in dia, subglobose. Involucre subglabrous, considerably shorter than florets; involucral bracts basally somewhat broad, later gradually becoming linear-subulate, spinescent, with prominent mid- rib, slightly divergent in upper part, sometimes outer bracts horizontal; inner scarious, somewhat colored, sometimes with curved, acute apex, with 3 distinct veins. Corolla pink or dark-pink, at tip irregularly in- cised, in ray florets corolla tube bent. Achene small, 3 mm long, 1 mm wide, slightly compressed, grayish-olive, lacking prominent longitudi- nal striations, but with weak transverse lines and a rather persistent tubercle; pappus bristles lacking distinct serration, longer than inner involucral bracts and covering 3/4 of corolla. Flowering July to Au- gust. Fields, roadsides, in desert and stony places——European Part: almost all regions, except the northernmost ones; Caucasus: almost all regions; Western Siberia: Upper Tobol. General distribution: northern Europe, Atlantic Europe, Mediterranean, Balkans-Asia Minor. De- scribed from western Europe. Type in London. Economic Importance. Young stems cleared of spines are used as raw vegetable in some countries. 17. C. navaschinii Bordz. in Izv. Kievsk. Bot. Sada, XII—XIII (1931) 141; Grossh. Fl. Kavk. IV, 176. Biennial. Stem to 75-100 cm high, ascending, branched from middle, finely sulcate, with long, multicellular hairs or arachnoid-hairy, winged; wings deeply lobed with spiny-toothed lobes. Leaves arach- noid-pubescent above, rarely beneath, particularly on veins; middle leaves sessile, oblong or oblong-elliptical, pinnately lobed; lobes ovate or deltoid-ovate, in turn 2—4 lobed or toothed, spinescent, with up to 3.5 cm-long spine, decurrent, semiamplexicaul; upper leaves acute with highly spiny lobes. Capitula subglobose, to 2-3 cm long, in clusters of 2-3, rarely solitary or many (7); peduncles short, white-tomentose, often lacking wings at tip. Involucral bracts densely arachnoid-hairy, imbricate, appressed or divergent-recurved in upper part, very narrow, with prominent midrib extended into thin spine; outer bracts oblong- lanceolate; innermost bracts membranous, oblong. Corolla white, to 15 cm long, significantly longer than involucre. Achenes olive, slightly compressed, oblong, narrowed downward, 4.5 mm long, to 1.5 mm wide; pappus white, to 12-14 mm long. Flowering July to August. High-mountain plant growing up to 3000 m.—Caucasus: Southern Transcaucasia. Endemic. Described from slopes of Aragats (Alagez) Mountain. Type in Kiev; paratype in Leningrad. 2 fon) 25 Note. This species was described by E.I. Bordzilovsky from his own collections. The species was collected again by the author of these lines from the same and other adjoining places in 1933. Bordzilovsky connected this species with C. flavescens C. Koch, which was described from Turkish Armenia from the vicinity of the town of Ardagan. Koch, comparing C. flavescens with C. hamulosus and C. acanthoides, considered that the difference of C. flavescens lies in its yellowish flowers. But Boissier linked C. flavescens with C. collinus and C. adpressus, noting that it is distinguished from the first species by short peduncles, winged almost up to capitula, and from the second species by the nature and size of the involucral bracts. From a more comprehensive study of C. navaschinii, based on the specimens avail- able in Erevan and identified by Bordzilovsky himself, and from the specimens of C. flavescens of Koch in the General Herbarium of Botanical Institute of the Academy of Sciences of the USSR, one can conclude that these species are very close to each other and are cer- tainly related to C. crispus, but they also show some similarity with C. uncinatus and C. hamulosus. In the herbarium of the Botanical Insti- tute of the Academy of Sciences of the USSR, there are two sheets from the collection of Meyer, which had been referred to C. uncinatus. Both specimens were collected by Frick in 1847 from Alagez Moun- tain, 1.e., in the same region from which C. navaschinii was also de- scribed. On one sheet there is a label where in Meyer’s handwriting is written “C. crispus flore albo,” then “crispus” had been crossed out and “uncinatus” written in by Meyer. In fact, both specimens should be referred to Bordzilovsky’s C. navaschinii. In this way, the species established by Bordzilovsky in 1931 was, for the first time, collected more than 100 years ago, and even at that time was cited by Meyer as a puzzling form, conforming neither to C. crispus not to C. uncinatus. 18. C. uncinatus MB. FI. taur.-cauc. III (1819) 553; DC. Prodr. VI, 623; Ldb. Fl. Ross. II, 720; Boiss. Fl. or. II, 518; Grossh. FI. Kavk. IV, 175; Klok. in Vizn. Rosl. URSR, 567.—C. uncinatus var. eu-uncinatus Rech. f. in Ann. Nat. Mus. Wien. 14-15 (1943) 16. Biennial. Stem straight, branched above or simple, ribbed-sulcate, arachnoid-hairy-pubescent, narrowly winged, wings interrupted, with small ovate lobes, strongly spinose along margin. Leaves arachnoid- hairy-pubescent above, tomentose beneath; lower leaves with winged petioles, 20—25 cm long, broad, pinnately lobed; lobes deltoid, reduc- ing downward; middle and upper leaves oblong, oblong-linear or lin- ear, slightly emarginate, with short sharp spine; peduncles usually long, tomentose-lanate, discontinuously winged almost to capitulum, less often wingless below capitula. Capitula solitary, large, 3.5—4.0 cm in dia, Zi 26 subglobose. Involucre hemispherical, constricted in middle, grayish- arachnoid-hairy; outer bracts considerably shorter than others, ap- pressed, somewhat broader than middle; middle bracts linear-subulate, appressed in lower half, uncinately recurved in upper half, but not stiff, with short spine; innermost bracts not spiny, some erect, some slightly recurved. Corolla dark pink or orange-pink, less often pinkish- purple. Pappus of white hairs, shorter than corolla. Achenes to 2.0—2.5 mm long, grayish-olive, slightly angular, with thin, longitudinal, brown and transverse sinuate striations, with unsegmented, indistinctly angu- lar tubercle. Flowering May to August. (Plate II, Fig. 1.) Steppes, hill-steppes, sometimes in lower mountain zone.—Euro- pean Part: Crimea, Lower Don, Lower Volga; Caucasus: Ciscaucasia, Dagestan, Western Transcaucasia. General distribution: Central Eu- rope, Asia Minor. Described from Crimea. Type in Leningrad. Note. In the Herbarium of the Botanical Institute of the Academy of Sciences of the USSR, there is one specimen of this species from Karpinsky’s collection, borrowed from Wilhelms’ collection from Abkhazia. This species is not reported from Abkhazia; however its presence there is not ruled out. Economic Importance. An ornamental plant because of the beau- tiful color of the capitula. 19. C. stenocephalus Tamamsch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVI (1954) 468.—C. uncinatus var. parviceps Iljin in schedis. Biennial. Stem ribbed, between ribs sulcate, slightly pubescent below along grooves, densely grayish-tomentose above, strongly branched, more or less uniformly winged. Lower leaves early decidu- ous, upper linear lanceolate, pinnately lobed, lobes deltoid, 3—5 toothed, terminating in sharp, short spine, bright green above with sparse, long, flexuous hairs, grayish-tomentose beneath, with prominent midrib; peduncles and upper part of stem densely white-tomentose, wingless, but often with cluster of spines below capitulum or with very narrowly lanceolate, prickly subtending leaf. Capitula not large, 2.5—3.0 cm long, 1.5 cm in dia. Involucre basally roundish, more or less narrowly ob- long, shorter than florets; involucral bracts subglabrous or slightly arachnoid-pubescent, mainly in distal half; outer bracts linear-lanceolate or subulate, terminating in stiff spine bent at right or wide angle, with prominent midrib and scarcely visible lateral veins; inner bracts ap- pressed, subulately spinescent, sparsely ciliate, pale with very distinct, sparsely pubescent midrib. Corolla limb intensely pink or lighter, irregularly incised. Achenes oblong, to 6 mm long, somewhat angular, slightly oblique, white, with brownish longitudinal striations but 28 2, lacking transverse pattern, with sessile, cylindrical, usually 5-lobed tubercle at apex; pappus much shorter than corolla. Flowering June to August. Steppes and low-lying areas.—European Part: Trans-Volga, Lower Volga; Western Siberia: Upper Tobol (southwestern part); Soviet Cen- tral Asia: Aralo-Caspian Region (northwestern part). Endemic. De- scribed from Krasnoarmeisk (former Sarepta). Type in Leningrad. 20. C. personata (L.) Jacq. Fl. Austr. [V (1776) 25; Gaertn. De fruct. II, 378.—C. arctioides Vill. Hist. pl. Dauph. II (1789) 22, non Willd.—Arctium personata L. Sp. pl. (1753) 816.—Ie.: Jacq. 1. c. tab. 348. Perennial. Stem ribbed-sulcate, up to 100 cm high, grayish-brown, narrow-winged on ribs; wings spinulose along margin; somewhat pu- bescent. Lower leaves 20—25 cm long, on equally long petioles; middle leaves 10-15 cm long, 11-12 cm wide; upper leaves gradually re- duced; lamina thin, dark green and subglabrous above, grayish-tomen- tose beneath over a larger part, less often gray from scattered arach- noid pubescence, pinnately veined with many thin veins, entire, less often pinnately parted, ovate, coarsely sinuate-toothed, sometimes ser- rate; teeth ciliate and prickly; peduncles terminal and axillary, spinu- lose to capitula, white-tomentose, ciliate, bearing many (3-4), rarely 1 capitulum, 1—3 subtending leaves below capitulum. Capitula medium- sized, 2.5 cm long, more or less clustered. Involucre shorter than flo- rets, glabrous; outer involucral bracts subulately extended upward from broader base, ciliate; middle and innermost bracts linear, gradually narrowed into subulate tips; all bracts with distinct veins on inner side, slightly recurved, with slightly twisted non-prickly cusp. Corolla pink- ish-violet or pale purple, limb irregularly incised, 1.0—1.5 cm long, slightly longer than pinkish-white anthers. Achenes 4 mm long, 1 mm wide, erect or slightly oblique, tapering toward base, often around tubercle with scar representing fallen corolla tube; pappus consider- ably shorter than corolla. Plant resembling burdock in appearance. Flowering July to August. (Plate II, Fig. 2.) Shady places, in spruce forests, among mountain shrubs.—Euro- pean Part: Upper Dniester. General distribution: northern Europe, At- lantic Europe. Described from western Europe. Type in London. Note. This species is highly variable in the shape of the lower leaves. Some European authors denote the forms with entire leaves as f. simplicifolius, and with pinnately lobed leaves as f. pinnatisectus, Series 3. Deflorati Tamamsch.—Almost nonprickly plants; peduncles long, wingless, lacking spines, densely grayish- or pinkish- 29 28 white pubescent; capitula medium-sized, 2.0—2.5—3.0 cm in dia, more or less flat-globose or oblong-globose. This series is richly represented in Western Europe; some species are characteristic only of the flora of the Caucasus. 21. C. glaucus Baumg. Enum. stirp. Transsilv. IT (1816) 58; Klok. in Vizn. Rosl. URSR, 565.—C. defloratus var. glaucus Beck, Fl. Nied. Osterr. II (1893) 1233, Gugler in Mitteil. Bayer. Bot. Ges. II, 139.— C. defloratus ssp. glaucus (Baumg.) Hegi, Ill. Fl. VI, 2 (1929) 851.— Ic.: Hegi, loc. cit. fig. 545. Perennial. Stem 15—75 cm high, erect, usually simple below, some- times 2-3 times branched from middle, with narrow, usually unlobed wings, almost lacking leaves for more than 2/3 height or with small leaves. Lower leaves thick, grayish-green on both sides, with bluish tinge, obovate, obtuse, undivided, covered with sharp-pointed, thin, stiff cilia at apex; upper leaves small, undivided, less often slightly lobed, linear-lanceolate; peduncles not ribbed, cylindrical, somewhat virgate, appressed grayish-tomentose with 1, less often 2—3 capitula. Capitula drooping at flowering. Involucre 1.0—1.5 cm in dia below, hemispherical; outer involucral bracts small; middle much longer, 4 times as long as wide, abruptly narrowed at apex to cartilaginous spinule. Corolla lilac or pinkish carmine, apically irregularly incised. Achenes 4 mm long, slightly compressed laterally, yellowish-brown, with distinct transverse striations, obovate; pappus considerably shorter than corolla. Flowering June to September. Mountains from 600 to 2,000 m, on stony limestone soils.—Euro- pean Part: Upper Dniester. General distribution: central and southern Europe. Described from Austria. Type in Vienna. 22. C. adpressus C.A.M. Verzeichn. (1831) 71; DC. Prodr. VI, 627; Ldb. Fl. Ross. II, 722; Boiss. Fl. or. III, 518 p. p.; Grossh. FI. Kavk. IV, 176. Biennial. Stem finely sulcate, white-arachnoid-hairy, weakly branched, winged almost to tip; wing lobes small, deltoid, spinulose, arachnoid-hairy-lanate. Leaves decurrent, oblong, slightly and irregu- larly lobed, lobes finely spinose; lamina thin, light-green, lanate- tomentose beneath, with distinct veins; peduncles 2-3, not winged, white-tomentose, each with solitary medium-sized capitulum. Involu- cre not prickly; involucral bracts distinctly imbricate, appressed, un- equal; outer bracts short, ovate, abruptly terminating in inconspicuous spinule; inner long, linear, scarious in upper part, with slightly diver- gent tip; all bracts arachnoid-hairy gray-pubescent along margin. Co- rolla pink or light-purple. Achenes grayish or light brown, 4 mm long, 30 29 with 5-angled tubercle at apex, sometimes with remnants of style, more or less obovate; pappus 1/2 as long as corolla. Flowering August to September. Mountain plant of alpine meadows and pastures.—Caucasus: Glavnyi [Main] Range (Eastern, Central). Endemic. Described from Kaishaur from Gutgora. Type in Leningrad. Note. This is quite a rare plant. The report that this species is widely distributed is based on the fact that it is easily confused with C. multijugus, from which it is well distinguished by the appressed, imbricate, arachnoid-pubescent involucral bracts. 23. C. multijugus C. Koch in Linnaea, XXIV (1851) 392.—C. laciniatus Ldb. FI. Ross. II (1846) 722; Tamamsch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XV, 385; Kapeller in Fl. Gruzii, VIII, 468, non Lam. (1817).—C. colchicus Alb. Prodr. Fl. Colch. (1895) 142.—C. kolakovskyi Sosn. in Vestn. Tifl. Bot. Sada, XLIX (1920) 37.—C. innocuus Kolak. Fl. Abkhazii IV (1949) 267 descr. ross. et auct. georg. non C.A.M. in schedis. Biennial. Stem ribbed-sulcate, thickened below with very sparse arachnoid pubescence, corymbosely branched above, broadly winged almost to tip; lobes 1-2 toothed, deltoid, with thin sharp spinules. Leaves oblong-ovate, deeply pinnately parted, terminal lobe thrice, pinnately parted, spinose-ciliate, short-spinescent, glabrous or finely arachnoid-pubescent above; peduncles mostly with solitary capitulum wingless, finely, sparsely or densely pubescent. Capitula medium-sized, with subglabrous involucre. Outer and middle involucral bracts linear, cuspidate, with distinct midrib, in upper half recurved; inner bracts acute, with scarious, colored tip, midrib distinct only in lower half. Corolla light purple. Achenes gray, with darker, fine, longitudinal stria- tions and small 5-angled sessile tubercle; pappus of white hairs, 1/2 as long as corolla or longer. Flowering July to August. Low-lying areas to mountain meadows and forest glades.— Caucasus: almost all regions. Endemic. Described from Makharadze (Ozurgeta). Type in Leningrad. Note. The C. innocuus of Kolakovsky and Kapeller belongs to this species. The specimens cited by these authors from Panyutin’s collections completely confirm this fact. However, the real C. innocuus, whose authentic specimen, also preserved in Leningrad, was carefully examined by me, is a quite unusual specimen. Collected once in Karabakh (Southern Transcaucasia) by Sovich, this bristle-thistle gives the impression of an anomalous or hybrid form. This specimen has a rosette of quite large (about 17 cm) oblong basal leaves; cauline leaves, 7-8 cm long, hardly up to 6 mm wide in the middle; a lamina slightly 31 30 crenate with hard cilia on the margin, which generally are not charac- teristic of bristle thistles or are quite rare; stem simple, single, with solitary capitulum. The specimens collected by Panyutin from Western Transcaucasia have well developed spines; leaves characteristic for the genus, the width of the smallest leaves exceeding 2 cm; a basal rosette of leaves absent, and stem branched near tip; involucral bracts and capitulum as a whole quite similar to those in C. multijugus. C.A. Meyer proposed the name C. innocuus, but refrained from publishing this species; his description was appended to Sovich’s her- barium sheet. On the basis of the character of the more or less glau- cous leaves, particularly the upper, entire linear ones with cilia, C. innocuus can be united with the European species C. glaucus Baumg. 24. C. seminudus MB. apud Willd. Sp. Pl. III, 3 (1803) 1657; ej. Fl. taur.-cauc. II, 271, III, 554; DC. Prodr. VI, 623; Ldb. FI. Ross. II, 720; Boiss. Fl. or. III, 520; Grossh. Fl. Kavk. IV, 176; Kapeller in FI. Gruzii, VIII, 471; Isaev in Fl. Azerb. VIII, 385.—Ic.: MB. Cent. II, tab. LXXV. Biennial. Plant to 50-60 cm high. Stem slender, simple and slightly ribbed below, finely sulcate, pubescent, slightly winged, wing lobes small with thin spines, branched above; branches straight, long. Leaves lanceolate, crenate-toothed, spinose-small-toothed, decurrent; lower leaves broad, their lamina thin, subglabrous above, tomentose-pubes- cent beneath; peduncles very long, more than 1/2 as long as entire stem, not winged, slender, cylindrical, finely sulcate, densely white- tomentose when young, later becoming irregularly spinose-pubescent. Capitula solitary, to 3 cm in dia and 2.0-2.5 cm long. Involucre subglabrous; involucral bracts very sparsely arachnoid-pubescent along margin, often purple; outermost bracts shorter, middle longer, subu- late, more or less spinescent; inner-most linear, acute, finely ciliate, purple outside, pearl-pink inside, with distinct midrib and pseudo- glandular hairs in upper part. Corolla pink or pale-purple, irregularly incised. Achenes grayish-yellow, with thin longitudinal brown stria- tions, and distinctly 5-lobed tubercle at apex; pappus shorter than corolla and equal to or slightly longer than involucral bracts. Flowering July to August. Dry places, among xerophytic shrubs.—European Part: Crimea, Lower Don; Caucasus: all regions; Soviet Central Asia: Karakum (rarely). General distribution: Balkans-Asia Minor, Iran. Described from Kizlyar. Type in Leningrad. Note. Like most of the species of this genus, it is a fairly polymor- phic species, widespread throughout the Caucasus. A particularly 32 31 interesting form is found in Pirchevan in the Zangelansk Region, with broad thin leaves (var. foliosus). 25. C. kerneri Simk. in Term. fuz. x (1886) 181; Klok. in Vizn. Rosl. URSR, 565.—C. nigrescens Baumg. Enum. stirp. Transsilv. II (1816) 56; Fuss, Fl. Transsilv. excurs. 362, non Willd.—C. hamulosus Czetz. in Erd. Mus. VI (1872) 12, non Ehrh.—C. transilvanicus Borb. in Oest. bot. Zeitschr. XXXV (1886) 274.—C. defloratus var. kerneri Gugler in Mitteil. Bayer. Bot. Ges. II (1908) 139; Hegi, Ill. Fl. VI, 2, 854. Biennial or perennial. Plant to 50-60 cm high stem ascending, straight, cylindrical, finely sulcate, almost always simple, leafy in lower 2/3, slightly pubescent, narrowly winged; wings spiny, merging with decurrent leaf. Basal leaves long-petiolate, 6-10 cm long, 3—4 cm wide, ovate or oblong; upper leaves reduced, to 2—3 cm long, narrowly lanceolate, often with long, almost linear terminal lobe; lamina pin- nately lobed or pinnately dissected into lanceolate lobes; lobes entire in lower part of leaf and unevenly crenate in upper part with short and thin spines, sometimes slightly incurved, glabrous, finely arachnoid- pubescent beneath, particularly on midrib; peduncles long, not winged, pinkish-white-tomentose, densely lanate above and less so beneath. Capitula medium-sized, 2.5-3.0 cm long, subglobose. Involucre con- siderably shorter than florets; involucral bracts glabrous, sometimes pubescent only on margin on inner side; outer bracts lanceolate, with short cusp and distinct midrib; middle and innermost bracts linear, acute, pinkish, lacking spiny cusp, with dark midrib. Corolla pink, to 2 cm long; limb irregularly incised. Achenes to 0.5 cm long, light- brown, with distinct, darker, transverse striations, and small apical tubercle; pappus hairs lacking distinct barbs, considerably shorter than corolla. Flowering July to August. Meadows and pastures.—European Part: Upper Dniester (Eastern Carpathians). General distribution: southern and Central Europe. De- scribed from Carpathians. Type in Budapest. Economic Importance. A very elegant plant that can be used as an ornamental. Section 3. Stenocephalus Rouy, FI. Fr. IX (1905) 70.—Capitula 1.2-1.7 cm, less often 2.0—2.5 cm long; involucre oblong, cylindrical, narrowly campanulate or narrowly obconical. Annuals, usually with readily falling capitula and small, slightly and almost uniformly in- cised lobes of narrow corolla. Type of section: C. pycnocephalus L. 33 a2 Series 1. Pycnocephali Tamamsch.—Involucral bracts pubescent, appressed or recurved, never lustrous, usually lacking scarious margin, sometimes with thicker tip, abruptly narrowed to short spine. 26. C. pyenocephalus L. Sp. pl. ed. 2 (1763) 1151; Geideman, Opred. Rast. Mold. SSR, 286.—C. pycnocephalus eupycnocephalus Jacq. Hort. Vindob. I (1770) 17.—C. tenuiflorus y. pycnocephalus DC. Prodr. VI (1837) 626.—Iec.: Jacq. loc. cit. tab. 44. Annual. All plants light-green; stem 90-100 cm high, usually straight, branched, less often simple, white-tomentose, narrowly winged, discontinuously spiny-winged. Leaves pinnately lobed or sinuate-pin- nate, with palmate or deltoid lobes and small divergent secondary lobes, terminating in subulate spine, spinose-ciliate with spines to 1 cm long, slightly arachnoid-pubescent above, more densely pubescent beneath; upper leaves decurrent. Capitula readily falling, on white-tomentose, narrowly winged peduncles in clusters of 2—5, almost cylindrical, 2.0— 2.5 cm long, 1—2 cm in dia. Outer involucral bracts lanceolate, gradu- ally and indistinctly extended into triangular, recurved spine, dorsally yellowish because of small pseudo-glandular hairs, ventrally greenish, with not so prominent midrib; middle and inner-most bracts narrowed to short spine, 3—5 veined, light-colored on margin, narrowly scarious, short-ciliate all over surface and margin. Florets purple; corolla 16.0- 16.5 mm long, limb almost uniformly incised, 5—6 mm long, exceeding inner involucral bracts and pappus; stamens including appendages 5— 6 mm long. Achenes 5—6 mm long, 1.5—2.0 mm wide, yellowish-gray, with indistinct, darker, longitudinal striations, lustrous, with round or slightly angular, sessile tubercle; pappus white, its bristles serrate, 2— 3 times as long as achene. Flowering May to July. Sunny places, on uncultivated and fallow fields —European Part: Upper Dniester (rarely), Bessarabia. General distribution: southern Europe. Described from southern Europe. Type in London. 27. C. beckerianus Tamamsch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XV (1953) 383.—C. cinereus var. amblyolepis Trautv. in Tr. Peterb. Bot. Sada, I, 1 (1871) 19.—C. knorringianus Tamamsch. loc. cit. 384 and in Fl. Turkm. VII (1960) 238. Annual. Plant to 70-80 cm high. Stem ribbed, slightly angular, on ribs discontinuously spiny-winged; on edges slightly sulcate, glabrous below or sparsely arachnoid-hairy or pubescent to tip, sometimes not winged at apex but with solitary long spines. Lower leaves decurrent, subsessile or on winged petioles, deeply and pinnately to sinuately lobed, oblong; lobes remote, deltoid, 3-toothed, with thin sharp spines; lamina thin, somewhat sparsely puberulent or subglabrous above, finely 34 33 grayish arachnoid-pubescent beneath; peduncles white or grayish- tomentose to capitula but always with small leaves below capitulum, sometimes modified into acute lanceolate cusp. Capitula small, 2.5 cm long, 1.0—1.5 cm in dia, ovate till flowering, more or less cylindrical at dehiscence, in clusters of 3(2—4), readily falling. Involucre lacking long gray hairs, appearing glabrous; involucral bracts with colored tips, with somewhat raised, violet, purple, brownish or bright spot, and brighter and prominent midrib; outermost bracts ovate, abruptly nar- rowed to short spine, less often narrowly lanceolate, except in upper part covered with yellow pseudo-glandular hairs; inner bracts longer, with lighter scarious appendage at tip, slightly fimbriate along margin. Corolla pink, scarcely exceeding involucre. Achenes small, lustrous white or light-gray, 1.0—1.5 (2.0) mm long, 1 mm wide, more or less flat, with indistinct, very sparse, longitudinal striations and transverse zigzags, with small tubercle lacking distinct lobes or edges; pappus hairs barely scabrous, longer than involucral bracts and equaling or exceeding corolla. Flowering May to August. (Plate III, Fig. 2.) Dry wastelands, in ravines, old cotton fields.—Caucasus: Eastern Transcaucasia; Soviet Central Asia: mountainous Turkmenia; General distribution: northern Iran. Described from Lenkoran. Type in Leningrad. 28. C. cinereus MB. FI. taur.-cauc. II (1808) 270, III (1819) 554; DC. Prodr. VI, 624; Ldb. FI. Ross. II, 721; Grossh. Opred. Rast. Kavk. 479.—C. pycnocephalus 56. cinereus Boiss. Fl. or. II (1875) 521.—C. pycnocephalus y. cinereus Schmalh. FI. II (1897) 98; Grossh. Fl. Kavk. IV, 174. Annual. Stem sulcate, short-grayish-pubescent, more or less winged and spiny below, not winged above, branched. Leaves slightly pubes- cent above, sometimes subglabrous, finely grayish-tomentose beneath; lower leaves to 20 cm long, 6-7 cm wide, on winged petioles, pin- nately-sinuately lobed, lower lobes acute deltoid, increasing in size, with long cusp; middle and upper leaves to 10—12 cm long, oblong, often with lanceolate upper lobe, usually less lobed, thin. Peduncles long, almost always not winged, densely white-tomentose, bearing 1, less often 2—3 capitula, but then middle capitulum usually larger than others. Involucre almost equaling florets, grayish-pubescent; capitula cylindrical, often on much longer secondary peduncles, involucral bracts erect, appressed in flowering and fruiting capitula, slightly divergent, but not recurved, lanceolate-linear, with indistinct midrib; inner bracts scarious, with short pseudo-glandular pubescence in lower part. Co- rolla pink, its limb uniformly incised. Achenes obovate, almost white or light gray, with or without fine, indistinct, longitudinal striations, 34 \ ! ny) 7 WY ng. Si "r ara Ne Ca SY We re) fae Plate III. 1—Carduus cinereus MB.; 2—C. beckerianus Tamamsch.; Tamamsch. 3—C._ nikitinii 36 35 very small, 3-4 mm long; pappus equaling or longer than inner involu- cral bracts and equaling corolla. Flowering June to July. (Plate III, Fig.19) Fallows fields and uncultivated places.—European Part: Lower Volga, Crimea; Caucasus: Ciscaucasia, Eastern and Southern Transcaucasia; Soviet Central Asia: mountainous Turkmenia. General distribution: Iran. Described from Kizlyar. Type in Leningrad. 29. C. arabicus Jacq. apud. L. Syst. ed. 14 (1784) 724; Collect. I, 56; Willd. Sp. pl. III, 3, 1647; DC. Prodr. VI, 626; Grossh. Opred. Rast. Kavk. 479.—C. arabicus var. albidus Grossh. Fl. Kavk. IV (1934) 174.—C. albidus MB. FI. taur.-cauc. II (1808) 269, III (1819) 554 p. p.; C.A.M. Verzeichn. 71; Grossh. Opred. Rast. Kavk. 479; Kapeller in Fl. Gruzii, VII, 473.—C. pycnocephalus C. Koch in Linnaea, XVII (1843) 42, non L.—C. pycnocephalus y. arabicus Boiss. FI. or. III (1875) 521.—C. pycnocephalus B. albidus Boiss. 1. c.—C. tenuiflorus 8. albidus DC. Prodr. VI (1837) 626.—C. cinereus Arénes in Not. Syst. Paris, XV, 4 (1959) 396, non MB. Annual. Stem slender, ribbed and slightly sulcate, grayish- or whitish-pubescent, almost not winged below, continuously broadly winged in upper half; wings with stiff spines along margin, 3-10 mm long. Leaves decurrent, 15—20 cm long, somewhat reduced upward but present up to apex; lamina thin, pubescent above with long, gray hairs or denser white hairs; finely grayish-arachnoid-hairy or densely white- pubescent beneath, unevenly pinnately lobed; lobes distinct, broadly deltoid, reduced toward base, irregularly sinuate-toothed, 5—7 toothed, with prominent midrib extended into spine, 2—3 cm long in subtending leaves; peduncles densely tomentose, winged or densely spinose, ax- illary with 2-5 small capitula clustered at base, sometimes readily falling; immature capitula borne racemosely on short secondary pe- duncles. Involucre narrowly conical-campanulate; involucral bracts in unopened capitula erect or slightly recurved, imbricate at flowering, somewhat sparsely or densely white-pubescent; outer bracts ovate or more or less oblong-ovate, mucronate, with short, straight or slightly recurved spine and prominent midrib; middle bracts obtuse, terminat- ing in spine; inner bracts oblong, narrowed into cuneate tip, with or without weak spine, white-scarious along margin, with 3—5 thin but distinct veins, darker in upper half than lower, covered, like outer bracts, with pseudo-glandular hairs. Corolla pink or bright purple, 11-— 12 mm long; corolla tube 6.5—7.0 mm long including limb, lobes 4—5 mm long. Achenes creamish or light-brown, lustrous, with thin longi- tudinal striations, S—6 mm long, 1.0—1.5 mm wide, with small decidu- ous tubercle. Flowering May to July. 37 36 Weedy places, long-fallow fields, dry habitats —European Part: Crimea, Trans-Volga, Lower Don, Lower Volga (rarely); Caucasus: Ciscaucasia, Dagestan, Eastern and Southern Transcaucasia; Soviet Cen- tral Asia: mountainous Turkmenia (rarely). General distribution: At- lantic Europe (eastern part), Armenia and Kurdistan, Iran. Described from Arabia. Type in Paris. Note. J. Arénes (1959) considers that the species C. albidus and C. cinereus, described by Bieberstein, should be regarded as forms of C. pycnocephalus belonging to ssp. pycnocephalus. Furthermore, he thinks it is possible to refer the first species (C. albidus) to var. pycnocephalus and the second to var. arabicus. Having at my disposal the type specimens of Bieberstein, I cannot agree with such an inter- pretation. C. albidus MB. and C. cinereus MB. are well distinguished from each other not only by the pubescence of the involucral bracts and the structure of the peduncles, but also by the relative proportions of the corolla parts, to which great taxonomic importance is accorded by Arénes himself, and also by the achenes, to which he did not pay attention. In C. cinereus the achenes are little more than 1/2 as long as in C. albidus, not only in the type specimens of Bieberstein, but also in the numerous specimens examined by me in the Caucasian Herbarium. Only in one specimen of C. cinereus, collected in the Nakhichevan ASSR, did I find achenes almost as long as in C. albidus. Without entirely supporting Arénes and his broad interpretation of C. pycnocephalus, I agree with him in one respect that the separate spe- cies status of C. albidus is not justified. Therefore, I combine this species with C. arabicus Jacq., but do not refer it to C. pycnocephalus ssp. pycnocephalus var. pycnocephalus, as Arénes did. So far as C. cinereus is concerned, it is hardly possible to include it in var. arabicus, because it differs in many characters from both C. pycnocephalus s. str. and C. arabicus sensu Jacquin. Rather, it could be identified with the Mediterranean species C. argentatus L., which is widely distrib- uted in the eastern Mediterranean region. The forms found in Nakhichevan, although close to this species, however, give the impres- sion of hybrid forms that have arisen, possibly, as a result of the crossing of C. cinereus with C. nervosus. Series 2. Tenuiflori Tamamsch.—Involucral bracts subglabrous or slightly pubescent, often green after flowering, lustrous, with narrow, more or less scarious margin. This series includes 4—5 species from Western Europe and two species in the USSR—in the Caucasus and Soviet Central Asia. 38 37 30. C. nikitinii Tamamsch. in Addenda XXVII, 600.—C. cinereus var. pusilla Tamamsch. in Fl. Turkm. VII (1960) 238, diagn. ross. Annual. Plant 5—15 cm high; stem slightly angular, covered with small leaves and spines to capitula, grayish-pubescent below, simple or occasionally with few branches. Leaves 2—4 cm long, on decurrent, winged, prickly. petioles, grayish-pubescent on both sides with long flexuous hairs and prominent, riblike, whitish midrib beneath, irregu- larly and weakly crenate, with shorter as well as longer spines; pe- duncles cylindrical, white-tomentose, with 1 or 2 sessile capitula. Capitula few-flowered (8—9 flowers), truncate, turbinate. Involucre 4— 5-seriate, distinctly imbricate; outer involucral bracts broadly ovate, with distinct midrib in upper half, narrowed into short spine; middle bracts oblong-ovate, inner longer, linear-lanceolate, darker at tip, with 3 distinct veins; all bracts greenish, glabrous, lacking long hairs in lower part, covered with short pseudo-glandular hairs, narrowly mem- branous along margin. Corolla 10-11 mm long, pale purple, slightly zygomorphic; corolla tube including limb 7-8 mm long. Achenes 5—6 mm long, 1.0-1.5 mm wide, obovate, slightly compressed, somewhat oblique in lower part, dull-white, with fine, darker, longitudinal stria- tions in upper part and large, up to 1.0—1.5 mm-long, cylindrical, long persistent, indistinctly 5-angled, brownish tubercle; pappus dull white, 2—2.5 times longer than achene, exserted from involucral bracts and longer than corolla. Flowering April to June. (Plate III, Fig. 3.) Dry talus slopes.—Soviet Central Asia: mountainous Turkmenia, Kara-Kum. Endemic. Described from Kopetdag. Type in Leningrad. 31. C. nervosus C. Koch in Linnaea, XVII (1843) 42, XXIV (1851) 391; Ldb. Fl. Ross. II, 722; Boiss. Fl. or. III, 521; Grossh. Fl. Kavk. IV, 175; Opred. Rast. Kavk. 479, p. p. Annual. Plants 30-40 cm high; stem ribbed, irregularly winged, narrowly winged above, wings broader and prickly below. Leaves lin- ear or oblong, with narrowly winged petiole, pinnately lobed, spiny along margin, lobes deltoid, 3—5-toothed, spiny, light green, with promi- nent midrib beneath. Peduncles long, bearing 3—6-capitula at tip, less often capitulum1!. Involucre more or less broad, cylindrical, in early stages bracts contiguous with short erect cusp; involucral bracts imbri- cate, subglabrous, ovate-lanceolate; outer bracts shorter, mucronate; inner acute, colored; all bracts covered all over, except midrib, with fine, yellowish pseudo-glandular hairs; innermost bracts very long, equaling pinkish corolla. Achenes 6 mm long, gray, oblong, slightly compressed and oblique, with quite distinct 5-angled tubercle; pappus of more or less silky grayish or dull-white, finely scabrous hairs, as long as inner involucral bracts and corolla. Flowering June to July. 39 38 Semidesert arid areas.—Caucasus: Southern Transcaucasia. General distribution: Iran. Described from Echmiadzin. Type was in Berlin. Note. This species has been interpreted quite differently by Cau- casian botanists. There are specimens in the Caucasian herbaria with a completely different habit and characters, which are identified by D.I. Sosnowsky and A.A. Grossheim as C. nervosus C. Koch. Initially, Koch himself (1843) reported that this species occupied an intermedi- ate position between C. onopordioides Fisch. ex MB. and C. tenuiflorus DC., but is much closer to the first species. In the addendum to “K Izucheniyu Flory Vostoka” [The Study of Flora of the East] (1851) he noted that the capitula of C. nervosus are considerably larger than those of C. onopordioides and compared them with the capitula of C. confertus and C. fasciculiformis from the affinity of C. pycnocephalus L. Boissier related C. nervosus to the annual bristle-thistles of the C. pycnocephalus type. In the Caucasian Herbarium of the Botanical In- stitute of the Academy of Sciences of the USSR, there is a specimen with a label on which in Meyer’s handwriting is written “C. onopordioides, Echmiadzin legit et determinavit Koch 1837,” and at the bottom, “C. nervosus.” I took this specimen as the basis for clari- fying the species C. nervosus. It must be stated that in this specimen, the capitula are really similar in size and form (more or less cylindri- cal) to those of the European species C. tenuiflorus, but they are not larger than those of C. onopordioides Fisch. ex MB. The presence of a large number of intermediate forms between the species, particularly in the last section, testifies to the extensive hy- bridization occurring in the genus Carduus. Below I cite a number of the intermediate forms found in the USSR, but it undoubtedly is pos- sible to find hybrids between other species too. The genera Galactites and Cirsium, close to the genus Carduus, apparently also have species that cross with bristle-thistle species. Attempts to erect the genus Carduicirsium are well-known in the literature. Sennen described three species that were later transferred to the genus Carduus by Fournier (1940). Arénes, in 1942, described Cirsiicarduus lutetianus—a hybrid between Cirsium vulgare (Savi) Ten. ssp. savianum and Carduus tenuiflorus. The hybrid between Carduus and Galactites, described as Carduiga-lactites ludoviciana, was transferred to the genus Galactites by Fournier. Natural hybrids found in our flora are as follows: 1. C. acanthoides L. x C. crispus L. 2. C. acanthoides L. x C. nutans L. 3. C. acanthoides L. x C. personata (L.) Jacq. 4. C. cinereus MB. x C. nervosus C. Koch. 40 39 . crispus L. x C. nutans L. . crispus L. X C. personata (L.) Jacq. . hamulosus Ehrh. x C. nutans L. _ hamulosus Ehrh. x C. onopordioides Fisch. ex MB. . nervosus C. Koch x C. onopordiodes Fisch. MB. 10. C. nutans L. Xx C. onopordioides Fisch. ex MB. 11. C. nutans L. x C. personata (L.) Jacq. Om NAY ea ao GENUS 1590. Alfredia Cass.'? Cass. in Bull. Soc. Philom. Paris (1815) 175. Capitula homogamous, large, many-flowered. Involucral bracts with more or less coriaceous or membranous, marginal, incised append- ages, spinescent in outer bracts, spatulate in inner ones, or outer bracts spinescent and inner with toothed, acute-tipped appendage. Receptacle densely covered with bristles. All florets tubular, actinomorphic, bi- sexual, with short tube and short almost equally 5-parted limb; corolla long, narrowly tubular with short tapering lobes distinctly incurved. Basal appendages of anthers setose, truncate, in lower part, short; fila- ments free, glabrous or slightly tuberculate. Style exserted, unbranched almost to tip, covered with short papillae. Pappus many-rowed; pappus bristles connate at base in ring, unequal, fragile, mostly pale yellow, barbed, inner bristle longer, slightly flattened apically; achenes obconical, 6 mm long, compressed, straight, glabrous, dull, with basal or slightly oblique hilum. Perennial herbs with long-petiolate lower leaves. Type of genus: A. crenua (L.) Cass. 1. All involucral bracts broad, membranous, golden, short, spinescent, toothed; leaves undivided, ovate-cordate; cauline leaves with winged petiole, basally expanded, amplexicaul......... 1. A. cernua (L.) Cass. + Involucral bracts coriaceous; outer bracts entire or spiny-ciliate, acuminate-spinescent; middle bracts discontinuously ciliate on marnean with ScamousjappenGapes.f4. .. ss: sca. aebewenapere ...i. ha iye...28. Ce k.. Lacs kek. fbb Seales chee, wekees aReeee eee a: 106. Leaves entire, thinly covered with small, basally torulose hairs above, arachnoid-hairy beneath, densely spinose-ciliate. Capitula solitary on long leafless peduncle............... 77. C. pannonicum (L. fil.) Link. Leaves entire, sometimes lower leaves pinnately divided, with few segments, more or less thinly arachnoid-hairy, spinose-ciliate; up- per leaves usually reduced. Capitula solitary or in clusters of 2-3 on long, few-leaved peduncles..................... 75. C. canum (L.) All. Florets red. Capitula solitary on short peduncles, usually in lax panicles. Cauline leaves broadly oblong-ovate or ovate, slightly lobate-toothed; middle and upper cauline leaves broadly amplexicaul, lower scarcely decurrent............... 87. C. depilatum Boiss. and Bal. Florets purple-red. Capitula solitary or in clusters of 2-3 in corym- bose-paniculate inflorescences. Leaves oblong, elliptical, pinnately lobed; middle and upper cauline leaves sessile, semiamplexicaul, lower somewhat decurrent................. 88. C. hygrophiloides Charadze Leaves on both sides glabrous, very rarely floccose-arachnoid, usu- ally entire, more or less elongate, less often remotely pinnately toothed, sessile, with small, appressed spines along margin. Stems not winged, less often lobately winged below................cccccccsssesseeeee BEE SES ache ak ach « PESRGIED. «0d, HOR ode na 108. C. setosum (Willd.) MB. 68 + Leaves more or less pinnately parted, sessile, sometimes with more or less decurrent prickles. Stem not winged, less often with narrow- winged, long-spinose internodes beloW.............:.csssssessssesesesseeeeesees 109. 109. Stem and leaves, particularly beneath, densely, grayishly and al- most tomentosely hairy; leaves shallowly sinuate-lobed, sometimes almost entire............... 109. C. incanum (S.G. Gmel.) Fisch. ex MB. + Leaves on both sides glabrous or thinly arachnoid beneath........ 110. 110. Leaves more or less small and stiff, with elongate lobes and teeth more or less strongly spinescent; lower cauline leaves sometimes decurrent on internodes, with long, thin spines. Involucral bracts purple-vidlets..tls...sccae.0n ee 111. C. arvense (L.) Scop. + Leaves stiff, coriaceous, with both sides glabrous or thinly grayish- arachnoid, shallowly sinuate-lobed, with teeth abruptly narrowed into small, hard spines. Involucral bracts usually stramineou.............. LOMA Rs Desh ae ie a teh ol yeaa 110. C. ochrolepideum Juz. Section 1. Caucasigena Charadze in Addenda XXVII, 600.—Sect. Epitrachys auct. non DC. in Duby, Bot. Gall. I (1828) 286; Boiss. FI. or. III (1875) 523 p. p.; C. Koch in Linnaea, XXIV, 396 p. p., pro gen.—Eriolepis Cass. in Dict. Sc. nat. XXV (1825) 172 p. min. p., pro gen.; DC. Prodr. VI (1837) 635 p. min. p., pro sect—Lophiolepis Cass. in Dict. Sc. nat. XXVII (1823) 180 p. min. p., pro subgen., XXXV (1825) 71, pro gen.; DC. Prodr. VI (1837) 634 p. p., pro sect.—Capitula drooping. Involucre more or less arachnoid-hairy; in- volucral bracts entire or spinose-ciliate along margin, oblong-ovate or lanceolate in lower part, linear-lanceolate above, strongly narrowed into long, hard spine. Florets red, less often whitish; corolla tube nar- row usually more or less surpassing limb. Achenes 5—6 mm long. Leaves large with broad lamina, more or less deeply sinuately-pinnately di- vided or lobed, densely covered above with spiny bristles. Mostly perennials with ascending, apically branched stems and more or less numerous capitula clustered in paniculate, less often capitate common inflorescence. Type of section: C. caucasicum (Adams) Petrak. Note. The species with drooping but not erect capitula and long- spinose involucral bracts are somewhat isolated, exhibiting definite genetic affinity with the ancient Colchid subsection Aggregata Petrak, which we include [as a series] in section Caucasigena. The range of the section if confined mainly to the Caucasus; only a few of the species grow in the adjoining areas of Asia Minor. The majority of the species belong to the xerophilous representa- tives of forest vegetation, which grow in open forests, in tall herb vegetation, and along forest edges from the mid-montane to subalpine 70 69 zone. Of these species, C. sommieri Petrak from Asia Minor is particu- larly distinguished by its more xerophilous habit. Series 1. Aggregata (Petrak) Charadze comb. nova.—Agegregata Petrak in Bibl. Bot. 78 (1912) 6, pro subsect.—Leaves sessile, pin- nately lobed or almost undivided. Capitula small, in capitate inflores- cence on apices of stem and branches, oblong. Involucral bracts more or less appressed, outer ones attenuated into long hard spines, small- toothed. 1. C. aggregatum Ldb. Fl. Ross. II (1846) 737; Boiss. Fl. or. III, 537; Grossh. Fl. Kavk. IV, 180; Kolak. Fl. Abkhazii, IV, 272; Charadze in Fl. Gruzii, VIII, 490; Dmitr. Opred. Rast. Adzh. 275.—C. aggregatum var. macrophyllum Alb. Prodr. Fl. Colch. (1895) 143.— Ic.: Dmitr. op. cit. Fig. 31. Perennial. Stem up to 1 m high, robust, ascending, dark green or somewhat purple, slightly angular, sulcate, scatteredly crisped hairy, leafy up to tip, with terminal capitate inflorescence of small capitula, weakly branched, with 1-2 slender axillary shoots bearing few unde- veloped capitula at apex. Cauline leaves green, densely covered above with long, thin, spiny bristles, densely grayish-arachnoid-hairy beneath; lower leaves oblong-ovate, deeply-pinnately divided into broadly ovate, sinuately biparted lobes, narrowed toward base to semiamplexicaul petiole, acute, spinose-ciliolate on margin, acuminate, with teeth ter- minating in yellowish spines; middle and upper cauline leaves usually like lower, 10—15 cm long, 5—7 cm wide, ovate, pinnately divided into broadly ovate lobes or almost undivided, sessile, amplexicaul. Lower leaves large, in var. macrophyllum Alb., with few lateral segments and 3-lobed terminal segment; middle and upper cauline leaves almost undivided or pinnately toothed; broadly lobate apical leaves at base of inflorescence surpassing it and usually recurved. Capitula terminal, 6— 10 in capitate inflorescence, less often solitary or 2-3 at tip of short, axillary branches, globose-ovate, 12-17 mm in dia, 15—20 mm long. Involucre sparsely arachnoid-hairy, greenish or purple; outer involu- cral bracts glabrous or with scattered arachnoid hairs, carinate, small, lanceolate, finely serrate, gradually attenuated into elongate, green, linearly-subulate point with long spine; inner bracts oblong-lanceolate, with shorter spines; florets red, less often white, 13-15 mm long, with long, narrow tube, limb to 5 mm long, 5-parted almost to middle. Pappus slightly shorter than florets, sordid white, with hairs almost plumose up to tip; inner hairs slightly enlarged above; achenes brown- ish-reddish-brown, 5—6 mm long. Flowering July to August. 7 — 70 Upper forest and lower alpine zones, in meadows and tall herb vegetation.—Caucasus: Western Transcaucasia (Gornaya Racha, Adzharia, Guria). General distribution: Asia Minor (Lazistan). De- scribed from Guria. Type in Leningrad. Note. It stands somewhat isolated in the section, apparently being an ancient colchid species. It has not yet been well studied. The variety C. aggregatum var. macrophyllum Alb., described from Abkhazia, is distinguished by the form of the leaves, purple-colored involucral bracts, and purplish florets. Similar specimens are also known from Adzharia. Series 2. Colchica Charadze.—Capitula oblong, up to 20 mm in dia, in clusters of 2—3 or solitary, forming corymbose-paniculate inflo- rescence. Involucral bracts sharply serrate, scabrous or ciliate from thin spiny bristles. 2. C. gagnidzei Charadze in Addenda XXVII, 600. Perennial. Stem erect, up to 1 m high, robust, ribbed, slightly sulcate, subglabrous, simple, with short branches near tip, purplish, leafy to apex. Leaves sessile, oblong, green above, with small dense bristles; bluish, more or less arachnoid-hairy beneath, with prominent subglabrous veins; lower leaves narrowed toward base into lobate long- spinose petiole, almost to base pinnately dissected into 2-lobed ovate- deltoid segments, on margin with thin appressed spines; lobes and teeth of segments terminating in hard, yellowish 5-7 mm-long spines; middle cauline leaves auriculate, semiamplexicaul, deflected, to 15 cm long, 7 mm wide, deeply sinuately pinnately divided into ovate-del- toid, 2-fid, toothed strongly spinescent lobes; upper leaves reduced, sinuately-pinnately lobed; apical leaf 1, at base of capitula small, re- curved, lanceolate, pinnately toothed, somewhat surpassing capitulum. Capitula subsessile, in clusters of 2—3 or solitary terminal on stem and short branches, oblong at beginning of flowering, later ovate-globose, forming corymbose common inflorescence. Involucre 10—20 mm in dia, at base weakly, in upper part more or less densely arachnoid- hairy; involucral bracts greenish, lanceolate, carinate, attenuated from middle into upward directed or spreading, yellowish, thin spines; lower bracts small, subglabrous; middle elongate, in upper part arachnoid- hairy; inner bracts linearly lanceolate, with elongated, filiform, curved acute point. Florets purple; corolla 20 mm long, limb parted almost up to half, equaling narrow tube. Pappus sordid white, with long, sca- brous tip of inner hairs; mature achene unknown. Flowering August. Subalpine tall herb vegetation at 1,750-1,900 m.—Caucasus: Western Transcaucasia. Described from basin of Mzymta River (Psekakho Ridge). Endemic. Type in Tbilisi. 7 nN 71 Note. It occupies a distinct position in the section. On the basis of its small capitula in clusters of 2-3 and upward-directed spines, it can only be close to C. aggregatum Ldb. 3. C. longiflorum Charadze in Addenda XXVII, 601. Perennial. Stem to 150 cm high, more or less strongly branched from middle, leafy up to tip, weakly sulcate, purplish, slightly bluish, scatteredly arachnoid-hairy in upper part. Leaves elliptical or oblong- elliptical, gradually reduced toward stem apex, 4-17 cm long, 1.5—7.0 cm wide, broadly, deeply pinnately divided into more or less oblong- lanceolate or deltoid lanceolate 2—3(5)-parted segments; lobes of seg- ments terminating in sharp, long, stiff, yellowish-white, 5-10 mm-long spines; all leaves dull-green above from somewhat scattered, yellow- ish-white bristles and very small scattered hairs, grayish-tomentose beneath, with prominent, thinly pubescent whitish veins; apical leaves near base of capitula, usually small, lanceolate, usually 2-3, shorter than involucre or almost as long. Capitula numerous, divergent or horizontal, on short peduncles, usually 1.5—2.0 cm in dia, with florets 3.0-3.5 cm long, 2-7, less often solitary at stem tip and on lateral leafy, usually more or less divergent branches surpassing stem apex, forming lax corymbose-panicles. Involucre subglabrous or with occa- sional scattered, arachnoid-hairs, usually more or less dark purple; outer bracts lanceolate, carinately convex, dorsally keeled in upper part, densely imbricate, narrowed from middle to long, upward di- rected, spreading, stiff, yellowish spines, ciliolate on margin, scabrous; inner bracts linear-lanceolate, scarious, acuminate, with thin, dark colored, curved filiform point, scabrous along margin, very finely, sharply serrate. Florets purple. Corolla 20-22 mm long, limb deeply 5- parted; corolla tube narrow, almost as long as or somewhat shorter than limb. Pappus 16 mm long, sordid white; inner hairs of pappus plumose, with somewhat thickened, scabrous tip; achenes 4.5 mm long, 2.3 mm wide. Flowering August to September. Mid-montane zone, post-forest glades.—Caucasus: Western Transcaucasia (ravine of Mzymta River). Endemic. Described from Krasnaya Polyana. Type in Tbilisi. Note. In the structure of its involucre this species is closer to C. gagnidzei Charadze and, possibly, is its xerophilized derivative. From the latter it is distinguished by leaves deeply incised into narrow seg- ments and numerous elongated capitula. 4. C. ezerkessicum Charadze in Addenda XXVII, 602. Perennial. Stem up to 200 cm high, ascending, from middle branched above, blackish-purple, slightly sulcate, leafy to apex, with 1p Mi AS we fe 73 Plate IV 1—Cirsium caucasicum (Adams) Petrak; 2—C. tomentosum C.A.M. 74 73 upward directed, somewhat divergent branches, subglabrous, with sparse, scattered, crispate appressed hairs. Cauline leaves sessile, nar- rowed toward apex; lower leaves with long, semiamplexicaul auricles, somewhat densely covered above with appressed spiny bristles, and crispate hairs along veins, pale green beneath and with scattered floc- cose hairs or thinly grayish-tomentose, 25 cm long and 10 cm wide, deeply sinuately-pinnately lobed with distinct deltoid-ovate, 2—3(5)- lobed segments, thinly spinose-ciliate along margin, lobes terminating in sharp, thin, 4-8 mm long spines, with up to 12 mm-long yellowish spines at base of leaves, oblong elliptical; other leaves, like lower, gradually reduced toward stem apex; middle leaves 17 cm long and 8 cm wide, upper oblong-lanceolate; apical leaves 4-5 cm long, sub- tending and somewhat surpassing capitula. Capitula few, oblong, 15— 22 mm in dia, with florets up to 3 cm long, nodding; terminal capitula solitary or 2 each on short, erect peduncles, axillary capitula undevel- oped or strongly reduced; capitula crowded at stem apex and divergent branches in short corymbose-paniculate inflorescence. Involucre gla- brous in lower part, sparsely arachnoid-hairy in upper, with somewhat recurved, small, yellowish spines; outer bracts small, oblong, convex, slightly carinate, yellowish-green, less often purple, terminating in sharp, falcately recurved, short spine, more or less densely spinose-ciliate on margin from upward directed, yellowish spines; inner bracts scarious, linear-lanceolate, terminating in sharp, thin, almost filiform, recurved cusp, on margin finely, sharply serrate-dentate, scabrous in upper part, slightly arachnoid-hairy or glabrous; florets lilac-red. Corolla to 22 mm long, with narrow tubes to 10 mm, limb parted up to 1/3. Pappus sordid-white with inner hairs plumose almost up to apex, with slightly enlarged scabrous tips. Flowering August to September. Subalpine zone, in tall herb vegetation and in secondary semi-tall herb meadows.—Caucasus: Ciscaucasia (basin of Kisha and Urushten rivers), Western Transcaucasia (in upper reaches of Mzymta-Achishkho River). Described from basin of Kisha River. Endemic, Type in Tbilisi. Note. It is distinguished from C. gagnidzei Charadze and C. longiflorum Charadze by ciliate outer involucral bracts with tiny spines on the margin and by shorter lower bracts tapered to short spines. Specimens from the northern as well as the southern slopes of the Greater Caucasus that deviate from the type are distinguished by a purple-colored involucre and weakly developed small spines on the margin of involucral bracts. Series 3. Caucasica (Petrak) Charadze comb. nova.—Sect. Epitrachys DC. subsect. Eriophora Petrak 4. Caucasica Petrak in Bibl. Bot. 78 (1912) 6.—Capitula large- or medium-sized, ovate or globose- 75 74 ovate, nodding. Involucre more or less densely arachnoid-hairy to tomentose. Involucral bracts attenuated into long, erect or recurved spines. 5. C. imereticum Boiss. Fl. or. III (1875) 537; Somm. and Lev. in Nuov. Giorn. Bot. Ital. N. S. II, 1, 14; Petrak in Tr. Tifl. Bot. Sada, XII, 1, 33; Grossh. Fl. Kavk. IV, 184; Charadze in Fl. Gruzii, VIII, 507; Dmitr. Opred. Rast. Adzh. 277; Kolak. in Rast. Mir. Kolkh. 204. Perennial. Stems erect, slightly sulcate, purple or greenish, gla- brous, finely arachnoid-hairy above, ascending, shortly-corymbosely branched near apex. Leaves oblong, deeply sinuately-pinnately divided into deltoid, lanceolate, irregularly 2—3-parted toothed segments; all lobes thinly spinose-ciliate on margin, tapered to 5-12 mm-long, hard spine; leaves densely covered above with small appressed bristles, grayish, loosely tomentose beneath; middle cauline leaves to 15 cm long, 9-0 cm wide, sessile, semiamplexicaul, upper ones reduced; subtending apical leaves small, remote from capitulum, linear- lanceolate, recurved. Capitula globose-ovate, nodding, terminal on stem and axillary branches, solitary or few, usually 2—5, subsessile. Involucre 2.5—3.5 cm in dia with long, reddish-purple spines, weakly arachnoid- hairy; outer and middle involucral bracts entire, lanceolate, slightly cari- nate, abruptly tapered to thin, long, purple, recurved spines, slightly arach- noid-hairy; inner bracts linear-lanceolate, scarious, tapered to long, thin, deflected spines; florets red. Corolla to 28 mm long, limb to 13 mm long. Pappus sordid-white with brownish, toothed tips of inner hairs. Achenes dark brown, 5.5 mm long. Flowering August to September. Herb slopes, at upper limits of forest zone —Caucasus: Western Transcaucasia (Imeretia, Adzharia, Abkhazia). Endemic. Described from Imeretia (between Olizmereti and Zekari). Type in Leningrad. Note. It is an obscure species described from incomplete speci- mens collected by Ruprecht from Imeretia. It is distinguished from C. adjaricum Somm. and Lev., by narrow, weakly pubescent involucral bracts, terminating in long purple spines. We had no opportunity to examine the specimen from Adzharia cited by Petrak as C. imereticum Boiss. This specimen is likened by Petrak to C. caucasicum ssp. cladophorum Petrak. 6. C. kemulariae Charadze in Zem. po. Sist. i Geogr. Rast. Tbil. 23 (1963) 112.—Iec.: Charadze ibid. Perennial. Rhizome thickened, woody; -stems ascending, erect, usually 1-3, 70-80 cm high, yellowish or reddish, slightly angular, sparsely arachnoid-hairy, simple or weakly branched above, leafy to tip. Leaves densely covered above with small, somewhat appressed, 76 75 spiny bristles, subglabrous beneath or on veins slightly arachnoid-hairy, less often scatteredly floccose, deeply pinnately divided, with more or less remote ovate-lanceolate or broadly lanceolate, unequally 2-fid, toothed segments, teeth and lobes tapered into spines 3.5-5.0 mm long, thinly spinose-ciliate; basal and lower cauline leaves on long, semiamplexicaul petioles, oblong-elliptical, almost up to base pinnately parted, petioles 21-25 cm long, lamina 9-10 cm wide; middle and upper cauline leaves sessile, semiamplexicaul, oblong-lanceolate to ovate-lanceolate, pinnately divided to middle or deeper into 2-parted lanceolate or deltoid-lanceolate lobes, up to 12 cm long, 5.5 cm wide, reduced to 2 cm in length toward apex. Capitula at beginning of flow- ering somewhat oblong, 1.5—2.0(2.5) cm in dia, solitary terminal on stem and its branches, short-pedunculate, few, more or less in compact corymbose-paniculate inflorescence. Involucre yellowish-floccose- arachnoid-hairy; involucral bracts narrowly lanceolate, carinate, gradu- ally tapered into spines thin, basally arachnoid-hairy and purple; outer bracts small, 3-4 mm long, longer toward inner side; inner bracts linear-lanceolate with thin, curved spines; florets purple, 26-27 mm long. Corolla limbs to 14 mm long, parted almost to 2/3. Pappus 20— 21 mm long, at base brownish, sordid-white above, with somewhat attenuated scabrous tips of inner hairs; achene dark brown, about 5 mm long. Flowering August to September. Stony habitats in mid-montane zone, in chestnut forest clearings.— Caucasus: Western Transcaucasia (Imeretia). Endemic. Described from upper reaches of Kvirila River. Type in Tbilisi. Note. So far it is known only from one locality on the basis of M.F. Sakhokia’s collections. In the structure of involucre it shows much resemblance to the western Caucasian species C. chlorocomos Somm. and Lev., from which it differs mainly by having purple and not whitish florets, smaller capitula, and thinly spiny leaves. From C. adjaricum ssp. nutans Petrak, to which it is similar in its purple-col- ored involucre and florets, it is distinguished by smaller capitula and leaves with small prickles. 7. C. chlorocomos Somm. and Lev. in Tr. Peterb. Bot. Sada, XII, 1 (1892) 154, XVI (1900) 257; Grossh. Fl. Kavk. IV, 184; Kolak. FI. Abkhazii, IV, 271; Charadze in Fl. Gruzii, VIII, 502.—Ic.: Somm. and Lev. op. cit. (1900) tab. XXV.—Exs.: Fl. cauc. exs. No. 373. Perennial. Stem up to 1 m high, slightly sulcate in upper part, with erect branches, more or less densely arachnoid-hairy or tomentose, with numerous capitula forming lax paniculate common inflorescence. Leaves more or less densely covered with small, appressed, spiny bristles above, snow-white beneath because of dense tomentose pubes- a 76 cence; lower and middle cauline leaves large, ovate-lanceolate, to 20— 25 cm long and 10—18 cm wide; all leaves deeply pinnately parted into broad, almost rhomboid, unevenly 5-fid, subacute lobes, of these shorter lobes toward leaf apex, bearing at apex yellowish, 5—10 mm-long spine; lower leaves narrowed toward base into elongated, spiny-toothed peti- ole; middle leaves sessile, auriculately expanded at base, with long spiny-toothed, semiamplexicaul; upper leaves on long axillary shoots strongly reduced, lanceolate, pinnately toothed, with long yellowish spines; apical leaves at base of capitula 1—2, subtending capitula, lin- ear-lanceolate, not surpassing capitula. Capitula terminal on stem and branches, solitary or in clusters of 2—3, nodding, pitcher-shaped, ex- panded at base, slightly narrowed in upper part, 1.5—3.0 cm in dia. Involucre densely arachnoid-hairy, almost tomentose with yellowish hairs, usually reaching tips of falcately recurved spines, forming check- ered pattern of involucre; involucral bracts lanceolate-linear, yellow- ish, terminating in recurved spines; spines considerably shorter, in lower bracts than middle ones, 10—11 mm, inner bracts almost linear, with acute point: florets pinkish-white, 25-28 mm long, with 12-15 mm_-long limb. Pappus hairs yellowish-white, with brownish, toothed tips of inner hairs; achene brown, to 5 mm long. Flowering July to August. Mainly in subalpine zone on talus, alluvial fans, occasionally in upper zone of dark coniferous forests along edges or in lower alpine zone up to 2,600 m.—Caucasus: Western Ciscaucasia (upper Kuban), Western Transcaucasia (Abkhazia, Svanetia). Endemic. Described from two specimens from Dzhodisvik Pass (between Svanetia and Abkhazia) and from Klukhor Pass. Syntypes in Florence, isotypes in Leningrad and Tbilisi. Note. In the series Caucasica, C. chlorocomos Somm. and Lev. is distinguished by white or slightly pinkish florets. The snow-white pubescence of the leaves and yellowish-tomentose involucre impart to this plant quite an ornamental appearance. The range of this species is restricted to the upper regions of the Kuban River in the western part of the Greater Caucasus, and on its southern slope this species is known from the upper reaches of the Chkhalta, Seken and Nenskrira rivers. Specimens slightly differing from the type have been collected in Western Transcaucasia (Tsebelda). There we also find the hybrid forms annotated in the herbarium by Yu. N.Voronov as xX C. tsebeldinum Woron. (C. lanceolatum x C. chlorocomos). The hybrid specimens are close to C. vulgare (Savi) Ten. and are distinguished by short decurrent leaves and erect capitula in contrast with the drooping capitula of C. chlorocomos Somm. and Lev. Speci- 78 a mens of the latter species from Tsebelda differ from the typical ones by involucral bracts with thinner apical spines, the somewhat spiny- ciliate margin of the lower involucral bracts, and the loose tomentose pubescence of the dorsal surface of the leaves. The pollen analysis to assess the hybrid nature of these specimens conducted by I.S. Shtepa does not provide any basis to consider them as hybrids. Specimens deviating from C. chlorocomos Somm. and Lev. have been identified by Petrak as C. chlorocomos Somm. and Lev. non typicum or by the authors of the Flora of Caucasus as C. tsebeldinum Woron. 8. C. caucasicum (Adams) Petrak in Tr. Tifl. Bot. Sada, XII, 1 (1912) 38; Grossh. Fl. Kavk. IV, 184; Charadze in Fl. Gruzii, VIII, 493.—C. caucasicum ssp. cladophorum Petrak in schedis p. p.; Grossh. Fl. Kavk., IV, 184 p. p.—Carduus caucasicus Adams in Web. and Mohr. Beitr. Naturk. I (1805) 64.—Cnicus horridus MB. FI. taur.- cauc. II (1808) 278, non Carduus horridus Adams (1805).—Cirsium horridum MB. FI. taur.-cauc. III (1819) 558; Spreng. Syst. veg. 3, 373; DC. Prodr. VI, 638; Boiss. Fl. or. III, 537, non Petrak (1912).— Exs.: Pl. or. exs. No. 223; Petrak, Cirsioth. Univ. Nos. 137, 138. Perennial. Stems to 100 cm high, ascending, thick, sulcate, weakly branched above, arachnoid-hairy to whitish-tomentose. Leaves green above, densely covered with very small appressed spiny bristles, more or less densely grayish- or whitish-tomentose beneath; almost all leaves pinnately divided usually to middle or deeper, with distant segments and elongate, lanceolate terminal segment, lateral segments oblong- ovate, sparsely spinulose along margin; segments usually irregularly 2- lobed, with elongated ovate and short deltoid-ovate lobes, very sparsely toothed, lobes and teeth narrowed to somewhat long, hard, yellowish, 8-15 mm-long spines; lower cauline leaves short-petiolate, oblong, sinuately-pinnately divided, basally narrowed to lobate, long-spiny petiole, to 30 cm long, 14 cm wide; other leaves sessile, semiamplexicaul, sinuately pinnately divided to middle or deeper, oblong or elongated-broadly lanceolate, middle leaves to 20 cm long, 8 cm wide, apical leaves strongly reduced, linearly lanceolate, lamina pinnately parted almost up to base, reduced lateral lobes abruptly ta- pered to long, hard spines. Capitula solitary or in clusters of 2—3 on long axillary shoots, less often small ones undeveloped; peduncles shorter than involucre, nodding, with 1-2 small, recurved, bracteal leaves at base of drooping capitula. Involucre (2.5)3—5 cm in dia, densely tomentose, surrounded by spreading or downward directed elongate spines; outermost involucral bracts tapered from broad base to linear-lanceolate, more or less arachnoid-hairy, recurved spines; outer bracts broadly lanceolate, glabrous, tapering to sharp, long, somewhat 7g 78 recurved spines, arachnoid-hairy in lower part; inner bracts linear- lanceolate with thin, curved, soft cusp. Florets purple-red, to 30 mm long, with limb slightly shorter than narrow tube, to 1/3 divided. Pappus yellowish-sordid-white, apically toothed tips of inner hairs longer, darker; achene blackish-brown, to 5 mm long. Flowering Au- gust to September (Plate IV, Fig. 1). Forest edges, stony habitats on slopes of the mountain gorges and river terraces.—Caucasus: Eastern Transcaucasia, Western Transcaucasia (Imeretia, rarely). General distribution: Asia Minor. Described from Georgia (gorge of Aragvi River). Type in Leningrad. Note. In southern Georgia (Bakuriani, Abastumani) this species varies considerably in the form of the leaves, length of the spines on the involucral bracts, their color and other characters. Forms transi- tional to C. adjaricum Somm. and Lev. are also observed. C. caucasicum ssp. cladophorum Petrak, described by Petrak from Adzharia, is closer to C. adjaricum in its involucral bracts and, in part, is referred by me to the latter species. The status of var. wwedenskyi Petrak [Petrak in Vestn. Tifl. Bot. Sada, No. 24 (1912) 4] reported from the former Artvin District remains unclear. The single specimen of this variety preserved in Tbilisi appears to be a teratological plant and is close to C. adjaricum in certain characters. J. Sojak [in Nov. bot. Hort. bot. Univ. Carol. Prag. (1961) 34] proposes for C. caucasicum var. wwedenskyi Petrak the new-combina- tion C. wwedenskyi (Petrak) Sojak. I feel it's premature to erect a new species on the basis of an undeveloped specimen. 9. C. adjaricum Somm. and Lev. in Nuov. Giorn. Bot. Ital. N. S. II, 1 (1895) 14, in adnot. nom. legit.; Petrak in Tr. Tifl. Bot. Sada, XII, 1, 34, descr. emend.; Grossh. Fl. Kavk. IV, 184; Charadze in FI. Gruzii, VIII, 493; Dmitr. Opred. Rast..Adzh. 277.—C. adjaricum ssp. nutans Petrak in Tr. Tifl. Bot. Sada, XII, 1 (1912) 37; Grossh. loc. cit.; Charadze, loc. cit. 493, in adnot.—C. noli-tangere Somm. and Lev. in Nuov. Giorn. Bot. Ital. N. S. I, 1 (1895) 11, non Borb. diagn. brev.; in Tr. Peterb. Bot. Sada, XVI, 255, descr. emend.—C. caucasicum ssp. nutans Petrak in schedis. (1913).—C. caucasicum ssp. cladophorum Petrak in Tr. Tifl. Bot. Sada, XII, 1 (1912) 38, quoad pl. ex Adsharia—C. puniceum Sojak in Nov. bot. Hort. bot. Univ. Carol. Prag. (1961) 37.—Ic.: Kharadze in Fl. Gruzii, VIII, Fig. 414. Perennial. Stem 30-100 cm high, sulcate, arachnoid-hairy, more or less branched above, densely leafy below but very sparsely above. Leaves. green above, densely covered with small spiny bristles, and sparse, more or less remote, thin, yellowish, usually long spines scat- tered throughout, grayish-tomentose beneath with prominent veins, 80 79 deeply sinuately pinnately divided into somewhat approximate, broadly -deltoid-ovate segments; lateral segments short, 2~3-parted into, diver- gent deltoid ovate lobes, attenuated in sharp, hard, to 15 mm-long spines, unevenly and remotely toothed with teeth bearing more or less long spines; lower cauline leaves oblong-ovate or oblong-elliptical, repandly pinnately divided almost to base, segments pinnately toothed, acute, narrowed toward base into very short, semiamplexicaul, long- spiny petiole, terminal segment, elongated, acuminate, leaves usually 15-25 cm long, 5—10 cm wide; basal leaves petiolate, 35 cm long, 18 cm wide; upper leaves reduced, ovate or ovate-oblong, sessile, semiamplexicaul, deeply pinnately divided, 6-7 mm long, to 4 cm wide; apical leaves linear-lanceolate, long-plumose-spiny, subtending capitula, usually recurved, almost as long as involucre. Capitula 2.5— 3.5 cm in dia, solitary terminal on stem and branches or on short axillary shoots near stem tip, nodding, purple, sparsely floccose in lower part, thinly arachnoid-hairy above, with black-purple or dark green spots at base of spines. Lower and middle bracts ovate, carinate, concave outside, with thick keel, abruptly narrowed to subulate tip falcately recurved at base, arachnoid-hairy or tomentose, terminating in long, yellowish, basally purple, usually recurved or erect spines; inner bracts lanceolate, with thinner, curved spines; florets purple, to 30 mm long; corolla limb divided to 1/3, almost as long as narrow tube or slightly shorter. Pappus more or less yellowish-sordid-white, with somewhat extended, apically scabrous inner bristles; achenes dark reddish-brown-brown, 5.5 mm long. Flowering August to September. Ravines of rivers, stony habitats, mid-montane to subalpine zone.— Caucasus: Western Transcaucasia (Adzharia). General distribution: Asia Minor (Lazistan). Described from Caucasus (Adzharia). Type in Florence. Note. In the Adzhar-Imeretin Range it replaces C. caucasicum (Adams) Petrak, which is common in Eastern Georgia and is reported in Bakuriani along the southern border of its range. From this locality, which is also the eastern boundary of the range of C. adjaricum, sev- eral forms transitional to C. caucasicum are known; some of them had been described as C. adjaricum ssp. nutans Petrak and were subse- quently called C. caucasicum ssp. nutans Petrak. We cannot rule out the possibility that at the confluence of their ranges these two species freely hybridize. Apparently such is the origin of C. adjaricum ssp. nutans Petrak. This subspecies has much shorter, generally recurved spines on the involucral bracts, as in C. caucasicum, but is purple- colored and floccose, as in C. adjaricum; leaves less spiny than in C. adjaricum, approaching the type of C. caucasicum. This form deserves more careful study. We also consider it better to refer C. 81 80 caucasicum ssp. cladophorum Petrak to C. adjaricum. The type ssp. cladophorum, described from Adzharia (Beshumi), is preserved in the herbarium of the Tbilisi Institute of Botany and is almost indistin- guishable from C. adjaricum in the structure of capitula; only the leaves are less deeply incised, with short and thin spines, which dif- ferentiates this form from both C. adjaricum and C. caucasicum. Apparently one of the deviating forms is the new species C. puniceum Sojak (in Nov. bot. Hort. bot. Univ. Carol. Prag. [1961] 35), described recently from Southern Transcaucasia (Adzharia, Shavshetia) on the border with Turkey. In this species the author notes characters transitional from C. adjaricum Somm. and Lev. to C. caucasicum (Adams) Petrak and speculates on its hybrid origin. Series 4. Fimbriata (Petrak) Charadze comb. nova.—Sect. Epitrachys DC. subsect. Ciliata Petrak 3. Fimbriata Petrak in Bibl. Bot. 78 (1912) 5 p. p.—Involucral bracts ciliate or with scattered short spines on margin, attenuate into spiny-ciliate apex with somewhat long spines, usually recurved in outer bracts. Leaves broadly elliptical to oblong-ovate, pinnatifid to pinnate. This series includes species of Caucasian origin, which in the Caucasus are forming closely related vicarious species, partly pen- etrating Asia Minor (Lazistan) and Iran. In the series Fimbriata, Petrak included species with ciliate involucral bracts, but which belong to different sections according to our division. I confine the scope of this series to only those species that can be referred to section Caucasigena Charadze, which are characterized by dropping and not erect capitula. 10. C. erythrolepis C. Koch in Linnaea, XVII (1843) 41; Ldb. FI. Ross. II, 726; Boiss. Fl. or. III, 528; Grossh. Fl. Kavk. IV, 180 p. p.; Charadze in Fl. Gruzii, VIII, 504; in Zam. po Sist. i Geogr. Rast. Tbil. 22, 71, descr. emend.—Epitrachys erythrolepis C. Koch in Linnaea, XXIV (1851) 398. Perennial. Stem ascending, up to 1.5 m high, densely leafy, simple, branched only above, sparsely leafy, dark-gray, sulcate, with scattered arachnoid and crispate hairs. Leaves biocolorous, dark-green above, sometimes with grayish spots, not densely spinose-hairy, at places thinly arachnoid-hairy, grayishly, loosely tomentose beneath, with prominent yellowish veins, ligneous toward apex of lobes, terminating in hard short spines; lamina pinnately dissected into irregularly 2-fid, oblong segments, with lanceolate, toothed lobes, terminating in hard, 3—4 mm- long spines; terminal segments of leaf elongate, acuminate, with 1-2 teeth on each side; all cauline leaves cuneately narrowed toward base, 82 81 basally expanded to long-spiny, semiamplexicaul auricles; middle cauline leaves 15-20 cm long, to 8 cm wide; upper leaves 5—6 cm long, 3.5 cm wide, deeply pinnately divided; apical leaves small, lan- ceolate. Capitula oblong, terminal, nodding, solitary or 2 each on long peduncles forming corymbose-paniculate common in florescence. In- volucre sparsely arachnoid-hairy, 2—3 cm in dia, elongate above, some- what narrowed at apex, with recurved, densely arachnoid-hairy, acute tip; involucral bracts dark purple, spinose; outer bracts small, lan- ceolate, subulately narrowed into long, hard spines, sparsely spinose; middle bracts with long, somewhat falcately recurved, spiny-ciliate and arachnoid-hairy subulate tip, terminating in short spine; inner bracts greenish-membranous, glabrous, linear-lanceolate, arachnoid hairy above, tapering to thin, sharp, horizontal tip. Florets dark purple-vio- let; corolla to 21 mm long, limb up to 14 mm long. Pappus dull white, inner hairs terminating in dark, scabrous-clavate tip; achenes reddish- brown, to 5 mm long. Flowering August to September. Forest glades in mid-montane zone.—Caucasus: Western Transcaucasia (Gornaya Racha), Eastern Transcaucasia (South-Ossetia). Endemic. Described from Ossetia. Type was in Berlin; isotype in Leningrad. Note. A species with a localized range, distinguished from other species of the series Fimbriata by the form of the deeply incised leaves, with elongate, slightly retrorse segments, as well as by the oblong capitula with a purple-colored involucre narrowed near the apex and short lower involucral bracts. The authors of the Flora of the Caucasus have wrongly referred specimens of C. buschianum Charadze and C. sosnowskyi Charadze to this species, which are easily distinguished from C. erythrolepis C. Koch by the form of the broader leaves and their segments and also by the globose-ovoid capitula. 11. C. tricholoma Fisch. and Mey. in DC. Prodr. VI (1837) 637; Fisch. and Mey. ex Hohen. in Bull. Soc. Nat. Mosc. VI, 251, nomen; ej. Enum. pl. Talysch. 281, descr.; Boiss. Fl. or. HI, 528 p. p.; Grossh. Fl. Kavk. IV, 528 p. p.; Charadze in Fl. Azerb. VIII, 408.—C. dimorphum Petrak ex Grossh. Fl. Kavk. IV (1934) 180, pro syn. Perennial. Stem up to 100 cm high, ascending, purple above, branched, thinly arachnoid-hairy. Cauline leaves oblong; middle leaves up to 12 cm long, 3.5—5.5 cm wide, pinnately incised into long, 2—3 fid segments, with broad sinuses; lobes of segments ovate or oblong, obtuse, with hard, up to 8 mm-long spines at tip, basally expanded into semiamplexicaul auricles, with long terminal segment of leaf, thinly spinose above, grayish-, finely-tomentose beneath; spinose-ciliate on margin. Capitula solitary, terminal on stem and its branches, to 4 cm 8 LoS) 82 in dia, projecting above leaves and nodding. Involucre finely arach- noid-hairy; outer involucral bracts narrowly lanceolate, spinescent; inner bracts linear-lanceolate, purple, spinescent. Florets purple, up to 17 mm long. Pappus yellowish-white, with longer inner hairs attenuate in toothed, somewhat flattened tip; achene to 5 mm long. Flowering July to August. Meadows, from mid-montane to subalpine zones.—Caucasus: East- ern Transcaucasia (Sarial Mountain in vicinity of Kirovabad, Murovdag Range). Endemic. Described from Sarial Mountain. Type in Leningrad; isotype in Tbilisi. Note. It is an inadequately studied species, known from the col- lections of Hohenacker from Sarial Mountain in Azerbaidzhan. This very species was later collected by A. Shelkovnikov from the Murovdag Range. De Candolle mistakenly indicated Darial instead of Sarial. Boissier in Flora Orientalis cited the specimens of Hohenacker thus: “Mons Sarial Georgiae.” However, this species does not grow in Geor- gia, although Caucasian authors have referred to it some species of the affinity of C. osseticum (Adams) Petrak, mainly from Western Transcaucasia. C. tricholoma Fisch. and May. is distinguished from all the species of this affinity by having the leaves deeply cut into nar- rower segments and the involucral bracts sparsely spinulose. 12. C. osseticum (Adams) Petrak in Tr. Tifl. Bot. Sada, XII, 1 (1912) 3 p. p.; Grossh. Fl. Kavk. IV, 180; Charadze in Fl. Gruzii, VIII, 489; in Fl. Azerb. VUI, 406.—C. osseticum ssp. eu-osseticum Petrak, loc. cit—C. fimbriatum Spreng. Syst. veg. 3 (1826) 373; DC. Prodr. VI, 635; Ldb. FI. Rossi II, 726; excl. pl. ex Imeretia; Boiss. FI. or. III, 528—Carduus osseticus Adams in Web. and Mohr. Beitr. Naturk. I (1805) 75.—Cnicus fimbriatus MB. F1. taur.-cauc. II (1808) 276.—Ic.: Kharadze in Fl. Gruzii, VIII, Fig. 413.—Exs.: Petrak, Cirsioth. Univ. No. 87. Perennial. Stem ascending, to 1 m high, with purple spots, sulcate, branched above, finely arachnoid-hairy. Leaves large, lower petiolate, others sessile; middle cauline leaves to 25 cm long, 14 cm wide, obo- vate, lamina pinnately divided to middle into broad, 2-fid, ovate seg- ments obtusely toothed with short-spinescent teeth, narrowed toward apex, auriculate, semiamplexicaul; all leaves with somewhat scattered, appressed, spiny bristles above, more or less thinly grayish- or whit- ish-tomentose beneath; upper leaves reduced; 1—2 small, usually re- curved apical leaves subtending capitula. Capitula solitary, terminal, few, in corymbose-paniculate inflorescence, 2.5—3.5 cm in dia. Involu- cre sub-glabrous in lower part, arachnoid-hairy above; outer and middle involucral bracts lanceolate, gradually tapered to linearly lanceolate, 84 83 long-spiny-ciliate tip terminating in more or less thin recurved spine; inner bracts linear-lanceolate with recurved spiny tips, basally arach- noid-hairy. Corolla bright-purple, 26-28 mm long, limb 14-15 mm long. Pappus yellowish-white with long, apically scabrous, inner hairs; achenes to 4 mm long. Flowering July to September. Forest and subalpine zones, among shrubs.—Caucasus: Dagestan, Eastern Transcaucasia (Kuba District, Kazbeg District), Southern Transcaucasia. Endemic. Described from Georgian Military Highway. Type unknown; isotype in Leningrad (?). Note. It is a widely but sporadically distributed species in the eastern Caucasus and Southern Transcaucasia. C. osseticum ssp. bornmulleri Petrak, reported by Petrak from Crimea, is so different from C. osseticum that it should have been included in section Epitrachys DC. as the separate species C. sublaniflorum Sojak. C. osseticum (Adams) Petrak was described from the Georgian Military Highway as Carduus osseticus Adams, for which the habitat was indicated to be between Lars and Darial. A specimen of this spe- cies is preserved in the Herbarium of the Botanical Institute in, Leningrad under the name C. fimbriatum MB. with the label: “Iberia. Herb. Mussin-Puschk. Acc. a D. Fischer 1835. Hb. Meyer.” I am inclined to consider this specimen an isotype of C. osseticum (Adams) Petrak, which was redescribed by Bieberstein under the name Cnicus fimbriatus, also from the Georgian Military Highway (“Ex Iberia Schlegelmilch”). Subsequently, it has been known through new collec- tions mainly from Transcaucasia. 13. C. buschianum Charadze in Addenda XXVII, 602; in FI. Gruzii, VIII, 488, descr. georg.—C. tricholoma auct. p. p. non Fisch. and Mey.: Somm. and Lev. in Tr. Peterb. Bot. Sada, XVI (1900) 253; Grossh. Fl. Kavk. IV, 180 p. p.—C. tricholoma var. aciculare Somm. and Lev. in Nuov. Giorn. Bot. Ital. N. S. II, 1 (1895) 11; in Tr. Peterb. Bot. Sada, XVI, 253; Grossh. loc. cit.—C. osseticum ssp. erythrolepis Petrak in Tr. Tifl. Bot. Sada, XII, 1 (1912) 4 p. p. non Cirsum erythrolepis C. Koch. Perennial. Stem to 100 cm high, somewhat sulcate, arachnoid- hairy, ascending, branched above. Lower and middle cauline leaves to 30 cm long, 13-15 (16) cm wide, broadly-elliptical, narrowed toward base; cauline leaves auriculate, semiamplexicaul, narrowed in upper part, acuminate, pinnately parted into ovate, less often biparted slightly antrorse segments, with usually large, spines 2-4 mm long, terminal on teeth, densely covered above with appressed spiny bristles, densely whitish-tomentose beneath; upper cauline leaves 4.5—9.5 cm long and 2-5 cm wide; apical leaves subtending capitula, linear. Capitula nod- 85 84 ding, terminal, solitary or in clusters of 2—4 at tips of elongated branches, globose-ovate, forming lax corymbose inflorescence. Involu- cre in lower part appressed, arachnoid-hairy, in upper part loosely arachnoid, to 4.5 cm in dia; outer involucral bracts usually strongly reduced, middle lanceolate, falcately recurved, more or less densely spiny-ciliate, tapered to sharp, short spines, less often spines long, hard, yellowish, recurved (f. aciculare (Somm. and Lev.) Charadze); inner bracts linearly lanceolate, scarious, tapered to long, recurved, spiny tip. Florets violet-red; corolla (23)25—27 mm long, limb to 17 mm long. Pappus sordid-white, inner hairs with elongate and toothed, somewhat enlarged tip; achenes to 5 mm long. Flowering August to September. Upper forest and subalpine zones, often in dark coniferous forest zone, and in ruble areas among tall herb vegetation.—Caucasus: Ciscaucasia (Balkaria, Karachai), Eastern Transcaucasia (western part of South Ossetia), Western Transcaucasia (Racha, Svanetia). Endemic. Described from South Ossetia. Type in Leningrad. Note. It is distinguished from the related species of the affinity C. osseticum (Adams) Petrak by comparatively larger and broader leaves and highly reduced, falcately recurved, almost tomentose, outer in- volucral bracts. Some specimens of this species are reported by au- thors of the Flora of the Caucasus as C. tricholoma Fisch. and Mey. or C. erythrolepis C. Koch, but C. buschianum Charadze is well distin- guished from both species by the form of the leaves and involucral bracts and from C. erythrolepis also by globose-ovate and not oblong capitula. 14. C. sosnowskyi Charadze in Fl. Gruzii, VIII (1952) 488, descr. georg.; in Zam. po. Sist. i Geogr. Rast. Tbil. 22, 73, diagn.—C. fimbriatum var. diversifolium Somm. and Lev. in Tr. Peterb. Bot. Sada, XVI (1900) 251.—C. osseticum ssp. tricholoma Petrak in Tr. Tifl. Bot. Sada, XII, 1 (1912) 4, non Cirsium tricholoma Fisch. and Mey.— C. erythrolepis auct. non C. Koch; Grossh. Fl. Kavk. IV (1934) 180 p. p.; Kolak. Fl. Abkhazii, IV, 269.—Epitrachys ossetica C. Koch in Linnaea, XXIV (1851) 398, non Cirsium osseticum (Adams) Petrak. Perennial. Stems 2—3, 50-60 cm high, simple or branched almost from base, ascending, more or less densely white-arachnoid-hairy. Leaves velvet-green above, densely covered with appressed, golden, more or less thin, spiny bristles, densely, grayish- and whitish-tomen- tose beneath; basal and lower cauline leaves elliptical or broadly lan- ceolate-elliptical, subacute, 15—16 cm long, 5.5—6.0 cm wide, narrowed in usually long, densely and thinly spiny-ciliate petiole, usually undivided, finely and sparsely toothed, less often pinnately parted, sparsely and thinly spiny-ciliate; cauline leaves sessile, auriculate, 8 lon 85 semiamplexicaul, obtuse or acute, lower cauline leaves subentire, sinu- ate-toothed, orbicular-ovate, other leaves deeply pinnately divided, broadly elliptical, sometimes almost lyrate, (7)9—12(17) cm long, 4.5— 7.5 cm wide, with somewhat elongate, deltoid-ovate, antrorse segments; apical leaves small, recurved. Capitula solitary, 2.0-2.5 cm in dia, nodding, in clusters of 2-5 in compact corymbose apical inflores- cence. Involucre purple or greenish, glabrous below, arachnoid-hairy above; outer involucral bracts scarious, deltoid-ovate, glabrous, with somewhat elongate, falcately recurved spiny tip, purple or greenish, carinate apex, covered from base with dense, long spiny cilia, spinescent. Florets purple, to 21 mm long, limb 12-13 mm long. Pappus sordid-white, with elongate, almost to tip plumose, inner hairs, somewhat thickened and scabrous at tip; achene blackish-brown, up to 5 mm long. Flowering July to August. (Plate VI, Fig. 2.) Mid-montane forest zone, on limestone rocks, stony alluvial de- posits, and among scrubs.—Caucasus: Western Transcaucasia. Endemic. Described from Western Georgia (Nakeral Range). Type in Tbilisi. Note. A unique species, easily distinguished from all other species of the series Fimbriata by having almost entire lower cauline leaves and more or less pinnately incised middle cauline leaves, velutinous and setose-hairy above and snow-white beneath. It is much closer to C. buschianum, from which it is distinguished by smaller capitula and acuminate, densely spinulose lower involucral bracts. Even Sommier and Levier, who described this species as a separate variety, paid attention to the peculiar structure of the leaves. 15. C. abkhasicum (Petrak) Grossh. Fl. Kavk. IV (1934) 184; Kolak. Fl. Abkhazii, IV, 270; Charadze in Fl. Gruzii, VIII, 502, excl. syn. C. tsebeldinum Woron.—C. osseticum ssp. abkhasicum Petrak in Tr. Tifl. Bot. Sada, XII, 1 (1912) 5. Perennial. Stem up to 100 cm high, branched in upper half, more or less densely leafy to top, sulcate, reddish-brown, finely arachnoid- hairy or floccose. Leaves green above, scabrous, densely covered with small appressed bristles, grayish-tomentose beneath, shallowly pinnately-parted, lobes broadly ovate with 4-6 mm-long, thin spines on tips of teeth and lobes, narrowed toward base, auriculate, semiamplexicaul, terminal segment oblong, acuminate; lower cauline leaves to 25 cm long, 15 cm wide; slightly lyrately pinnately incised, oblong-elliptical; middle cauline leaves oblong-ovate, to 12 cm long, pinnatifid in broadly ovate, 2-, less often 3-parted, somewhat antrorse lobes with round-ovate sinuses; apical leaves subtending capitula, lin- ear-lanceolate, shorter or almost equaling capitula. Capitula nodding, 87 8 oo 86 solitary, in clusters of 2—4, or terminal on stems, 2.0—2.5 cm in dia. Involucre thinly grayish-arachnoid-hairy; lower involucral bracts strongly reduced, gradually elongating from outer to inner ones, gla- brous, lustrous, abruptly narrowed in brownish-green or purple, falcately recurved, arachnoid-hairy tip, spinescent with more or less short spine; inner bracts linear-lanceolate, spinescent with thin recurved spine. Corolla red, to 24 mm long; limb to 13 mm long. Pappus brownish- white with brush-like spreading, toothed, somewhat elongate, inner hairs; achenes 4.5 mm long, dark brown, with thin yellowish stripes. Flowering August to September. Forest zone on stony slopes.—Caucasus: Western Transcaucasia (Abkhazia). Endemic. Described from Achangvar Pass in basin of Bzyb River. Type in Tbilisi. Note. The species was described from incomplete specimens col- lected by N. Albov in Abkhazia. An obscure species, known from the basin of the Bzyb River (Shvimpindzh, vicinity of Achangvar and other places). It is distinguished by involucral bracts that lack the spines, characteristic for species of this series. In spite of this, in the general appearance, form, and spiny nature of the leaves, this species must be referred to the series Fimbriata, although it occupies an intermediate position between it and the series Caucasica. 16. C. fominii Petrak in Tr. Tifl. Bot. Sada, XII, 1 (1912) 23; Grossh. Fl. Kavk. IV, 180; Kolak. Fl. Abkhazii, IV, 272; Charadze in Fl. Gruzii, VIII, 503.—C. tricholoma auct. non Fisch. and Mey.: Alb. Prodr. Fl. Colch. (1895) 143.—Ic.: Kolak. Rast. Mir. Kolkh. (1961), Bigs 73: Perennial. Stem dark-purple or greenish with purple spots, slightly angular and weakly sulcate, floccose-arachnoid-hairy, to 1 m high, slightly branched above. Middle cauline leaves to 10 cm long, to 4 cm wide, ovate-elliptical, deeply pinnately divided into 2-fid deltoid-ovate lobes with broad sinuses; lamina tapered toward apex, narrowed at base, semiamplexicaul, covered above with appressed, yellowish, spiny bristles, grayish-tomentose beneath, with prominent, subglabrous veins, lobes acute, spinescent with hard spines 7-9 mm long, unequally spiny- ciliate; apical leaves 1-2, linearly lanceolate, spiny-ciliate, toothed, recurved. Capitula solitary, terminal on stem and short branches, nod- ding, 3—4 cm in dia. Involucre more or less densely grayish-arachnoid- hairy; outer involucral bracts lanceolate, spinescent with short recurved spine; middle bracts spinescent with sharp, elongate, recurved spine, sparsely spiny-ciliate; inner bracts linear-lanceolate, spinescent with recurved, thin spine. Florets purple; corolla to 27 mm long, limb al- most equaling narrow tube or somewhat shorter. Pappus sordid-white, 8 \o 87 inner bracts somewhat longer and apically somewhat enlarged, sca- brous; mature achenes unknown. Flowering July to August. Meadows in subalpine zone, forest edges Caucasus: Western Transcaucasia (Abkhazia; Yaps-Khu, Atsgyrkha, Mamdzyshkha, Dzishra mountains). Endemic. Described from Abkhazia. Type in Tbilisi. Note. The species was described from an incomplete specimen preserved in the herbarium of the Institute of Botany, Academy of Sciences of the Georgian SSR. Material similar to the type was later collected in Abkhazia by various authors. We have supplemented the description of the species from the collections of A.A. Kolakovsky from Yaps-Khu (Bzyb Range). Petrak allies this species with C. caucasicum (Adams) Petrak and C. osseticum (Adams) Petrak, hy- pothesizing a hybrid origin of C. fominii. However, the first of these species does not grow in Western Georgia, and the presence of abun- dant, well developed pollen in C. fominii, which was reported for the first time by A. Kolakovsky in Flora Abkhazii [Flora of Abkhazia], casts doubt on a hybrid origin of this species. The habit characters and presence of spiny cilia on the involucral bracts bring C. fominii close to species of the series Fimbriata, although the presence of hard spines and the form of its leaves put this species in a somewhat isolated position rather closer to the series Caucasica. 17. C. caput-medusae Somm. and Lev. in Nuov. Giorn. Bot. Ital. N. S. II, 1 (1895) 10; in Tr. Peterb. Bot. Sada, XVI (1900) 253; Petrak in Tr. Tifl. Bot. Sada, XII, 1, 6; Grossh. Fl. Kavk. IV, 179; Kolak. FI. Abkhazii, IV, 270; Charadze in Fl. Gruzii, VIII, 507.—Ic.: Somm. and Lev. loc. cit. (1900) tab. XXIV; Charadze in FI. Gruzii, VIII, Fig. 415.—Exs.: Petrak, Cirsioth. Univ. No. 88. Perennial. Stem 40-100 cm high, sulcate, greenish, with purple spots, thinly arachnoid-hairy, highly branched above with spreading branches and numerous small capitula in corymbose-paniculate com- mon inflorescence. Leaves more or less densely covered above with appressed spiny bristles, usually with scattered, solitary, long spines, thinly grayish-tomentose beneath, with more or less prominent veins; all leaves oblong or broadly lanceolate, acute; basal leaves to 20 cm long, 10 cm wide, petiolate; cauline leaves with broad, semiamplexicaul auricles; middle leaves 15 cm long, 7 cm wide, deeply pinnately in- cised into 2—3-parted, unevenly toothed, broadly lanceolate or deltoid- ovate, sub-acute segments with 4-5 mm long, thin spines on margin and hard spines to 12 mm long, terminal on teeth; apical leaves small, lanceolate, long-spinose-pinnately toothed. Capitula nodding, small, solitary, globose, 1.5—2.0 cm in dia by end of flowering, surrounded by up to 2 cm long, yellowish, remote spines; terminal capitula on 90 88 somewhat short branches, and axillary on long branches. Involucr with smooth, long, divergent spines, narrowed above, with recurve arachnoid-hairy bracts; involucral bracts lanceolate, more or less longs densely-pectinately-spiny-ciliate on margin; outer and middle bract dark-purple, spinescent; inner bracts sublinear, spinescent with thi recurved spine, arachnoid-hairy above. Corolla purple, 15—16 mm long limb almost 2 times as long as narrow tube. Pappus sordid-white, to 1 mm long, with slightly elongate, brownish, toothed tips of inner hairs achenes 2.5 cm long (immature). Flowering July to August. Forest and subalpine zones, along forest edges and in trample meadows.—Caucasus: Western Transcaucasia (Svanetia, Megrelia Adzharia, Abkhazia). Endemic. Described from Upper Svaneti: (Mestia). Type in Florence. Note. A rare species described from Upper Svanetia. A topotyps of this species, collected by D.I. Sosnowsky, is preserved in Tbilisi From seeds of these plants sent by Sosnowsky to Petrak, specimen: were grown and subsequently reported on by Petrak in his work Cirsiotheca Universa. Among the species of the series Fimbriata this species is unique in the form of its small, globose, little-opened ca- pitula, and the form of its involucral bracts. Section 2. Epitrachys DC. in Duby. Bot. Gall. I (1828) 286 p. max p.; Koch Syn. (1837) 392; C. Koch in Linnaea, XXIV, 396 p. p., pro gen.; Boiss. Fl. or. HI, 523, emend. p. max p.—Eriolepis Cass. in Dict. Sc. nat. XXXV (1825) 172, pro gen.; DC. Prodr. VI, 635 p. p., prosect.—Lophiolepis Cass. in Dict. Sc. nat. XLI (1826) 313, pro gen.; DC. Prodr. VI, 634 p. p., pro sect— Leaves more or less densely covered above with tiny spiny bristles. Capitula erect, on short, less often long peduncles, somewhat surpassing apical leaves, or subsessile, subtended by apical leaves, more or less surpassing capitula. Involucre arachnoid-hairy, less often subglabrous; involucral bracts gradually elongated from outer to inner, smooth or spiny-ciliate on margin; outer bracts usually narrowed from middle attenuated into upward directed or decurved spines, less often spines shorter than bracts. Corolla limb irregularly 5-parted almost to middle, almost equal or half as long as narrow tube. Pappus shorter than florets. Achenes (4)5—7 mm long. Type of section: C. italicum DC. Note. Segregated and published by De Candolle for the first time in 1828 (DC. apud Duby, 1. c.), the section included several species growing in Southern Europe, particularly C. italicum DC., C. lanceolatum (L.) Scop., C. eriophorum Scop., and others. Later, Boissier included in the section a number of Eurasian and Caucasian species. I am restricting the scope of this section some- 89 what by including species with nodding capitula in the separate section Caucasigena Charadze. Species of the section Epitrachys DC. are distributed in arid and mountainous regions of the Mediterranean, West Asia, the Caucasus, and occasionally Soviet Central Asia. Detailed intrasectional division was proposed by Petrak (Petrak in Bibl. Bot. 78 [1912] 5), who erected a number of subsections. Petrak considers the mountains of Soviet Central Asia, with one species—C. turkestanicum (Rgl.) O. and B. Fedtsch., as the eastern boundary of the section Epitrachys. In 1938 he described (Petrak in Fedde, Repert, XLIII, 271) C. interpositum Petrak from Yunnan with characters intermediate between the sections Onotrophe (Cass.) DC. and Epitrachys DC. The most significant dis- tinguishing character between the two sections is the presence of spiny bristles on the upper surface of the leaves in Epitrachys DC. and their absence in Onotrophe (Cass.) DC. The latter section, including the type of the genus, must be named Cirsium. Subsection 1. Congesta Petrak in Bibl. Bot. 78 (1912) 5.—Cauline leaves sessile, coriaceous, stiff. Capitula ovate-oblong, clustered in capitate inflorescence. Involucre glabrous or with scattered arachnoid hairs; involucral bracts scabrous or smooth on margin, closely ap- pressed, scarious, spinescent with hard, long, spreading or upward- directed spines. Achenes large, 6—7 mm long. Type of subsection: C. congestum Fisch. and Mey. ex DC. Note. Petrak considers four Western Asian species, two of which penetrate also into southern regions of Transcaucasia, to comprise this subsection. Petrak notes the unique position of the species of this subsection, which approach species of the subsection Bracteosa Petrak. As elements of the section Epitrachys DC., species of this subsection are distinguished by having the stiffest leaves and largest achenes. 18. C. congestum Fisch. and Mey. ex DC. Prodr. VI (1837) 641; Boiss. Fl. or. III, 525; Petrak in Izv. Kavk. Muz. VIII, 1-2, 37; in Nabelek, It. turc.-pers. II, 33; Grossh. Fl. Kavk. IV, 178; Charadze in Fl. Turkm. VII, 241; in Fl. Azerb. VIII, 405.—Exs.: Sint. It. transcasp. pers. No. 959. Perennial. Rhizome thick, woody, with brown scaly remnants of old leaves at base of stem; stem hard, ascending, slightly angular, sulcate, usually glabrous or with scattered arachnoid hairs, simple or weakly branched in upper part, 50(60)—120(150) cm high. Leaves coriaceous, stiff, sparsely spinose above with very thin arachnoid pubescence, subglabrous beneath, with scattered, short, curly hairs and prominent veins, more or less deeply pinnately lobed, with thin, sparse 90 9 _— spines on margin and to 15—18 mm long, hard, yellowish spines termi- nal on deltoid, acuminate lobes; basal leaves oblong, 30—45 cm long, 10-12 cm wide, long-petiolate, pinnatifid into broadly deltoid, 3—5- parted, acuminate, spinescent lobes; cauline leaves to 25 cm long, 7— 8 cm wide, 10-15 cm long toward stem apex, somewhat distant, re- curved, ovate or broadly-lanceolate, at base broader, semi-amplexicaul, at apex tapered to lanceolate segment, leaf sinuately-pinnatifid in 2— 3-parted short-deltoid lobes with long, stiff spines. Capitula oblong, 2.5-3.5 cm long, 1.5—2.5 cm in dia, conically narrowed at base, subsessile, in axils of lanceolate, pinnatifid, long-spiny leaves, usually slightly surpassing capitula; capitula numerous, clustered at stem tip, less often axillary, in capitate inflorescence. Involucre glabrous, with upward directed, spreading, usually long spines; involucral bracts sca- brous on margin, very thinly spiny-ciliate or toothed, at base of ca- pitula usually short-subulate; outer involucral bracts subulately nar- rowed from ovate-lanceolate base into yellowish, more or less thin, hard, upward directed or slightly recurved spines, 2—3 times as long; inner bracts linearly lanceolate, scarious, narrowed into somewhat short spines. Florets pale-pink, 22—24 mm long; corolla limb irregularly incised almost to middle, somewhat shorter than tube. Pappus yellow- ish from basally almost scaly, lustrous, sparsely plumose hairs and apically somewhat cusp-like enlarged inner hairs; achenes to 7 mm long. Flowering August to September. Clayey-rubbly slopes, in mountain steppe and semi-steppe zone, from 700-1200 m.—Caucasus: Southern Transcaucasia (Daralagez, Nakhichevan ASSR, Megri); Soviet Central Asia: mountainous Turkmenia (Karakala District). General distribution: Armenia and Kurdistan, Iran. Described from Kurdistan from Sovich’s specimens. Type in Leningrad. Note. De Candolle considered C. consanguineum DC., described from the collection of Belanger from Iran (DC. Prodr. VI, 641), as a separate species. Later, Boissier (Fl. or. III, 526) cites it as C. congestum B. consanguineum (DC.) Boiss, distinguishing it from the typical form by its longer capitula. 19. C. sorocephalum Fisch. and Mey. Ind. Sem. Hort. Petrop. 2 (1835) 33; in Hohen. Enum. pl. Talysch. 281; DC. Prodr. VI, 636 p. p. excl. pl. Meyeriana: Somm. and Lev. in Nuov. Giorn. Bot. Ital. N. S. II, 1 (1895) 9; Boiss. Fl. or. III, 526; Ldb. Fl. Ross. II, 727; Petrak in Izv. Kavk. Muz. VIII, 1-2, 37, descr. emend.; Grossh. Fl. Kavk. IV, 178; Charadze in Fl. Azerb. VIII, 405. Perennial. Stem strong, ascending, (40)50—75 cm high, sulcate, 92 glabrous or more or less scatteredly arachnoid-hairy, simple, less often 93 91 in upper part with few axillary branches. Leaves stiff, more or less coriaceous, usually glabrous, with scattered small, erect spines above, glabrous beneath or at places thinly arachnoid-hairy, with very promi- nent veins; lower leaves oblong or oblong-lanceolate, to 30 cm long and 10 cm wide, narrowed to winged petiole, sparsely spiny-toothed, short-acuminate, deeply sinuately pinnately lobed with broadly ovate, 2—5-parted, deltoid lobes; teeth tapered into hard, yellowish, to 20 mm long, spines; upper cauline leaves gradually reduced, middle to 10 cm long, 5 cm wide, ovate-oblong, sessile, auriculate, semiamplexicaul, remotely sinuately pinnatifid to 1/3, with deeply 2—3-parted lobes, lobes lanceolate-deltoid spinescent; apical leaves narrowly lanceolate, deeply sinuate pinnatipartite, with short triangular lobes spinescent, with long stiff spines, 2-3 times as long as capitula. Capitula ovate- oblong, 20-30 mm long, 14—20 mm in dia, narrowed toward base, in clusters of 4—6(10), terminal on stem and short proximate branches, in umbellate-capitate inflorescence, apical leaves projecting above ca- pitula. Involucre glabrous; lower and middle bracts imbricate, middle bracts coriaceous, slightly carinate; bracts gradually becoming longer from outer ovate to inner linear-lanceolate, scabrous on margin, very thinly spiny-ciliate, gradually tapered into prickly short spine, some- times with brown spot in upper half. Florets pale-pink, 21 mm long; corolla tube scarcely longer than limb. Pappus sordid-white, at base yellowish, consisting of thin scaly hairs, inner hairs with long toothed tips; achenes brownish, with dark striations, 6.5 mm long. Flowering July to August. Dry stony slopes, from mid-montane to subalpine zone.—Caucasus: Talysh (Lenkoran). General distribution: Afghanistan. Described from Talysh. Type in Leningrad; isotype in Tbilisi. Note. The range of this species is still unclear. So far it is known from Talysh and is reported for Afghanistan (Petrak in Koje and Rechinger, Symp. Afgh. II [1955] 180; Kitamura, F/. Afgh. [1960] 408). We do not rule out the possibility of its occurrence also on the Iranian upland. De Candolle (DC. Prodr. V1, 636) cited the specimen of C. aduncum Fisch. and Mey. ex DC. (= C. fallax Fisch. and Mey.) from Meyer’s herbarium under this name. Subsection 2. Bracteosa Petrak emend. Charadze.—Subsect. Bracteosa Petrak 2. Rigida Petrak in Bibl. Bot. 78 (1912) 6, excl. 1. Phyllocephala Petrak.—Subsect. Microcephala Petrak, ibid. 6.—Leaves sessile or scarcely decurrent on stem, more or less coriaceous, stiff, glabrous or thinly arachnoid-hairy. Capitula usually medium-sized or small, globose-ovate or ovate. Involucre glabrous or weakly arach- noid-hairy; involucral bracts usually entire, abruptly attenuated into 94 92 straight or more or less spreading spines, almost as long as involucral bracts, less often shorter or slightly longer. Type of subsection : C. bracteosum DC. Note. Petrak includes two series in the subsection Bracteosa Petrak: 1) Phyllocephala Petrak and 2) Rigida Petrak. From the first group, C. bornmiilleri Sint. ex Bornm. grows on the territory of the USSR; it so markedly differs in the structure of the involucral bracts and, in part, the pappus and achenes that it can be treated as a separate subsection. It seems more natural to include C. aduncum Fisch. and Mey. ex DC. in the subsection Bracteosa, which is considered by Petrak as the separate subsection Microcephala Petrak. The subsection Bracteosa Petrak includes the group of Caucasian- Western Asian species. 20. C. bracteosum DC. Prodr. VI (1837) 641; Boiss. Fl. or. III, 532 p. p. excl. var. B.; Somm. and Lev. in Nuov Giorn. Bot. Ital. N. S. II, 1, 12; Petrak in Nabelek. It. taurc.-pers. II, 33; Charadze in FI. Azerb. VI, 404.—C. strigosum Fisch. and Mey, ex DC. Prodr. VI (1837) 640 p. p. quoad pl. ex Armenia et Persia; Petrak in Izv. Kavk. Muz. VIII, 1-2, 56, non MB.—C. strigosum var. gymnocalycina Trautv. in Tr. Peterb. Bot. Sada, II, 2 (1873) 555. Perennial. Stems slightly angular, sulcate, usually glabrous or with scattered crisped hairs, simple, branched above with long, upward spreading axillary branches. Cauline leaves numerous, to 80 mm long, 15 mm wide (excluding spines), divergent, slightly rugose above, cov- ered with numerous tiny, more or less appressed spines, midrib very prominent beneath, convolute, lanceolate, distantly pinnately lobed, sessile, slightly decurrent with auricles, with long, acute, more or less narrowly lanceolate terminal segment, spinescent with 5-6 mm long stiff spine; leaf lobes 2-parted, with small deltoid lobes, proximal lobules almost reduced, strongly attenuated into long, hard, more or less thin, sharp, straw-yellow spines 10—12(15) mm long, ciliate, with thin, hard, 2-4 mm-long spines on margin; upper cauline leaves to 4 cm long, broadly lanceolate; apical leaves subtending capitula, 3-S, narrow-lanceolate, more or less pinnately dentate, usually 2—3 times as long or almost equal to capitulum. Capitula terminal and axillary, solitary,’more or less numerous; terminal capitula sessile, to 25 mm long, 20 mm in dia, less often larger; axillary capitula 8-12 mm long, on elongated lateral branches and with terminal capitula form short racemes in corymbose-paniculate common inflorescence. Involucre glabrous or with occasional arachnoid-hairs; involucral bracts oblong- lanceolate; lower bracts almost ovate-oblong, glabrous, carinate, along margin scabrous, usually tapered from middle in thin, carinate, more 95 93 or less falcately recurved tip, strongly extended into thin, spines 2—4 mm long; inner bracts scarious, spinescent, with more or less recurved scaly tip. Florets purple, 15—18 mm long; corolla tube somewhat longer than limb. Pappus sordid-yellowish-white, of few hairs, inner hairs plumose in lower part and clavate-toothed above; achene brown, to 5 mm long. Flowering July to August. Among phrygana vegetation in mid-montane zone.—Caucasus: Southern Transcaucasia (Nakhichevan ASSR). General distribution: Armenia and Kurdistan, Iran. Described from Iran from Belanger’s specimens without precise indication of locality. Type in Geneva. Note. In Leningrad (Herbarium of the Botanical Institute of the Academy of Sciences of the USSR), the Sovich specimen from Kurdistan (Pere) is preserved, which is cited by De Candolle as C. strigosum Fisch. and Mey. ex DC.; it also is cited by Boissier under the name C. bracteosum DC. Plants of Southern Transcaucasia are similar to this specimen. I also examined the Aucher-Eloy specimen (Elbourz, No. 4813) cited by Boissier and identical to Transcaucasian specimens. However, the Kotschy specimen from Kuh-Daena is con- siderably different from the above-mentioned specimens. The specimen from Kazikibaran from Radde’s collections (upper reaches of the Araks) was described by Petrak (op. cit.) as C. strigosum (MB.) MB. and is cited by Trautvetter as C. strigosum var. gymnocalycina Trautv. According to the diagnosis, the Transcaucasian specimens differ by the more or less fine arachnoid pubescence and the dimensions and structure of the capitula. This species occupies an intermediate position between C. strigosum (MB.) MB. and C. aduncum Fisch. and Mey. ex DC. It is distinguished from the first species by ovoid- or globose-oblong capitula and long-acuminate, erect, but not falcately recurved, involucral bracts, and from the second species by oblong capitula and attenuated, thin, spinescent involucral bracts. The variability of C. bracteosum has not yet been adequately stud- ied. I did not have the opportunity to examine its type; hence it is not possible to discuss the specific identity of close forms growing on the territory of Transcaucasia and Western Asia. 21. C. strigosum (MB.) MB. FI. taur.-cauc. III (1819) 558 p. p.; in Fisch. cat. Hort. Gorenk. 35; Link, Enum. pl. II, 300; C.A.M. Verzeichn. 69; DC. Prodr. VI, 640 p. p.; Boiss. Fl. or. III, 536 p. p. excl. B. khorassanicum Boiss; Ldb. FI. Ross. II, 728; Somm. and Lev. in Nuov. Giorn. Bot. Ital. N. S. II, 1, 14; Grossh. Fl. Kavk. IV, 181; Opred. Rast. Kavk. 483; Charadze in F1. Azerb. VIII, 403.—C. rigidum Petrak in Vestn. Tifl. Bot. Sada, Vyp. 24 (1912) 5 p. p. quoad pl. ex Ata-Czai, non DC.—Carduus strigosus MB. Beschreib. Land. Casp. 96 94 Meer. (1800) 194, diagn.—Cnicus strigosus MB. FI. taur.-cauc. II (1808) 277. Perennial. Stem erect, strong, up to 50 cm high, branched above, slightly angular, glaucous, glabrous or with scattered arachnoid hairs, to tip densely leafy. Leaves glaucous, stiff, upper surface not densely covered with more or less erect, tiny spines, glabrous or with scattered arachnoid hairs, glabrous beneath or finely arachnoid-hairy on veins, with very prominent network of veins, and short, upward directed spines on margin, ovate-oblong or broadly-lanceolate; basal and lower cauline leaves pinnately incised into remote 2-parted segments, with oblong lobes strongly narrowed into hard spines 2—5(7) mm long, narrowed toward base into spiny petiole; other cauline leaves sessile, auriculate, amplexicaul, long-acuminate, lobes terminating in hard spines, upto 12 cm long, 5 cm wide, sinuately-pinnately shallow cut in 2-parted, ovate or ovate-deltoid lobes, upper lobes reduced, lower oblong-ovate, strongly spinescent; leaves of lateral branches somewhat reduced, sur- passing axillary capitula on very short peduncles; apical leaves lin- early-lanceolate, adpressed to capitula and usually surpassing them. Capitula numerous, terminal and axillary, in corymbose-paniculate common inflorescence, globose-ovate, narrowed toward base, short- pedunculate, 1.5—2.5 cm in dia; axillary capitula often reduced; usu- ally undeveloped. Involucre glabrous, less often sparsely arachnoid- hairy; involucral bracts appressed, imbricate, gradually becoming longer from outer to middle, oblong or ovate-elliptical, lustrous, straw-yel- low, with dark-green stripe, carinate, tapered above middle to carinate, dark-colored, more or less falcately recurved tip strongly spinescent with thin, up to 4 mm long spines; inner bracts linearly lanceolate, scarious, tapered to recurved sharp points. Florets pale-pink; corolla tube narrow, almost 2 times as long as limb. Pappus sordid-white, at base yellowish, with toothed, somewhat clavate tips of elongate inner hairs; achenes to 5 mm long. Flowering July to August. Found up to mid-montane zone, on dry rubbly slopes and in river ravines.—Caucasus: Eastern Transcaucasia (Kuba District). Endemic. Described from Eastern Transcaucasia (Shemakha). Type in Leningrad. Note. This species is usually confused with C. rigidum DC., from which it is easily distinguished by its numerous capitula, usually not rising above the subtending leaves, and much more highly branched, densely leafy stem. Boissier refers Bunge’s specimen from Iran (Khorasan) to this species under the name B. khorassanicum, which is distinguished by its larger capitula, and, apparently, is a separate spe- cies. Under the name C. strigosum Fisch. and Mey. ex DC., De Candolle mentions the specimen from Kurdistan (Pere), cited by Boissier as C. bracteosum DC. The entire group of species of the affinity C. bracteosum DC. undoubtedly needs a critical review. 97 95 22. C. rigidum DC. Prodr. VI (1837) 640; C. Koch in Linnaea, XVII, 41; Ldb. FI. Ross. II, 728; Petrak in Vestn. Tifl. Bot. Sada, Vyp. 24, 5 p.p. quoad pl. ex Ardanucz; Grossh. and Schischk. Sched. ad Pl. or. exs. 14; Grossh. Fl. Kavk. IV, 183; Opred. Rast. Kavk. 484; Charadze in FI. Gruzii, VIII, 514, descr. georg.—C. strigosum MB. FI. taur.-cauc. III (1819) 558 p. p.—C. arachnoideum Boiss. FI. or. III (1975) 536 p. p. quoad. pl. Stevenianas, non MB.—C. aspinellum Sojak in Nov. bot. Hort. bot. Univ. Carol. Prag. (1961) 36.—Cnicum strigosus Stev. in Mém. Soc. Nat. Mosc. IV (1812-1813, reimpr. 1830) 102, non MB.—BE&pitrachys rigida C. Koch in Linnaea, XXIV (1851) 400.—Exs.: Pl. or. exs. No. 48. Perennial. Stem at base woody, usually solitary, ascending, slightly sulcate, sparsely arachnoid-hairy, in lower part white-arachnoid-floc- cose, branched almost from middle, 20-60 cm high. Leaves more or less stiff, usually subglabrous, above, with scattered, appressed, tiny spines, glaucous beneath, usually with prominent thin veins, more or less occasionally arachnoid-hairy on midrib, margin ciliate from tiny, thin, upward directed spines; basal and lower cauline leaves oblong, almost to base pinnately parted into distant, deeply 2-parted, oblong- elliptical segments, narrowed in narrow-winged, long, floccose-white- hairy petiole, acuminate, with petiole 23 cm long, 8—10 cm wide; leaf segments almost to base 2-parted, with oblong, short-acuminate, elon- gate lobes, upper of them usually reduced, with hard spines; middle cauline leaves sessile, broadly amplexicaul, toothed with more or less reduced, almost ovate-oblong segments; leaves in upper part of stem and on elongate branches more or less remote, usually stiff, almost ovate, broadly amplexicaul, sessile, strongly spinescent with more or less thickened spines 7-8 mm long, shallow sinuately pinnately di- vided into broadly ovate lobes; their secondary lobes deltoid-ovate, strongly spinescent. Capitula terminal on stem and branches, solitary, erect ovate or globose-ovate, 35-40 mm long, 25—35 mm in dia, usu- ally rising above apical leaves, sometimes with 1—2 linear-lanceolate leaves, distant from capitula, ciliate, small, spinescent. Involucre gla- brous, with occasional, scattered, arachnoid hairs and imbricate bracts; outer bracts ovate, middle linearly lanceolate, coriaceous, glabrous, straw-yellow, lustrous, gradually tapered into sharp, dark-green, cari- nate, somewhat recurved tip, strongly spinescent with spines 2-4 mm long; inner bracts scarious, linearly lanceolate, acuminate, falcately recurved, scaly tip. Florets pink, 28 mm long, with corolla tube up to 15 mm long. Pappus at base yellowish, sordid-white above, inner hairs apically scabrous, slightly enlarged, clavate; achenes light brown, 5.5 mm long, 3.3 mm wide. Flowering July to August. Dry slopes, in mid-montane zone.—Caucasus: Western Transcaucasia (vicinity of Tbilisi, Gori), Southern Transcaucasia 98 96 (Meskhetia). General distribution: Armenia and Kurdistan (Ardanuch). Described from Aragvi River ravine. Type in Geneva; isotype in Leningrad. Note. C rigidum DC. has been confused with C. strigosum (MB.) MB. and C. arachnoideum (MB.) MB. From the former it is clearly distinguished by the shape of the capitula and short-acuminate involu- cral bracts; from C. arachnoideum (MB.) MB. it differs significantly and can be compared only in the form of the lower cauline leaves. Steven’s specimen, from which De Candolle described C. rigidum DC., was included by Marschall-Bieberstein in C. strigosum (MB.) MB., the plant characteristic for the eastern Caucasus. Later, Petrak (op. cit.) described C. strigosum (MB.) MB. in detail from two specimens, one of which is C. rigidum DC. from the former Artvin District (Ardanuch) and the other is C. strigosum (MB.) MB. from the Kuba District. C. rigidum DC. is distributed in Central Transcaucasia, pen- etrating to Lazistan. In 1961, J. Sojak described this same species as C. aspinellum Sojak (1. c.), comparing it with C. strigosum (MB.) MB., from a speci- men collected by Grossheim and Kolakovsky from the vicinity of Tbilisi (Olginskoe). 23. C. aduncum Fisch. and Mey. ex DC. Prodr. VI (1837) 636; Ldb. FI. Ross. II, 728; Petrak in Tr. Tifl. Bot. Sada, XII, 1, 20; Grossh. and Schischk. Sched. ad FI. or. exs. 25; Grossh. Fl. Kavk. IV, 183; Opred. Rast. Kavk. 484; Charadze in Fl. Azerb. VIII, 403.—C. fallax Fisch. and Mey. in Hohen. Enum. pl. Talysch. (1838) 280; Boiss. FI. or. III, 535; Ldb. Fl. Ross. II, 728; Somm. and Lev. in Nuouv. Giron. Bot. Ital. N. S. II, 1, 13.—C. sorocephalum DC. Prodr. VI (1837) 636 p. p. non Fisch. and Mey.—C. calvescens Sojak in Nov. bot. Hort. bot. Univ. Carol. Prag. (1961) 35.—Exs.: Pl. or. exs. No. 96. Perennial. Root ligneous. Stem strong, ascending, sulcate, more or less scatterdly crisped-hairy, at base with dark-brown remnants of basal leaves to 60 cm high, densely leafy, strongly branched above, with somewhat flexuous branches bearing numerous small capitula in dense corymbose paniculate common inflorescence. Leaves very sparsely arachnoid-hairy above, with somewhat dense, appressed, tiny spiny bristles, glabrous beneath, with prominently raised midrib and usually long arachnoid-hairs on veins, deeply sinuately pinnately divided into linearly lanceolate segments, spiny-ciliate on margin, with long acute terminal segment, lateral segments 2-parted, proximal lobes short, strongly spinescent, with up to 12 mm long spines; basal leaves usu- ally with more or less elongate, winged petiole; cauline leaves oblong, 9-18 mm long, 4—9 mm wide, sessile, semiamplexicaul, almost to base 99 97 sinuately-pinnately parted, with oblong-lanceolate 2-parted segments, 2-5 cm long, with strongly reduced deltoid-lanceolate proximal lobe with spinescent lobules; apical leaves narrowly lanceolate, pinnately lobed, long-spiny, closer to capitula, usually not surpassing capitula. Capitula terminal, in clusters of 2-3 on very short peduncles, less often solitary, ovate, 15—20 mm in dia. Involucre sparsely arachnoid- hairy; outer involucral bracts ovate, inner oblong-lanceolate, straw- yellow, slightly carinate, glabrous, weakly arachnoid-hairy on margin in upper part, abruptly narrowed to erect or recurved spine; inner bracts linearly lanceolate, acuminate, with recurved scaly apex. Florets red, 17-19 mm long; corolla tube narrow, to 12 mm long, limb 7—8 mm long, 2/3 parted, usually recurved in ray florets. Pappus yellowish, at tip sordid-white, with few plumose hairs of almost same length; inner hairs scabrous at tip, somewhat clavate; achenes to 5 mm long, dark brown. Flowering July to August. (Plate V, Fig. 1.) Rubbly habitats and among scrubs, from mid-montane to subalpine zone.—Caucasus: Southern Transcaucasia (Novo Bayazet, Agmagan, Karabakh, Nakhichevan ASSR), Talysh (Zuvant). General distribution: Armenia and Kurdistan. Described from Karabakh. Type in Leningrad. Note. The status of C. aduncum Fisch. and Mey. ex DC. is still not entirely clear. Boissier, the first to unite this species with C. fallax Fisch. and Mey., described from Hohenacker’s specimens from Zuvant (Lenkoran), preferred the latter name over the prior name, C. aduncum Fisch. and Mey. ex DC. The specimen of C. fallax Fisch. and Mey. from Lenkoran was wrongly cited by De. Candolle under the name C. sorocephalum Fisch. and Mey. The question of separate status for C. fallax still needs to be re-examined on the basis of new material. Petrak (op. cit.) showed the separateness of these two species, allying C. fallax with the Western Asian species C. bracteosum DC. and C. fraternum DC. The presence of transitional forms between the type specimens of C. aduncum and C. fallax does not allow one to delimit these species. The length of the spines on the involucral bracts, pubes- cence, size of the capitula, and other characters vary, but the structure of florets deserves special attention. As a characteristic feature, Petrak mentions C. fallax as having florets with an abrupt transition from the narrow corolla tube to the limb, whereas in C. aduncum the corolla tube is gradually expanded: There is a basis to presume that the phenomena of sexual dimor- phism are observed in this particular case. Subsection 3. Phyllocephala (Petrak) Charadze comb. nova.— Subsect. Bracteosa Petrak 1. Phyllocephala Petrak in Bibl. Bot. 78 (1912) 6.—Capitula ovate, narrowed above, subsessile, surrounded by bracteal leaves surpassing them. Leaves coriaceous; cauline leaves 100 98 sessile, with large amplexicaul auricles, somewhat decurrent. Involu- cral bracts numerous, membranous, appressed, spine scent; inner hairs of pappus long-plumose only in lower part, scabrous above. Note. It differs from the subsection Bracteosa Petrak by having the involucral bracts completely closed up to the end of flowering and tiny, thin, apical spines at the tips of the bracts. Type of subsection: C. phyllocephalum Boiss. 24. C. bornmiilleri Sint. ex Bornm. in Fedde, Repert. VIII (1910) 260; O. and B. Fedtsch. Perech. Rast. Turk. IV, 282; Charadze in F1., Turkm. VII, 241.—Ic.: Charadze in Fl. Turkm. VII, Plate XX VIII.— Exs.: Sint. It. transcasp.-pers. No. 690. Perennial. Root thick, producing solitary, less often many stems. Stem ascending, to 60 cm high, glabrous or rarely arachnoid-hairy, sulcate, divergently branched above, basally with blackish-brown fi- brous remnants of old leaves. Leaves coriaceous, with divergent spiny bristles above, glabrous or with fine, thin, arachnoid-pubescence be- neath, glaucous with prominent whitish veins, deeply pinnately di- vided into 2—3-parted, linear-lanceolate segments, strongly spinescent, spines up to 7 mm long; basal leaves to 18 cm long, long-petiolate, petiole spiny-toothed; cauline leaves to 9 cm long, narrowed into lin- ear-lanceolate tip, basally broadened, auriculate, semiamplexicaul, ob- liquely decurrent; apical leaves almost up to base incised into nar- rowly lanceolate, long-spiny segments, subtending capitula, and slightly surpassing or almost 2 times as long. Capitula solitary or in clusters of 2—4, terminal on stems and branches, less often axillary, ovate, cu- neate, 2.5 cm long and 1.5—2.0 cm in dia, short-pedunculate, borne in elongate paniculate common inflorescence. Involucral bracts numer- ous, appressed, imbricate, glabrous at apex, roundish in upper portion, floccose-arachnoid-hairy on margin, with short, filiform purple spine at apex. Corolla purple, 25-27 mm long; limb to 15 mm long. Pappus sordid-white, to 18-20 mm long; inner hairs clavate, toothed, long- plumose only in lower part, short-plumose to setose above. Achenes reddish-brown-brown, somewhat cuneately narrowed toward base, to 6 mm long. Flowering July to October. Zone of mixed herb-grass steppe, among cushion-plant formations and in old fields from 1,200 to 2,000 m.—Soviet Central Asia: moun- tainous Turkmenia. General distribution: northern Iran. Described from Kopetdag (Sulyukli). Type in Brno. Note. It is somewhat separate from all other species of the section by having ovate capitula, imbricate, tight involucral bracts with fili- form, small spines, inner hairs of pappus long-plumose only in the lower part, and achenes narrowed toward the base and cuneately trun- 101 99 cate. It was described from Bornmiiller’s specimens collected on the border with Iran and published by Sientenius. Bornmiiller notes that his species may be compared with species of the genus Notobasis, but morphologically it is close to the Lebanese species C. phyllocephalum Boiss. C. bornmiilleri also differs somewhat from species of the sec- tion Epitrachys DC. by the structure of the pollen grains, which, ac- cording to I.S. Shtepa, have a thinner exine. Subsection 4. Armata Charadze in Addenda XXVII, 603.— Subsect. Eriophora Petrak in Bibl. Bot. 78 (1912) 6 p. p. (3. Horrida Petrak, 5. Lappacea Petrak) excl. 1. Eriocephala Petrak, 2. Ferocia Petrak, 4. Caucasica Petrak.—Subsect. Ciliata Petrak. 1. Serrulata Petrak, ibid. 5 p. p——Stem densely leafy. Leaves deeply pinnately incised, more or less coriaceous, with strongly spinescent lobes. Ca- pitula usually large, less often medium-sized, erect, surrounded by somewhat long apical leaves, solitary, in clusters of 2—3, or in panicu- late inflorescence. Involucre usually densely arachnoid-hairy or with thin arachnoid-pubescence; involucral bracts entire, smooth or sca- brous, less often with occasional spines on margin, usually narrowed from middle to erect or divergent long spines. Type of subsection: C. lJappaceum (MB.) MB. Note. The species of this subsection are widely distributed in Asia Minor, Iran, Afghanistan, southern Transcaucasia, Dagestan, and occa- sionally in the mountains of Soviet Central Asia. Petrak includes in the subsection Eriophora Petrak mainly the eastern European and Medi- terranean species of the affinity of C. eriophorum (L.) Scop. with the Caucasian-Western Asian series of species. However, it seems more natural to consider Eriophora Petrak to have two vicariant subsec- tions, i.e., subsect. Eriophora Petrak in the narrow sense and subsect. Armata Charadze. The members of both of these subsections are well differentiated by the structure of their capitula and involucral bracts. Series 1. Lappacea (Petrak) Charadze comb. nova.—Subsect. Eriophora Petrak 5. Lappacea Petrak in Bibl. Bot. 78 (1912) 6 p. max. p.—Capitula medium-sized, in more or less lax racemose or race- mose-paniculate inflorescence, less often clustered at tips of stems in racemose-capitate inflorescence. A row of narrowly-lanceolate, small, greenish bracteal leaves, to half as long as involucre usually develop- ing at base of involucre. Florets red or pink. 25. C. lappaceum (MB.) MB. FI. taur-cauc. III (1819) 558 p. p.; DC. Prodr. VI, 637 p. p.; Ldb. Fl. Ross. II, 729 p. p.; Boiss. Fl. or. II, 531, p. p. excl. syn.; Somm. and Lev. in Nuov. Giorn. Bot. Ital. N. S. II, 1, 12; Charadze in Fl. Gruzii, VIII, 500; in Fl. Azerb. VIII, 102 100 399.—C. lappaceum ssp. eulappeceum Petrak var. bibersteinii Petrak in Tr. Tifl. bot. sada, XII, 1 (1912) 10.—C. lappaceum var. biebersteinii Grossh. Fl. Kavk. VI (1934) 183.—C. penicillatum C. Koch in Linnaea, XVII (1843) 41.—Carduus lappaceus MB. Beschreib. Land. Casp. Meer. (1800) 193.—Cnicus lappaceus MB. F1. taur.-cauc. II (1808) 277, excl. syn. Carduus horridus Adams.—Epitrachys penicillata C. Koch in Linnaea, XXIV (1851) 399. Perennial. Stem 30—40 cm high, sulcate, finely whitish arachnoid- tomentose, ascending, simple or with few axillary branches above. Leaves densely spinose above, white-tomentose beneath on veins, more or less deeply pinnately incised; basal leaves oblong-elliptical, to 20 cm long, 7-8 cm wide, with short, toothed-winged, spiny petiole, al- most to base sinuately pinnately lobed into oblong-lanceolate segments; segments unequally 2-parted, slightly recurved, rarely spiny-ciliate, lobes apically strongly narrowed into spines 12—15 mm long; terminal segments elongated ternate with short lateral lobes; lower cauline leaves like basal, narrowed toward base, sessile, upper ones smaller, 7-13 cm long, ovate-lanceolate, sessile, semiamplexicaul, divergent from stem, with broad, yellowish, thick vein above, sinuately-pinnately parted into unequal 2-parted ovate-deltoid, long-spiny lobes, proximal lobules more or less completely reduced, spinescent. Capitula solitary, terminal on stem and usually on elongated axillary branches, 15-25 mm in dia, globose-ovate, surrounded by narrowly-lanceolate apical leaves, slightly or almost 2 times surpassing capitula, in racemose or corymbose-pan- iculate common inflorescence. Involucre subglabrous or sparsely floc- cose-arachnoid-hairy; involucral bracts at base of capitula leafy, green- ish, linear-lanceolate, spinescent, half as long as capitula; outer bracts ovate-lanceolate, narrowed from middle into recurved tip, thinly spinescent; inner bracts linear-lanceolate, spinescent with recurved tip. Corolla red, 22 mm long; limb 5-parted almost to middle, somewhat shorter than narrow tube. Pappus sordid-white, with thin fragile bristles, inner bristles with longer filiform tips, scabrous above; achenes 5.5 mm long, grayish-brown or almost reddish-brown. Flowering August to September. (Plate V, Fig. 2.) Mid-montane to subalpine zone, on herb slopes and among shrubs.—Caucasus: Eastern and Southern Transcaucasia. Endemic. Described from Iberia and Shirvan Mountains. Type in Leningrad. Note. The species is usually interpreted broadly by authors of the Western Asian flora. However, the forms similar to the type have a limited range, and apparently are not found beyond the Caucasus. The majority of the varieties of this species are distinct species close to C. lappaceum: also, the specimens from Turkey and Iran identified as C. lappaceum (MB.) MB. apparently do not belong to this species. 10 pS 101 26. C. anatolicum (Petrak) Grossh. Fl. Kavk. IV (1934) 182 p. max. p. excl. var. ferox (Boiss.) Grossh.; Charadze in Fl. Gruzii, VIII, 501; in Fl. Azerb. VIII, 400.—C. anatolicum var. microcephalum Grossh. and var. ramosum Grossh. Fl. Kavk. Iv (1934) 182.—C. lappaceum E. microcephalum Boiss. Fl. or. II (1875) 532 p. p—C. lappaceum var. lineariloba Trautv. in Tr. Peterb. bot. sada, IV, 1 (1876) 158.—C. lappaceum ssp. anatolicum Petrak in Tr. Tifl. Bot. Sada, XII, 1 (1912) 12 p. p.; in Nabelek, It. turc.-pers. II, 33, diagn. brev.—C. lappaceum ssp. anatolicum var. microcephalum Petrak in Tr. Tifl. bot. sada, XII, 1 (1912) 14 p. p.; in Izv. Kavk. Muz. VIII, 1- 2, 56.—C. lappaceum ssp. anatolicum var. ramosum Petrak in Tr. Tifl. bot. sada, XII, 1 (1912) 14; in Izv. Kavk. Muz. VIII, 1-2, 56. C. lanciflorum y. subnudum DC. Prodr. VI (1837) 638. Perennial. Stem 20—50 (80) cm high, ascending, weakly branched above, more or less sulcate, densely arachnoid-hairy, sparsely leafy. Leaves divergent from stem, usually very densely setose above with appressed, spiny bristles, midrib and lateral veins yellowish, distinct against background of pale green lamina, whitish-tomentose beneath, usually with raised subglabrous, thick midrib; all leaves pinnately in- cised almost to base into narrowly lanceolate or linearly-lanceolate acuminate segments, strongly spinescent with hard, yellow, spines 3— 9(12) mm long; segments 2-parted to base, apical lobe reduced, spinescent, recurved; terminal segments usually short-spinescent; basal and lower cauline leaves narrow, oblong, with winged petiole, usually 20 cm long, 5—6 cm wide, upper cauline leaves shorter, sessile, at base usually not broadened, deeply pinnately divided into more or less deltoid-lanceolate or ovate-lanceolate lobes; apical leaves 3—4, at base of capitula, linear-lanceolate, almost 2 times as long as capitula. Ca- pitula globose-ovoid, 15—35 mm in dia, terminal or axillary, subsessile, short-pedunculate or on long lateral branches, few, in racemose-pan- iculate common inflorescence. Involucre at base usually glabrous, sparsely arachnoid-hairy above, less often more densely; outermost involucral bracts narrowly lanceolate, transitional to small apical leaves; outer bracts ovate-lanceolate, glabrous, straw-yellow, weakly carinate, narrowed from middle in more or less falcately recurved, arachnoid hairy, spinescent tip; inner bracts linear-lanceolate, narrowed to fine recurved, spiny corolla. Corolla pink 16 mm long, limb scarcely longer than narrow tube, deeply 5-parted. Pappus yellowish, with fragile bristles, inner bristles narrowed to scabrous, filiform tips; achenes reddish-brown, 5—6 mm long. Flowering August to September. (Plate V, Fig. 3.) 102 Ne WY 4 iil i NY Ye A bs wy 5 N ae / salt NAN S\N S\ — 103 Plate V. 1—Cirsium aduncum Fisch. and Mey. ex DC.; 2—C. lappaceum (MB.) MB.; 3— C. anatolicum (Petrak) Grossh. 105 103 Dry mountain meadows, usually from mid-montane to subalpine zone.—Caucasus: Southern Transcaucasia General distribution: Asia Minor, Armenia and Kurdistan, Iran. Described from Turkish Arme- nia. Type in Vienna. Note. It differs from the other species of the series Lappacea by having the leaves deeply incised into very narrow segments and small solitary capitula on more or less elongate branches. Assessment of the taxonomic importance of the varieties of C. anatolicum (Petrak) Grossh. needs special study. 27. C. megricum Charadze in Addenda XX VII, 603; in Fl. Azerb. VIII (1961) 402, descr. ross.—C. lappaceum var. tomentosum auct. fl. cauc. p. p. non Boiss. Perennial. Stem 30-100 cm high, simple ascending, tomentose, densely leafy to tip. Leaves elongate-oblong, more or less densely spinose above, or spines very short and leaf surface scabrous and thinly arachnoid pubescent, white-tomentose beneath; lower cauline leaves pinnately incised into linearly lanceolate, acuminate segments, basally narrowed into more or less long toothed-winged petiole; leaf segments unequally 2-parted, lobes and teeth strongly spinescent with 10-15 mm-long, hard spines; upper cauline leaves sessile, semi- amplexicaul, deeply pinnately parted. Capitula surrounded by narrowly linear, greenish apical leaves, slightly surpassing them, terminal on stem and in axil of apical leaves, subsessile or on thick, densely whit- ish-arachnoid-hairy, short peduncles. Involucre ovate, subglabrous or arachnoid hairy, 1.2—2.0 cm in dia; lower involucral bracts at base of capitula greenish; others scarious, ovate to lanceolate, abruptly nar- rowed into acute, somewhat falcately recurved tip, spinescent, glabrous below, arachnoid-hairy above, weakly scabrous on margin. Florets pinkish; pappus yellowish-whitish; achenes dark colored. Flow- ering August to September. Meadows in subalpine zone.—Caucasus: Southern Transcaucasia (Zangelan, Megri). Described from vicinity of Megri. General distri- bution: Possibly grows in Iran. Type in Azerbaidzhan. Note. The species is close to C. anatolicum (Petrak) Grossh. in its pubescence and small capitula, but the latter are in terminal clusters on the stem as in C. schelkownikowii Petrak. It is well distinguished from the latter species by having not grayish but dense white-tomentose pubescence on the leaves and by narrow, petiolate lower cauline leaves. Specimens of this species have been identified as C. Jappaceum var. tomentosum (Boiss.) Petrak. 106 104 28. C. schelkownikowii Petrak in Izv. Kavk. Muz. VIII, 1-2 (1914) 44; Grossh. Fl. Kavk. IV, 182; Opred. Rast. Kavk. 483; Charadze in Fl. Azerb. VIII, 401; Papava in Vestn. Gos. Muz. Gruzii, XV A, diagn, repet.—C. laniflorum B. szowitsianum DC. Prodr. VI (1837) 638—C. lappaceum f. subracemosum Ldb. Fl. Ross. II (1845) 729.—C. lappaceum y. tomentosum Boiss. Fl. or. III (1875) 531 p. p. C. lappaceum var. tomentosum Petrak in Tr. Tifl. Bot. Sada, XII, 1 (1912) 10 p. p.; Grossh. Fl. Kavk. IV, 183 p. p.—Ic.: Petrak in Izv. Kavk. Muz. VIII, 1-2, Plate 2. Perennial. Stem thick, hard, simple, ascending, sulcate, more or less densely grayish-arachnoid-hairy, densely leafy, with terminal ca- pitula. Leaves usually recurved, more or less densely setose above with thin, spine-like bristles, with prominent veins broadened at base; cauline leaves broadened at base, auriculate, ovate-oblong, deeply- sinuate-pinnately parted with long spiny teeth, semiamplexicaul; leaf segments 2—3-parted with short upper teeth recurved; lower lobes large, lanceolate- or ovate-deltoid, obtuse or short-acuminate, on margin spiny- ciliate; all lobes strongly spinescent with thick, yellowish stiff spine 8—15 mm long, upper cauline leaves like lower, but somewhat shorter. Capitula terminal, in clusters of 3—9, in racemose inflorescence, sessile, ovate-globose, upper solitary capitulum 3.5 cm in dia and 3.6—4.0 cm long, apical leaves almost 2 times surpassing capitula. Capitula in axils of long upper leaves 2—5, smaller usually 2.5 cm in dia, 3 cm long, sessile or on short peduncles. Lowermost involucral bracts many, linear-lanceolate, undivided, spiny-ciliate on margin, somewhat shorter or almost as long as involucre; outer involucral bracts glabrous below, densely grayish-arachnoid-lanate above, ovate at base, from middle abruptly narrowed into linearly lanceolate, carinate apex, tapered into more or less erect spine; inner bracts linear-lanceolate, long-attenuate into yellowish spine 2—4 mm long, horizontal or somewhat falcately recurved. Corolla purple; limb 5-parted to middle, almost equaling narrow tube or somewhat shorter. Filaments denticulate-hairy. Pappus sordid white with unequal bristles, inner bristles almost 2 times as long as outer, lanate over 2/3, filiform at tip, toothed; achenes imma- ture. Flowering July to September. Subalpine and alpine zones.—Caucasus: Southern Transcaucasia (Karabakh, Nakhichevan ASSR), Talysh. General distribution: Tran and Afghanistan. Described from vicinity of nomadic settlement of Ala-Khanchala in Karabakh. Type in Tbilisi; isotype in Leningrad. Note. It is distinguished from C. Jappaceum (MB.) MB. by clus- tered, subsessile, grayish-lanate capitula. The specimen of Sovich from Karabakh cited by De Candolle as C. laniflorum B. szowitsianum DC. and later included by Boissier under C. /appaceum y. tomentosum 107 105 Boiss. belongs to this species. The specimens of Sovich from Karabakh are cited by Ledebour under the name C. lappaceum B. subracemosum DC. They are distinguished from the type by somewhat smaller ca- pitula and acuminate leaf lobes. The Lenkoran specimens are distin- guished also by a more xerophilous habit. This species is distinguished from C. macrobotrys (C. Koch) Boiss. by relatively small capitula and pubescence. C. schelkownikowii Petrak is reported from Afghanistan (Petrak in Kgje and Rechinger, Symb. Afgh. II [1955] 178; Kitamura, Fl. Afgh. II [1960] 408). The range of this species is not yet suffi- ciently clear. Possibly the Afghan specimens are closer to C. pseudolappaceum Charadze, which was described from Kopetdag. 29. C. macrobotrys (C. Koch) Boiss. FI. or. III (1875) 532; Petrak in Tr. Tifl. bot. sada, XII, 1, 19; in Izv. Kavk. Muz. VIII, 1-2, 54; in Nabelek, It. Turc.-pers. II, 33; Grossh. Fl. Kavk. IV, 182; Opred. rast. Kavk. 483 (sub. C. macroptero Sphalm. [sic.]); Charadze in Fl. Azerb. VII, 401.—C. macrobotrys var. longilobum Bordz. in schedia, quoad. pl. e. Sarykamysch.—C. lappaceum y. macrocephalum Ldb. F1. Ross. II (1846) 729.—Epitrachys macrobotrys C. Koch in Linnaea, XXIV (1851) 399. Perennial. Stem 40-150 cm high, robust, ascending, more or less densely, grayish-arachnoid-hairy, sulcate, simple, with large solitary, terminal and axillary capitula in short racemose common inflorescence. Leaves green above with midrib straw-yellow, broadened at base, and prominent lateral veins, setose with appressed or slightly divergent, spiny bristles and with occasional flexuous hairs; sparsely grayish- tomentose beneath; cauline leaves oblong, middle ones 10—15 cm long, sessile, auriculate, weakly decurrent, remotely pinnately parted into deltoid-ovate or deltoid-lanceolate, unequally two-parted lobes, with short upper recurved lobe; terminal segment linearly lanceolate; all lobes and teeth more or less strongly spinescent, with 6-12 mm long, hard spines; apical leaves in inflorescence almost linearly lanceolate, with strongly reduced segments and short spines, surpassing capitula. Capitula (2.5)3.0—3.5(5.5) cm in dia, subsessile, in axils of apical leaves, less often lateral, on short peduncles. Involucre weakly floccose-lanate or subglabrous below, densely arachnoid-hairy above; lower involu- cral bracts many, narrowly linear, greenish-brown, spinescent, usually more than 1/2 as long as involucre, rarely floccose below, long arach- noid hairy above; outer bracts glabrous, carinate; oblong-ovate, sca- brous above on margin, abruptly narrowed from middle into falcately recurved, narrowly linear spiny apex with long floccose hairs; inner bracts glabrous, linearly lanceolate, falcately recurved. Corolla pink, 23 mm long; corolla tube narrow, gradually transitional to broad limb; 108 106 limb lobed up to 1/3. Pappus yellowish, to 20 mm long, inner hairs with somewhat elongate, scabrous tips; achenes 6 mm long, 3 mm wide, brownish. Flowering August to September. Subalpine zone, in meadows.—Caucasus: Eastern Transcaucasia (between Shakhdag and Khinalug peaks), Dagestan (Kurush), Talysh. General distribution: Asia Minor (Geksu, Khinish and Abusardag), Armenia and Kurdistan (Kars, Sarykamysh, Verakdar near Van, be- tween Bashkale and Kochan). Described from Asia Minor. Type in Berlin. Note. It is distinguished from C. schelkownikowii Petrak by the considerably larger capitula, form of the leaves, and nature of the pubescence. The range of this species is not entirely clear and it all needs refining. Series 2. Cosmeliana Charadze.—Subsect. Eriophora Petrak 5. Lappacea Petrak in Bibl. Bot. 78 (1912) 6 p. min. p.—Florets yellow. Capitula medium-sized, densely arachnoid-hairy, solitary, terminal on stem and branches, sessile; apical leaves considerably surpassing ca- pitula. Leaves pinnate, with approximate, linear, deeply two-parted segments. 30. C. cosmelii (Adams) Fisch. ex Hohen. in Bull. Soc. Nat. Mosc. VI (1833) 251; Petrak in Tr. Tifl. bot. sada, XII, 1, 6; Ldb. Fl. Ross. II, 730; Trautv. in Tr. Peterb. bot. sada, IV, 1, 158; Grossh. Fl. Kavk. IV, 182; Opred. Rast. Kavk. 483; Charadze in F1. Gruzii, VIII, 502; in Fl. Azerb. VII, 400.—C. lappaceum MB. F1. taur.-cauc. III (1819) 558 p. p.; Boiss. Fl. or. II, 531 p. p—C. lokense Conr. and Fr. in Bull. Herb. Boiss. III (1895) 468.—Carduus cosmelii Adams in Web. and Mohr. Beitr. Naturk. I (1805) 66.—Exs.: Pl. or. exs. No. 97; Herb. fl. cauc. No. 197; Petrak, Cirsioth. Univ. No. 95. Perennial. Stem strong, ascending, (15)40—60 cm high, somewhat flexuous, spreadingly branched from middle, indistinctly angular, sul- cate, purple, arachnoid-hairy, to tip densely leafy. Leaves setose above with spiny bristles, grayish-tomentose beneath with prominent, weakly arachnoid, whitish midrib, linear or lanceolate-elliptical, sinuately-pin- nately incised almost to base, with approximate, elongated segments; segments 2—5-parted, with 2 linearly-lanceolate, almost equal lobes; anterior lobe usually recurved with deltoid-lanceolate teeth, strongly spinescent with yellowish spines 5-7 mm long, on margin spiny-ciliate and somewhat recurved; all cauline leaves narrowed toward base to winged-toothed, long-spiny petiole; petioles long in lower and short in upper leaves; terminal segments linearly-lanceolate, strongly spinescent; cauline leaves somewhat shorter above, on the average 15—20 cm long, 109 107 4—5 cm wide; apical leaves many, linear or linearly lanceolate, some- what exceeding capitula, narrowed toward base, acute, with short lat- eral lobes, spinescent. Capitula globose-ovate, solitary, terminal, less often 2 on stem and long axillary branches, in paniculate common inflorescence. Involucre 1.5—3.5 cm in dia, thinly arachnoid-pubes- cent; outer involucral bracts ovate-lanceolate, inner linearly lanceolate abruptly narrowed, spinescent with slightly recurved, long spine; lower most bracts few, narrowly linear-lanceolate, arachnoid-hairy almost from base, outer and inner ones glabrous, lustrous, scarious, arach- noid-hairy at base of spines. Florets yellow, 25 mm long, with narrow, tube to 15 mm long; limbs 5-parted. Pappus yellowish-white, to 20 mm long, inner hairs with long scabrous tips; achenes 6.5 mm long, up to 3 mm wide, light reddish-brown with dark brown striations. Flow- ering July to September. Mid-montane to subalpine zone. Along roads in forest zone and in mountain meadows. Common weed of mountain pastures, particularly in Southern Transcaucasia.—Caucdsus: Eastern, Southern and West- ern Transcaucasia (reported from Adzharia). General distribution: Asia Minor. Described from region between Tbilisi and Dusheti. Type in Leningrad. Note. Hybrids are known. 1. C. x trifurcum (C. cosmelii x C. caucasicum) Petrak in Izv. Kavk. Muz. VIII, 1-2 (1914) 59.—Exs.: Petrak, Cirsioth, Univ. No. 140. Described from Bakuriani, where it was collected among related forms. Petrak (1. c.) reports that this hybrid is of special interest because hybrids are very rare among species of the section Epitrachys DC. in contrast to the members of the section Onotrophe (=Cirsium). The pollen studies conducted by I.S. Shtepa showed that almost 70% of the pollen is not fully viable. C. cosmelii is widespread in Southern Transcaucasia, whereas the southern boundary of C. caucasicum runs through the Trialeti Range, where hybrids are observed. 2. C. cosmelii x C. ciliatum (=C. ciliatiforme Petrak): Grossh. FI. Kavk. IV (1934) 189. Caucasus: Reported from Southern Transcaucasia (former Akhalkalak District). In the structure of the capitula, particularly the involucral bracts, it is more similar to C. szowitsii (C. Koch) Boiss. than to C. ciliatum (Murr.) Moench; it approaches C. cosmelii in its leaves with the proxi- mate segments cut almost to the base into slightly unequal, narrow, oblong lobes. 110 108 3. C. x succinctum Ldb. FI. Ross. II (1846) 732. Stem ascending, sparsely pilose with crispate hairs, with dark purple spots, sulcate, branched above. Lower cauline leaves oblong, to 25 cm long, sparsely spiny-setose above, grayish-tomentose beneath, on mid- rib subglabrous, narrowed into petiole, deeply pinnately incised into oblong segments; lateral segments unevenly 2—3-parted, with anterior short ovate-deltoid lobes; all lobes strongly spinescent with up to 7 mm-long, hard spines; other leaves gradually reduced upward, sessile, semiamplexicaul; apical leaves divergent, surpassing capitula. Capitula 3-4 cm in dia, solitary, terminal on stem and axillary branches, in corymbose-paniculate common inflorescence. Involucre covered with thin arachnoid pubescence; involucral bracts straw-yellow, sometimes dark-colored, carinate, outer and middle ovate to oblong-ovate, nar- rowed above middle into long, herbaceous, recurved tip, spinescent; inner bracts linearly lanceolate, with thin, recurved spiny tip. Florets red. Caucasus: Eastern Transcaucasia (Sarial). Described from Sarial Mountains in Azerbaidzhan from Hohenacker’s specimens. Noted on the labels of C. succinctum is: “... together with C. tricholoma Fisch. and Mey.” In the form of the leaves it is almost indistinguishable from C. tricholoma. Authors of the Caucasian flora have considered it a synonym of C. lappaceum (MB.) MB., which it approaches in its erect capitula and form of the involucral bracts, while differing by having large capitula and an arachnoid-hairy involucre. In having apical leaves that surpass the capitula, it is similar to C. cosmelii. Possibly, it is a hybrid between C. cosmelii and C. tricholoma. Series 3. Arachnoidea Charadze.—Subsect. Ciliata Petrak 1. Serrulata Petrak in Bibl. Bot. 78 (1912) 5 p. min. p.—Capitula basally slightly cuneate, solitary or 2—3 on short peduncles. Involucral bracts usually scabrous on margin, smooth, less often with occasional spiny setae. Basal and lower cauline leaves pinnately incised, upper pin- nately parted or pinnately lobed. The species of this series are distributed in the Caucasus and Crimea. They differ from species of the series Lappacea by the shape of the capitulum that is usually projected above the apical leaves. 31. C. arachnoideum (MB.) MB. FI. taur.-cauc. III (1819) 657; DC. Prodr. VI, 639; Ldb. Fl. Ross. II, 732; Boiss. Fl. or. II, 536 p. p. excl. syn.; Lipsky, Fl. Kavk. 358, excl. var. B.; Grossh. Fl. Kavk. IV, 183, excl. var.; Opred. Rast. Kavk. 484.—Cnicuus arachnoideus MB. FI. taur.-cauc. II (1808) 277.—Ic.: Rchb. Pl. crit. IV, tab. 359. 111 109 Perennial. Rhizome thick, woody, producing 1—3 erect stems; plant 70-80 cm high. Stem yellowish or reddish, weakly angular, sparsely crisped hairy, simple or weakly branched above, leafy to tip. Leaves densely spinulose above, with appressed spinules, subglabrous beneath, along veins weakly arachnoid hairy, less often thinly arachnoid. Basal and lower cauline leaves with long petioles, semiamplexicaul; lower leaves including petioles 21—25(37) cm long, 9—10(18) cm wide, ellip- tical-elongate, pinnately divided almost to base, basal segments broad, with elongated broadly lanceolate terminal lobe and deltoid-ovate lat- eral lobes; middle and upper cauline leaves auriculately broadened, semiamplexicaul, middle ones to 12 cm long and 5.5 cm wide, nar- rowed at apex, lanceolate-elliptical to ovate, usually deeply sinuate- pinnately divided into 2-parted elongated, lanceolate or deltoid-ovate lobes, spinescent, on margin finely spiny-ciliate; apical leaves small, approximate to capitulum, usually deflexed. Capitulum ovate, termi- nal, somewhat projected above leaves, solitary, at apices of stem and branches, on short peduncles, sub-sessile, aggregated in compact, corymbose paniculate inflorescence. Involucre 15—20(25) mm in dia, 18-20 mm long, basally cuneate at beginning of flowering, subapically attenuate, often purple, thinly arachnoid-hairy; outer bracts narrow, lanceolate, carinate, gradually narrowed, spinescent with thin purple spines, sometimes with occasional spines on margin; outer bracts usu- ally 3-4 mm long, gradually longer inward; inner ones membranous, linearly lanceolate, with incurved fine tip. Florets purple, to 26 mm, in broader part to 14 mm long, limb cut up to middle. Pappus basally brownish, sordid-white above, inner hairs with scabrous tips; achenes to 5.5 mm long, silver-gray, with dark streaks. Flowering July to August. Commonly in limestone regions. Mid-montane to subalpine zone, in meadows, along forest edges, in crops; widely distributed as com- ponent of weedy tall-herb vegetation Caucasus: Ciscaucasia (Cen- tral and Eastern parts), Dagestan. Described from Beshtau. Endemic. Type in Leningrad. Note. It is distinguished from the other species of the series by having very small outer involucral bracts and weakly pubescent leaves. 32. C. euxinum Charadze in Zam. po Sist. i Geogr. Rast. Tbil. 23 (1963) 108.—C. laniflorum Grossh. Opred. Rast. Kavk. (1949) 482, non MB.—C. arachnoideum var. incanum Lipsky in Schmalh. FI. II (1897) 102; Lipsky, Fl. Kavk. 358; Grossh. Fl. Kavk. IV, 183; Kolak. Fl. Abkhazii, IV, 271; Charadze in Fl. Gruzii, VIII, 508. Perennial. Stem to 70 cm high, weakly angular, ribbed, floccose- crisped-hairy or almost white-tomentose-hairy, erect, more or less 112 110 branched above. Leaves thinly spinose above, more or less densely white-tomentose beneath, with prominent midrib; basal leaves oblong- ovate, up to 25 cm long, 16 cm wide, deeply sinuate-pinnately divided into broad approximate segments, in turn apically bifid into deltoid lobes, ciliolate, spinose on margin with obtuse teeth and segments spinescent bearing spines to 3 mm long, narrowly decurrent on petiole as wing, petiole almost as long as lamina; cauline leaves oblong, nar- rowed at both ends, sessile, semiamplexicaul, sinuate-pinnately parted to middle or deeper into elongate, unequally bifid, ovate-oblong seg- ments, apically toothed, teeth spinescent; middle cauline leaves to 20 cm long, to 11 cm wide, upper ones 6 cm long and to 4 cm wide. Capitula numerous, terminal and axillary, ovate-globose, basally weakly cuneate, 1.5—2.0(2.5) cm in dia, with arachnoid-hairy involucre. In- volucral bracts imbricate, outer ones smaller, lanceolate, narrowed into thin, deflexed purple spine, glabrous, scabrous on margin, weakly arach- noid-hairy above, inner ones linearly lanceolate, tapered into fine cusp. Florets red, to 27 mm long, in expanded part to 13 mm. Pappus sordid- white, basally yellow, with inner hairs apically narrowed into slightly clavate, scabrous tip; achenes 5 mm long, 2.5 mm wide, brown with short dark streaks. Flowering July to August. Limestone cliffs.—Caucasus: Western Transcaucasia (Anapa, vi- cinity of Novorossiisk, Gelendzhik, Gagra). Endemic. Described from vicinity of Novorossiisk. Type in Leningrad. Note. This species is usually treated as a variety of C. arachnoideum (MB.) MB. with which, however, it has very little in common. It is closer to the Crimean forms of C. laniflorum (MB.) MB., from which it is distinguished by globose capitula and smaller involucral bracts. In the east, the range of C. euxinum extends to the Gagra District. This species is one of the indicators of the Crimean- Western Caucasian floristic links. 33. C. laniflorum (MB.) MB. FI. taur.-cauc. III (1819) 557; DC. Prodr. VI, 638, excl. var.; Ldb. Fl. Ross. II, 731 p. p.; Charadze in Zam. po Sist. i Geogr. Rast. Tbil. 23, 106.—Cnicus laniflorus MB. F1. taur.-cauc. II (1808) 276. Perennial. Stem ascending, usually to 50 cm high, weakly angular, sulcate, pale purple, more of less thinly arachnoid-hairy, below inflo- rescence almost white-tomentose, with deflexed leaves, weakly branched above. Leaves deeply sinuate-pinnately parted, densely spinose with slightly appressed spines, sometimes very finely arachnoid-hairy, white- tomentose beneath, with ligneous midrib; lower cauline leaves with long spinose petiole; middle cauline leaves narrowed into more or less short, toothed and long spinose along margin, amplexicaul petiole, 113 111 elongated-elliptical, 25 cm long, 13 cm wide, incised almost up to base into linearly lanceolate unequally 2-parted segments, with 1-2 lateral deltoid-ovate teeth, leaf lobes narrowed into spines 2-4 mm long; upper cauline leaves sessile, semiamplexicaul, to 12 cm long, pinnately parted into ovate or ovate-oblong segments, tapered into spines usually upto 8 mm long. Capitula terminal, usually solitary, on short peduncles, subsessile, slightly elongate, with florets 2.5—3.5 cm long, 2.0—2.5 cm in dia, wide open at end of flowering, to 3 cm in dia, clustered in 2-3 or solitary, on short, axillary and terminal peduncles, forming elongated racemose-corymbose inflorescence. Involucre floc- cose-arachnoid-hairy; involucral bracts narrowly lanceolate, spinose along margin, almost abruptly tapered into long, thin, hard, more or less squarrose spine. Corolla pink, 23-24 mm long, broader part more or less longer than narrow tube, cut upto 1/3, geniculately bent in outer florets. Pappus sordid-white, basally yellowish, to 22 mm long, more or less dense; inner pappus hairs slightly longer than outer, apically slightly thickened, scabrous; achenes to 5 mm long, 2.5 mm wide, yellowish, with blackish-purple streaks. Flowering August to September. Herb slopes, along roads in pine and juniper forests, occasionally in Yaila [mountain pastures]_—European Part: Crimea. Endemic. De- scribed from Crimea. Type in Leningrad. Note. It differs from the species of the series Arachnoidea and Lappacea by the long squarrose spines of involucral bracts and is sharply differentiated by the degree of incision of the lower and upper cauline leaves. 34. C. sublaniflorum Sojak in Nov. bot. Hort. bot. Univ. Carol. Prag. (1961) 36.—C. fimbriatum ssp. bornmiilleri Petrak in Oest. bot. Zeitschr. X (1910) 395.—C. serrulato-lanceolatum Boiss. Fl. or. II (1875) 553 p. p.; Zelenetzky in Zap. Novoross. Univ. 102, 351 p. p. Perennial. Stem ascending, up to 80 cm high, sulcate, more of less purple, arachnoid-hairy, branched from middle, densely leafy below, sparsely above. Leaves with appressed spines above, more or less loosely white-tomentose-beneath; lower cauline leaves oblong-ellipti- cal, more or less deeply sinuate-pinnately divided into ovate-oblong or oblong lobed, basally lobes with coarse deltoid-lanceolate teeth, spi- nose-ciliate along margin strongly tapered to yellow; 2-6 mm-long, hard spines; upper leaves gradually reduced toward apex, oblong or lanceolate-oblong, shallowly pinnately parted, sessile basally auriculately expanded, semiamplexicaul; apical leaves 1—2 at base of capitula, small lanceolate, usually progeosthetic, pinnately-toothed, with long-spinescent teeth. Capitula solitary, terminal and axillary, ovate or 114 W2 globose-ovate, 2—3.5(5) cm in dia. Involucre densely floccose-arach- noid from slightly yellowish hairs; outer involucral bracts glabrous, tapered to greenish tip, more or less recurved, tapered into spine 3—5 mm long. Corolla 26—28 mm long, in broader part incised to middle, usually 13 mm long. Pappus sordid white, inner hairs with longer scabrous tip; achenes 5 mm long, brownish. Flowering July to August. Among pear forests, in glades, in pine forests, on northern slopes of Yaila—European Part: Crimea (Karadag, Crimean State Reserve, Massandra, Lesser and Greater Chuchel, Mangup-Kale, Salamlar, Simferopol and others). Endemic. Described from Crimea (Karadag). Type in Leningrad. Note. Petrak, who described this plant as C. fimbriatum ssp. bornmiilleri Petrak, compared it with C. fimbriatum ssp. tricholoma (Fisch. and Mey.) Petrak. The latter is a separate species and belongs to the series of closely related species grouped around C. osseticum (Adams) Petrak, for which nodding capitula are characteristic. C. sublaniflorum Sojak with its erect capitula and structure of the leaves is closer to the Crimean species C. laniflorum, differing from it by having a broad, less incised lamina, a densely arachnoid-hairy involu- cre and larger capitula; it also differs by having involucral bracts that terminate in an herbaceous, deflexed tip, with a more or less short small spine. Possibly, specimens of this species were the ones cited by Boissier as hybrid forms between C. serrulatum and C. lanceolatum, under the name C. serrulato-lanceolatum. Zelenetzky supported this view. The ecological conditions and the distribution of this species still need to be investigated. Sojak segregated the specimen cited by Petrak as C. fimbriatum ssp. bornmiilleri Petrak from Karadag as C. sublaniflorum Sojak. Sojak considers closely related forms that differ from this specimen by hav- ing attenuated spinescent involucral bracts with occasional spines on the margin to be C. laniflorum (MB.) MB. Specimens with entire in- volucral bracts were described by him under the name C. tauricum Sojak. I attach a broader meaning to C. sublaniflorum Sojak, by in- cluding under it all specimens with involucral bracts having an attenu- ated herbaceous tip with a more or less short spine. Moreover, the spininess of the margin of the lower involucral bracts varies and some- times is almost unexpressed. The typical form of C. sublaniflorum (MB.) MB. should be considered the widespread one in southern Crimea; its characteristic features are the form of the involucral bracts, with strongly elongate, yellowish, squarrose tips, a narrower, slightly elongate capitulum, and deeply pinnatifid leaves. Identical specimens are pre-served in the Marschall-Bieberstein herbarium in Leningrad among Steven’s collections. UNS) 113 35. C. tauricum Sojak in Nov. bot. Hort. bot. Univ. Carol. Prag. (1961) 35. Perennial. Stem erect, 60-80 cm high, sulcate, arachnoid-hairy, more or less snow-white, weakly branched above. Cauline leaves co- riaceous, oblong-elliptical, sessile, semiamplexicaul, not decurrent, distant and sinuately pinnately incised; leaf lobes oblong, unequally 2- parted, short-tapered to spines 3—4 mm long; lamina thinly spinose- ciliate, finely setose above, more or less densely snow-white arach- noid-hairy beneath; leaves gradually reduced above, sinuate-toothed. Capitula 4, without or with 1 bracteal leaf somewhat distant from capitulum, broad, spinose along margin, reduced; capitulum ovate-glo- bose, erect, 2.5—3.0 cm in dia. Involucre densely grayish-arachnoid- hairy; outer and middle involucral bracts lanceolate, entire, with small papillae (under a hand lens), more or less abruptly narrowed from middle, narrow part 5-7 mm long, with short yellow stramineous spine, more or less erect or somewhat deflexed, never arcuate; spines of inner involucral bracts longer and deflexed. Corolla purple, to 30 mm long, about 12.5 mm long in broader part. Pappus to 18 mm long, sordid-white; mature achenes unknown. Flowering August. Montane herb slopes.—European Part: Crimea (Yaila, Ai-Petri). Described from Crimea. Type in Prague. Note. It is a small montane race of the C. laniflorum (MB.) MB. alliance With its numerous involucral bracts with thin spines it approaches the Caucasian species C. euxinum Charadze. Series 5. Pugnacia Charadze.—Subsect. Eriophora Petrak 3. Horrida Petrak in Bibl. Bot. 78 (1912) 6 p. p.—Involucral bracts almost subulately narrowed into long, erect spines; capitula more or less large, surrounded by apical leaves. Leaves deeply pinnately incised into ovate or oblong segments, on both sides or only beneath whitish- arachnoid-hairy or white-tomentose, less often subglabrous. It includes mainly Caucasian species found in the Greater Caucasus region. One species.—C. cephalotes Boiss.—is of the Caucasus and Asia Minor. 36. C. cephalotes Boiss. Diagn. pl. or. ser. I, 6 (1845) 102; FI. or. III, 533; Somm. and Lev. in Nuov. Giorn. Bot. Ital. N.S. II, 1, 12; Charadze in Fl. Gruzii, VIII, 499; in Zam. po Sist. i Geogr. Rast. Tbil. 20, 46.—C. Horridum DC. Prodr. VI (1837) 638, quoad pl. ex Erzerum, non MB. (sec. Boissier).—C. horridum var. tomentosum auct. non Boiss.: Petrak in Tr. Tifl. Bot. Sada, XII, 1 (1912) 33; in Vestn. Tifl. Bot. Sada, XII, 1(1912) 33; in Vestn. Tifl. Bot. Sada, Vyp. 24, 3.— C. horridum ssp. tomentosum Petrak Izv. Kavk. Muz. VIII, 1-2 (1914) 116 114 48 p. p. quoad pl. ex Adsharia—C. tomentosum Grossh. Fl. Kavk. IV (1934) 181 p. p.; Opred. Rast. Kavk. 483 p. p. non C.A.M.—C. munitum Somm. and Lev. in Nuov. Giorn. Bot. Ital. N.S. II, 1 (1895) 12 p. p.; in Tr. Petersb. Bot. Sada, XVI (1990) 255 p. p.; non MB.—C. munitum var. stenophyllum Somm. and Lev. op. cit. (1895) 12; op. cit. (1900) 256. Perennial. Stem basally surrounded by dark brown petioles of old leaves, thick, ascending, sulcate, more or less grayish-arachnoid-hairy, simple or with 1—2 axillary branches above, to tip leafy, (15)20-—60 cm high. Leaves grayish above from more or less sparse arachnoid pubes- cence and sparse, short, spiny bristles, grayish-arachnoid-hairy to white- tomentose beneath, deeply pinnately divided into broad, ovate or round- ish-ovate, less often oblong, 2-parted almost to middle, lobes with 3—5 teeth, tapered to hard spines 4—5 mm long, with sparse squarrose spines along margin; basal leaves to 25(30) cm long, 8(12) cm wide, elon- gate-elliptical, narrowed into elongate spinose-toothed petiole, deeply pinnately divided, with approximate segments; cauline leaves sessile, semiamplexicaul, linear-oblong or linear-lanceolate, pinnately cut into roundish-ovate segments with obovate sinuses between them; apical leaves at base of capitula numerous, broad, oblong, more or less pin- nately parted, as long as or longer than capitula. Capitula solitary, terminal on stem and branches, subsessile in axils of apical, reduced leaves, 3.5-5.5 cm long, 3—5 cm in dia. Involucre densely floccose, grayish-arachnoid-hairy; involucral bracts elongate, outer ovate-lan- ceolate, inner lanceolate-linear, gradually narrowed into elongate, erect spines, glabrous beneath, carinate, blackish-brown above, almost to tip grayish-arachnoid-hairy. Florets purple, 30-35 mm long; corolla with long, narrow tube, abruptly broadening to limb 11—13 mm long. Pap- pus brownish-yellowish with longer, almost filiform, apically scabrous inner hairs; achenes brown, 5 mm long. Flowering July to September. Rubbly and stony places, in subalpine and lower alpine zones.— Caucasus: Western Transcaucasia (mostly mountains of Adzharia and Guria, occasionally in Upper Svanetia). General distribution: Asia Minor, Armenia and Kurdistan. Described from Erzerum. Type in Geneva. Note. The species is variable in the form of the leaves and the pubescence and spininess of the upper side of the lamina. Possibly it forms hybrids with forms of C. pugnax Somm. and Lev., which it adjoins along the northern boundary of its range in the Greater Caucasus region. Authors of the Caucasian flora usually refer specimens of C. cephalotes Boiss. to C. tomentosum C.A.M., from which they are well distinguished by their large, usually solitary capitula and the form of the leaves. Probably, Sommier and Levier referred this same species to 117 BES C. munitum (MB.) MB., describing it as C. munitum var. stenophyllum Somm. and Lev. 37. C. pugnax Somm. and Lev. in Nuov. Giorn. Bot. Ital. N.S. II, 1 (1895) 12; in Tr. Peterb. Bot. Sada, XVI (1900) 256.—C. pugnax var. araneosum Somm. and Lev. 1. c. (1900) 257.—C. horridum (Adams) Petrak in Tr. Tifl. Bot. Sada, XII, 1 (1912) 19, non MB. (1819).—C. horridum auct. non MB.: Charadze in Fl. Gruzii, VIII (1952) 497, descr. georg.; in Zam. po Sist. 1 Geogr. Rast. Tbil. 20, 47; in Fl. Azerb. VIII, 397.—C. horridum ssp. macrocephalum Petrak in Izv. Kavk. Muz. VIII, 1-2 (1914) 46 p. p. non Cirsium macrocephalum C.A.M.—C. macrocephalum Grossh. Fl. Kavk. IV (1934) 181 p. p.; Opred. Rast. Kavk. 483 p. p. non C.A.M.—C. munitum MB. FI. taur.- cauc. III (1819) 559; DC. Prodr. VI, 639; Ldb. Fl. Ross. II, 731; Boiss. FI. or. III, 533, excl. var. B. and y.—C. munitum var. hypopolia Trautv. in Tr. Peterb. Bot. Sada, V, 2 (1878) 448.—Carduus horridus Adams in Web. and Mohr, Beitr. Naturk. I (1805) 66.—Cnicus munitus MB. FI. taur.-cauc. II (1808) 279. Perennial. Plant with somewhat ascending, thick stems, 25—50 cm high. Stem sulcate, more or less sparsely arachnoid-hairy, weakly branched above, leafy to tip. Leaves stiff, oblong-broadly lanceolate, upper ones oblong, more or less densely setose above with short spiny bristles and glabrous or sparsely arachnoid-hairy, glabrous beneath, on main veins arachnoid-hairy or entirely, more or less densely, grayish- arachnoid-hairy, usually deeply pinnately divided or in lower leaves shallowly parted into unequal 2—3-parted or 2—3-toothed broadly ovate segments, with lobes apically deltoid, pointed, tapered to hard, more or less thick, yellowish spines, 6-10 mm long, sparsely short spiny along margin; basal and lower cauline leaves basally narrowed into long spiny toothed petiole, acute; middle and upper leaves basally weakly narrowed, semiamplexicaul, middle leaves to 15 cm long, 5 cm wide. Capitula usually 3-6 cm in dia, subsessile, terminal on stem and branches, usually 1—4, surrounded by apical leaves, more or less pro- jected above them. Involucre densely whitish-arachnoid-hairy; outer involucral bracts ovate-oblong, glabrous below, lustrous, arachnoid- hairy above, weakly carinate, tapered to long, straight or slightly bent spines; outermost bracts abruptly narrowed from broad base to long arachnoid-hairy spines, inner ones linearly lanceolate, gradually nar- rowed into thin, curved, elongate spine. Florets purple-red, 25-35 mm long; corolla with long narrow tube, abruptly transitional to broad, 13 mm long, to 1/3 cut up limb. Pappus brownish, with scarcely thick- ened, elongate tips of inner hairs; achenes 5.5 mm long, dark-brown- ish-reddish-brown. Flowering July to September. 118 116 Mid-montane to alpine zone, in meadows and pastures, as weed, and along roads.—Caucasus: Ciscaucasia, Dagestan, Eastern Transcaucasia, Western Transcaucasia (northern part). Described from Dout Pass. Endemic. Type in Florence. Note. The species is highly polymorphic. The size of the capitu- lum is variable, as is the pubescence of the leaves (from more or less glabrous to grayish-arachnoid-hairy), etc. It is possible that C. pugnax forms hybrids at the boundary of its range with C. tomentosum C.A.M. in the Eastern Caucasus and with C. cephalotes Boiss. in the Western Caucasus. This species has been described under different names, first by Adams as Carduus horridus Adams, then by Bieberstein as Cnicus munitus MB. In all probability Sommier and Levier redescribed this same species aS C. pugnax Somm. and Lev. At the same time, they cited specimens of C. cephalotes Boiss. from Western Transcaucasia under the name C. munitum (MB.) MB. The first combination, C. horridum (MB.) MB. (basionym Cnicus horridus MB.), was used by Bieberstein for C. caucasicum (Adams) Petrak and is an earlier homonym of C. horridum (Adams) Petrak. For this reason the latter name, based on the basionym Carduus horridus Adams, must be rejected. The next name, in chronological order, of this species—Cirsium munitum (MB.) MB. (basionym Cnicus munitus MB.)—must also be rejected, as having been given to the same speci- men from the Kazbek District. This leaves C. pugnax Somm. and Lev. as the valid name. This species was described from several specimens from the basin of the Teberda River and from the Kazbek District. Sommier and Levier consider the form with glabrous leaves from the Dout Pass as the type of this species. Among the pubescent forms (var. araneosum) Sommier and Levier cite a specimen from Daryal, from where Carduus horridus Adams and Cnicus munitus MB. also were described. 38. C. balkharicum Charadze in Zam. po Sist. 1 Geogr. Rast. Tbil. 22 (1961) 69. Perennial. Whole plant white-arachnoid-hairy, producing several stems. Stems strong, erect, usually to 75 cm high sulcate, branched above, purple, weakly white-arachnoid-hairy, leafy to tip. Leaves large, deflected from stem, greenish above, scabrous from tiny, almost in- conspicuous, appressed spines, arachnoid-hairy, white tomentose be- neath, only along midrib thinly arachnoid-hairy, oblong-ovate or ellip- tical, broad, shallow or deeply sinuate-pinnately parted into unequal 2—3-parted, ovate segments, lobes broadly deltoid tapered to hard, 5— 10 mm long, yellowish spines; basal and lower caulines leaves to 25 cm long, 11 cm wide, narrowed at both ends, other cauline leaves 117 sessile, semiamplexicaul, middle ones 15—20 cm long, 5—8 cm wide; apical leaves at base of capitula strongly reduced, shorter than or very slightly surpassing capitula. Capitula roundish-ovoid, (4)5—6(7) cm in dia, solitary, terminal on stems and some axillary branches. Involucre densely whitish-arachnoid-hairy; lower involucral bracts small, abruptly narrowed into long arachnoid-hairy spines, outer ones ovate-lanceolate, scarious, greenish-yellowish, weakly carinate, glabrous, along margin floccose-hairy, abruptly narrowed into long, thin, basally slightly curved, arachnoid-hairy spines; inner bracts linearly lanceolate, glabrous, nar- rowed into strongly curved, thin tip, along margin floccose- and arach- noid-hairy above. Florets pinkish-lilac, large; corolla 40 mm long, in broader part 17-18 mm long, cut up to 1/3. Pappus yellowish-whitish, with soft hairs, inner hairs apically narrowed and toothed, with slightly clavate tips; achenes marbled, greenish, with blackish-brown stripes, to 6 cm long. Flowering August to September. Rubble slopes and meadows, subalpine to alpine zone, as weed.— Caucasus: Central Caucasus (in upper reaches of Urukh, Balkar Cherek and Bezingi Cherek, and Chegem rivers). Endemic. Described from basin of Bezingi Glacier. Type in Tbilisi. Note. It differs from the polymorphic C. pugnax Somm. and Lev. by the white-tomentose pubescence of the entire plant, wider, shal- lowly pinnatifid leaves, and the usually larger capitula, which pro- jected above the cauline leaves. Series 6. Tomentosa Charadze.—Subsect. Eriophora Petrak 3. Horrida Petrak in Bibl. Bot. 78 (1912) 6 p. p—Capitula medium- sized, subsessile, terminal on stem and branches, usually clustered in 2-3 or solitary, surrounded by more or less longer apical leaves. Leaves narrow, oblong or linear-oblong, pinnately divided, with more or less deltoid-ovate lobes, white-tomentose, occasionally more or less gla- brous. 39. C. tomentosum C.A.M. Verzeichn. (1831) 69; DC. Prodr. VI, 637; Ldb. Fl. Ross. II, 730; Grossh. Fl. Kavk. IV, 181 p. p.; Opred. Rast. Kavk. 483 p. p.; Charadze in Zam. po Sist. i Geogr. Rast. Tbil. 20, 45; in Fl. Azerb. VIII, 396.—C. munitum y. tomentosum Boiss. Fl. or. III (1875) 533.—C. horridum var. tomentosum Petrak in Tr. Tifl. Bot. Sada, XII, 1 (1912) 20.—C. horridum ssp. tomentosum Petrak in Izv. Kavk. Muz. VIII, 1-2 (1914) 48 p. p.—C. horridum ssp. isophyllum Petrak, op. cit. (1914) 49.—C. isophyllum Grossh. Fl. Kavk. IV (1934) 181; Opred. Rast. Kavk. 483.—Ic. Petrak, op. cit. (1914) 1-2, plate III.—Exs.: Herb. Fl. cauc. No. 199. 119 120 118 Perennial. Rhizome thick, ligneous, producing 1-5 aerial shoots. Stem branched from base, ligneous, covered with brown-reddish-brown remnants of petioles of old leaves, ascending, weakly sulcate, white- tomentose, (20) 40-60 cm high, simple or branched above. Leaves white-tomentose beneath, scabrous above from small, scattered spiny bristles, grayish- or whitish-arachnoid-hairy to white-tomentose; lower cauline leaves 20-26 cm long, 3—5 cm wide, basally narrowed into short petiole, acuminate, linearly oblong or linearly lanceolate, pin- nately parted; leaf segments unequally 2—3 parted into ovate-deltoid or lanceolate-deltoid lobes, narrowed into long, yellowish, hard spines 5— 8 mm long, occasionally spines to 12 mm long, sparsely spiny-ciliate along margin; middle and upper cauline reduced toward stem top, lin- early lanceolate to oblong-lanceolate, shallowly pinnately parted, sessile, basally slightly broadened, semiamplexicaul, with weakly decurrent auricles; apical leaves at base of capitula, narrow, pinnately parted, with distant lobes, slightly or almost 2 times surpassing capitulum. Capitula globose-ovate, 2.5 cm in dia, subsessile, usually solitary or in 2-3, terminal, on stem and branches. Involucre more or less floccose- arachnoid-hairy; involucral bracts greenish-yellow, glabrous, lustrous, weakly carinate above, outer ones ovate or oblong-ovate, tapered to more or less bent or almost straight, basally arachnoid-hairy, hard, yellowish spines; inner bracts oblong-lanceolate, narrowed into subulately acute tip. Florets red or pink, usually 17 mm long; limb 5- parted to middle, almost 1/2 as long as corolla tube. Pappus brownish- white, with soft hairs, inner hairs narrowed into scabrous tip; mature achenes not known. Flowering August to September. (Plate IV, Figes2%) Alpine zone in rubbly places, along glacial cirques, to 3,200 m.— Caucasus: Central Caucasus (Terek River gorge), Eastern Transcaucasia (Tfandag, Shakhdag), Dagestan (sources of Avar Kois and Samur rivers), Southern Transcaucasia (Murovdag, Sevan and Zangezur ranges). General distribution: Armenia and Kurdistan. Described from Eastern Caucasus. Type in Leningrad. ; Note. It differs from all the species of the series Tomentosa by having white-tomentose pubescence over the entire plant; the speci- men from the Megrin District, which requires further investigation, especially stands out by its dense snow-white hairs. 40. C. daghestanicum Charadze in Zam. po Sist. 1 Geogr. Rast. Tbil. 22 (1961) 68; in Fl. Azerb. VIII, 398. Perennial. Stem hard, ascending, to 100 cm high, branched above with divergent branches, more or less densely arachnoid-hairy, leafy to tip. Leaves divergent, oblong, more or less broadly elliptical, pinnati- 121 119 fid-sinuate, with 2—-3-parted broadly ovate, approximate lobes, tapered to hard, yellowish spines, up to 12 mm long, loosely grayish tomentose beneath, tuberculate above, sparsely spinose, sometimes arachnoid- hairy; lower cauline leaves narrowed toward base, usually to 25 cm long, 8 cm wide, acuminate; middle and upper cauline leaves auricu- late, semiamplexicaul, middle ones to 17 cm long; apical leaves at base of capitula, strongly reduced. Capitula on short peduncles, soli- tary, terminal, on stem and branches, forming lax corymbose-panicu- late inflorescence, 3—4 cm in dia, basally with 2—3 linear-lanceolate, plumosely spiny-toothed apical leaves, somewhat surpassing capitula. Involucre white-arachnoid-hairy; involucral bracts from outer oblong- lanceolate to inner linear-lanceolate, tapered to erect or, in inner bracts, curved long spines, glabrous below, smooth above, weakly arachnoid- hairy, with dark spots, along margin scabrous. Corolla pinkish, 23 (24) mm long, limb 8-9 mm long. Pappus yellowish-white, with long, apically serrate and clavately thickened inner hairs; achenes to 4 mm long, dark reddish-brown. Flowering August to September. Rubbly talus, in subalpine zone.—Caucasus: Dagestan (basin of Samur River). Endemic. Described from Gashelk Range. Type in Tbilisi. Note. In its broad leaves and solitary capitula, it is closer to the series Pugnacia Charadze but differs by having narrower outer involu- cral bracts, which allies it with the series Lappacea Petrak. 41. C. ketzkhovelii Charadze in Zam. po Sist. i Geogr. Rast. Tbil. 22 (1961) 64. Perennial. Stem ascending, 30—40 cm high, thinly grayish tomen- tose, simple or weakly branched above, densely leafy to tip. Leaves oblong or linear-oblong, divergent, green above, more or less densely spiny-setose, with thin appressed bristles, sometimes thinly arachnoid- hairy, whitish beneath, finely tomentose, pinnatifid into 2—3-parted, ovate lobes with 8-10 mm long teeth at apex. Cauline leaves oblong, apically narrowed, semiamplexicaul, 15—23 cm long, 3.5—5.0 cm wide; nearer stem tip somewhat reduced, lower leaves basally narrowed; apical leaves at base of heads linear or lanceolate, pinnatipartite, long spiny, somewhat surpassing capitula. Capitulum on short peduncles, solitary or 2-3, terminal or axillary, in corymbose paniculate inflorescence. Involucre 2.5—3.0 cm in dia, thinly arachnoid-hairy, with erect or slightly deflexed, long yellowish spines; involucral bracts from outer ovate to inner linear-lanceolate, glabrous, lustrous, tapered: to thin, slightly curved, long spine, greenish-yellowish, dark brownish-green above, along margin scabrous, at base of spine long-arachnoid-hairy. Florets purple-lilac; corolla to 21 mm long, limb 10 mm. Pappus brownish- white, its outer hairs plumose up to tip, inner hairs long, serrate and 122 120 clavately thickened at apex; achenes brownish-reddish-brown, 4.5 mm long. Flowering August to September. Alpine zone, on rubbly talus.—Caucasus: Dagestan (sources of Avar Kois), Eastern Transcaucasia (Kakhetia, upper reaches of Argun River). Endemic. Described from upper reaches of Tetr Durudzh in Kakhetia. Type in Tbilisi. Note. It differs from C. tomentosum C.A.M. by the leaves that are white-tomentose only beneath and green and densely spiny-setose above and by the apical leaves slightly surpassing the capitula. From C. daghestanicum Charadze it differs by having narrower leaves that are densely white-tomentose beneath and a compact general inflorescence. 42. C. tindaicum Charadze in Zam. po Sist. i Geogr. Rast. Tbil. 22 (1961) 66. Perennial. Stem hard, ascending, usually simple, 40-50 cm high, dark purple, subglabrous, with scattered, crisped, articulate hairs, leafy to tip. Leaves divergent, linearly lanceolate, semiamplexicaul, thin- acuminate, middle cauline leaves to 18 cm long, 4—6 cm wide, sinuately pinnatifid into 2—3-parted lobes, lanceolate, segments narrowed into hard spines as long as or scarcely surpassing them, green on both sides, very thinly appressed-spiny above, with prominent, stramineous, basally broadened midrib and lateral veins, more or less glabrous beneath, sparsely arachnoid-hairy, more distinctly on midrib, lateral veins very prominent, whitish, thick, continuing in stiff, thick, to 12 mm long spines; apical leaves, slightly surpassing capitula, narrowly lanceolate, pinnately parted. Capitula 13-18 mm in dia, terminal, 5-8, clustered on short peduncles, axillary capitula reduced, solitary on peduncles somewhat longer than capitula, in racemose-corymbose inflorescence. Involucral bracts lanceolate, imbricate, glabrous, along margin scabrous, greenish-yellow, scarious, above blackish-purple, tapered to thin, yellowish, somewhat divergent spine, somewhat sur- passing involucral bracts. Florets pale-pink, to 20 mm long, limb about 9 mm. Pappus sordid-white, basally yellowish, with strongly elongated, scabrous tips of inner pappus hairs; achenes to 5 mm long, dark brown. Flowering July to September. Rubbly talus, in alpine zone.—Caucasus: Dagestan (upper reaches of Andii Kois, in Avar Kois Gorge), Eastern Transcaucasia (mountain- ous Tushetia). Endemic. Described from Dagestan (Gaer-meer Moun- tain near the village of Tindi). Type in Tbilisi. Note. This species is readily distinguished from other species of the series Tomentosa Charadze by the narrow leaves that are green on both sides, almost concolorus, glabrous or thinly arachnoid-hairy with hard spines, and also by the pale pink color of the florets. With its 123 121 clustered small capitula it approaches C. tomentosum C.A.M., differ- ing from it by the absence of white-tomentose pubescence and by having a compact racemose inflorescence with more numerous capitula. Series 7. Macrocephala Charadze.—Capitula large, solitary; in- volucral bracts tapered to straight, thick spines; pubescence of multi- cellular slightly scaly, crisped hairs; leaves subglabrous or grayish- arachnoid-hairy, with sparse spiny bristles above. 43. C. macrocephalum C.A.M. Verzeichn. (1831) 70; DC. Prodr. VI, 640; Ldb. FI. Ross. II, 730; Charadze in Zam. po Sist. 1 Geogr. Rast. Tbil. 20, 44; in Fl. Azerb. VIII, 395.—C. horridum ssp. prasinolepis Petrak in Izv. Kavk. Muz. VIII, 1-2 (1914) 47.—C. prasinolepis Grossh. Fl. Kavk. IV (1934) 181; Opred. Rast. Kavk. 483.—C. horridum 8. eriocephalum Boiss. Fl. or. III (1875) 533.—C. tomentosum Trautv. in Tr. Peterb. Bot. Sada, X, 1 (1887) 118, non C.A.M. Perennial. Stem up to 40 cm high, ascending, yellowish, thick, covered with long, intertwined arachnoid hairs, leafy to tip, simple or weakly branched at apex, with large, solitary capitula. Cauline leaves sessile, basally rounded, auriculate, semiamplexicaul, narrowed toward apex, ovate or oblong-ovate, 12-15 cm long, 6-7 cm wide; basal leaves oblong-lanceolate, abruptly narrowed into more or less elongated peti- ole, 25 cm long, 9-10 cm wide; all leaves shallowly sinuately pin- nately divided in 2—3-parted, ovate or deltoid-ovate segments, lobes acuminate, with yellowish, ligneous lateral veins, prominent on both sides and narrowed into hare, thick spines to 10 mm long, more or less arachnoid-hairy above or glabrous, usually with scattered spinose bristles less often lacking bristles, more or less grayish-arachnoid- hairy beneath to almost white-tomentose, occasionally arachnoid-hairy only along veins; apical leaves reduced,.crowded at base of capitula, broadly lanceolate, remotely pinnately toothed with long spines. Ca- pitula 5S—6 cm in dia, globose, solitary, terminal on stem and slender branches, subsessile, surrounded by apical leaves scarcely surpassing them. Involucre unevenly grayish-hairy; involucral bracts glabrous below, lustrous, yellowish-green, outer ones ovate, inner linearly lan- ceolate, weakly carinate, narrowed from middle into long greenish tip, tapered to erect, yellowish, thick spine, densely pubescent above, along margin, with long crisped hairs, at base with scaly hairs; outermost bracts oblong-lanceolate and pilose almost from base. Florets reddish, 33-36 mm long; limb 15 mm long, 5-parted to middle. Pappus 26—28 mm long, sordid-white, inner pappus hairs apically scabrous, weakly clavate, long; achenes dark brown, to 7 mm long. Flowering August to September. 124 122 Rubbly talus and stony slopes in alpine and subalpine zones.— Caucasus: Eastern Transcaucasia (Kuba District), Dagestan. Endemic. Described from Eastern Caucasus (Tfandag). Type in Leningrad. Note. The name C. macrocephalum C.A.M. has been broadened by authors of the Caucasian flora to include C. pugnax Somm. and Lev. (= C. horridum (Adams) Petrak). The type of C. macrocephalum, from Tfandag Peak, differs sharply from C. pugnax in several charac- ters, mainly in the nature of the pubescence of the involucral bracts, and in having broad leaves. The distribution of C. macrocephalum in Dagestan and the Eastern Caucasus is highly restricted. Despite this, the species varies greatly in leaf pubescence. Petrak described the pubescent form as ssp. prasinolepis Petrak, which was later elevated by Grossheim to the rank of species as C. prasinolepis (Petrak) Grossh. A densely arachnoid-hairy specimen of this species was cited by Trautvetter as C. tomentosum C.A.M. Series 8. Argillosa Charadze.—Capitula solitary, surrounded by apical leaves surpassing them. Leaves coriaceous, stiff, green above, snow-white beneath, shallowly sinuately-pinnately lobed. 44. C. agrillosum V. Petr. ex Charadze in Zam. po Sist. i Geogr. Rast. Tbil. 20(1958) 50; Grossh. Opred. Rast. Kavk. 458; Charadze in Fl. Azerb. VIII, 395.—Cnicus trautvetteri Wink]. in Schedis. quoad pl. e loc. Chunzach. Perennial. Stems robust, erect, branched above, 60—100 cm high, more or less densely arachnoid-hairy, almost snow-white, solitary or 2—5. Leaves stiff, coriaceous, elongate-elliptical or ovate-oblong, snow- white-tomentose beneath, sparsely spinose-setose above, narrowed above into hard spines to 10 mm long, semiamplexicaul, shallowly and remotely sinuately pinnately lobed, with 2-parted ovate-deltoid lobes, tapered to hard spines, 8-10 mm long, and sparsely spinose along margin; lower cauline leaves 15—25 cm long, 4—8 cm wide, others gradually reduced toward stem tip. Capitula terminal, solitary or in 2— 3, surrounded by apical leaves surpassing them, basally broadened, 2.5 cm in dia, with florets about 4 cm long. Involucre scatteredly floccose- arachnoid-hairy; involucral bracts scarious, outer ones oblong-ellipti- cal or ovate, yellowish, abruptly narrowed into elongate, yellowish spine, almost as long as bract; inner bracts linearly lanceolate, nar- rowed into cusp. Corolla pinkish-violet, to 30 mm long; limb 12.5 mm long. Pappus sordid-white, to 22 mm long, inner pappus hairs apically serrate, scarcely thickened at tip; achenes light brown, to 5.5 mm long. Flowering August to September. (Plate VI, Fig. 1.) 125 123 Mid-montane to subalpine zone, in rubbly places, among phryganoid vegetation, mainly in limestone areas.—Caucasus: Eastern Transcaucasia (Kuba District), Dagestan. Endemic. Described from Alta-Agach in Eastern Caucasus. Type in Baku. Note. A characteristic species of the central limestone region of Dagestan, somewhat isolated from the Caucasian-Western Asian spe- cies of the series Tomentosa Charadze, it is one of the xerophytes of Dagestan. Because of its snow-white pubescence on the stems and leaves, it sometimes has been confused by Caucasus authors with C. tomentosum C.A.M. or with C. sinuatum (Trrautv.) Boiss., although the latter species is sharply distinguished by the structure of its involu- cral bracts. Series 9. Turkestanica Charadze.—Subsect. Eriophora Petrak 3. Horrida Petrak in Bibl. Bot. 78 (1912) 6 p. p——Outer involucral bracts with erect, hard spines. Capitula large, arachnoid-hairy, surrounded by long-spinose apical leaves. Leaves with long thick spines. The series includes two species found in the mountains of Soviet Central Asia. The presence of narrow, long-pointed straight spines on the lower involucral bracts shows its closeness to the species of the series Lappacea, differing by having larger, densely arachnoid-hairy capitula and deltoid leaf segments bearing strongly thickened hard spines. The latter characters give a reason to ally this series with the series Pugnacia, with which Petrak compared C. turkestanicum (Rgl.) Petrak. He split this species from C. eriophorum s. 1., which differs considerably in the structure of the involucral bracts. 45. C. turkestanicum (Rgl.) Petrak in Oest. Bot. Zeitschr. LX, 9 (1910) 351.—C. eriophorum y. turkestanicum Rgl. in Tr. Peterb. Bot. Sada, VI, 2 (1880) 319.—C. eriophorum var. turkestanicum O. and B. Fedtsch. Perech. Rast. Turk. IV (1911) 283. Perennial. Stem 40-80 cm high, thick, simple, more or less densely white tomentose, densely leafy. Leaves with numerous appressed spine- like bristles above, more or less thinly arachnoid-hairy, with yellowish prominent veins and midrib broadened at base, densely grayish-tomen- tose beneath, with very prominent thick and ligneous midrib and lat- eral veins extending to apices of lobes as hard, stramineous spines, oblong or ovate-oblong, deeply sinuately pinnately divided into broadly deltoid-lanceolate or deltoid-ovate segments, spiny-ciliate on margin, to 15-17 cm long; apical segments oblong, lanceolately tapered into hard spines up to 10 mm long, lateral segments usually nonuniformly 2-parted, with more or less reduced anterior lobe; all lobes and teeth tapered to thick, hard spines 10-12 mm long; basal and lower cauline 126 124 leaves more deeply pinnately incised, to 25 cm long, narrowed in more or less short, narrow-winged spinulose petiole, middle and upper leaves sessile, with long semiamplexicaul auricles, 2-parted into deltoid lobes, tapered to long, hard spines; rosette of small leaves often in axils of upper leaves surrounding undeveloped axillary capitula; apical leaves at base of capitula, morphologically similar to remaining leaves, al- most as long as or surpassing capitula. Capitula 3—4, globose-ovate, 5—6 cm in dia, subsessile, clustered at stem apices in shortly racemose inflorescence, surrounded by deeply pinnately divided apical leaves surpassing them. Involucre densely grayish arachnoid-hairy; outermost involucral bracts scabrous above, gradually narrowed from broad base into long straight spines, densely arachnoid-hairy, other bracts from outer ovate-oblong to inner oblong-broadly lanceolate, glabrous, scari- ous, carinate, gradually narrowed above middle into linearly lanceolate, slightly curved tip, weakly arachnoid-hairy above. Florets pink, 30—32 mm long; limb as long as narrow corolla tube or somewhat longer, to middle 5-parted. Pappus golden, sordid-white above, with scarcely long, scabrous tips of inner pappus hairs; achenes brown, to 6 cm long. Flowering August to September. Stony slopes, from 2,200 to 3,500 m.—Soviet Central Asia: Tien Shan, Pamiro-Alai. Endemic. Described from Darvaz. Type in Geneva; paratype in Vienna; isotype in Leningrad. 46. C. pseudolappaceum Charadze in Zam. po Sist. 1 geogr. Rast. Tbil. 23 (1963) 115; in Fl. Turkm. VII, 240, descr. ross.—C. lappaceum var. ferox auct. fl. As. Med. non Boiss.—C. Jappaceum auct. non MB.: O. and B. Fedtsch. Perech. Rast. Turk. IV (1911) 283.—C. lappaceum . monocephalum Bge. Beitr. Kenntnis Russl. Stepp. Centr.-As. (1851) 190.—Exs.: Sint. It. transcasp.-pers. No. 1097. Perennial. Stem basally with brown fibrous remnants of old leaves, ascending, ligneous, 40-70 cm high, densely arachnoid-hairy or al- most tomentose, sulcate, simple or with short axillary shoots above, leafy up to apex. Cauline leaves oblong, more or less divergent, sinuately pinnately parted into broadly deltoid-lanceolate, remote seg- ments, densely spiny-setose above with thin, more or less appressed or erect bristles, densely white-tomentose beneath, with thick midrib and lateral veins, tapered to long, hard spines; basal and lower cauline leaves narrowed to more or less long petiole; all leaves lanceolate, cauline leaves sessile, auriculate, semiamplexicaul, spinose-ciliate on margin, with slightly revolute margin; apical leaves pinnately divided into narrow lobes, slightly narrowed, tapered into long, hard spine, surrounding capitula, almost 1.5 times as long. Capitula terminal or axillary, subsesille, solitary, in short racemose inflorescence. Involu- 127 125 cre covered with weakly floccose, white-arachnoid hairs, terminal capitula 2.5—-3.5 cm in dia, axillary sometimes 1.5 cm in dia with florets 4 cm long; outer involucral bracts broad at base, subulate, with straight spine, 1/2 as long as involucre, more or less arachnoid-hairy, from outer oblong-ovate to inner linearly lanceolate, glabrous, green- ish-yellow, weakly carinate, narrowed from middle into thin, arach- noid-hairy spines. Florets pink, to 30 (32) mm long; corolla tube to 16 mm long. Pappus basally golden-yellow, sordid white above, to 23 mm long, with elongate scabrous tips of inner hairs; achenes 5 mm long, brown. Flowering July to September. Juniper zone, on gypsiferous hills and dry mountain slopes to 2,800 m.—Soviet Central Asia: mountainous Turkmenia, Pamiro-Alai Re- gion (Guzar, Baisun). General distribution: Iran-Afghanistan. Described from Arvaz in Kopetdag. Type in Alma-Ata. Note. It differs from C. turkestanicum (Rgl.) Petrak by having oblong, smaller capitula, floccose pubescence on the involucre, and leaf segments long-attenuated into hard spines. From the series Lappacea, with which this species usually has been allied, it differs by having broader lower involucral bracts, a different leaf form with distant segments, and other characters that give this plant a unique appearance. Subsection 5. Eriophora Petrak in Bibl. Bot. 78 (1912) 5 p. p— Subsect. Eriophora 1. Eriocephala Petrak, 1. c—Leaves grayish arach- noid-hairy beneath; cauline leaves sessile, not decurrent. Capitula usu- ally large, occasionally, medium-sized. Involucral bracts apically scarious, colored, more or less broadened, fimbriate along margin, tapered to thin, more or less short, spines, scabrous on margin or with occasional spinous cilia. The subsection includes species distributed in the Mediterranean region and replaces the subsection Armata Charadze. 47. C. polonicum (Petrak) Ijin in Izv. Glavn. Bot. Sada Akad. Nauk SSSR, XXX, 3-4 (1932) 350; Majevskii, Fl. ed. 8, 603; Klok. in Vizn. Rosl. URSR, 568; Geideman, Opred. Rast. Mold. SSR, 289.— C. eriophorum ssp. decussatum Petrak var. polonicum Petrak in Bibl. Bot. 78 (1912) 35.—C. eriophorum var. spathulatum auct. non Naeg. and Lohr.: Malz. in Sitzb. Naturf. Ges. Uni. Jurj. XVI, 2 (1907) 129; Schmalh. Fl. Yugo-Zap. Ross. 317; Fl. II, 101.—C. eriophorum var. platyonichium auct. non Wallr.: Fedtsch. and Fler. Fl. Evrop. Ross. (1910) 1009.—C. eriophorum auct. fl. ross. non Scop.: Bess. Enum. pl. Vohlyn. (1822) 32; Ldb. Fl. Ross. II, 731 p. p.—Ic.: Petrak in Bibl. Bot. 78, fig. 10.—Exs.: Fl. polon. exs. No. 957 (sub C. decussato Janka); GRF No. 115. 12 oo 126 Biennial. Stems robust, crisped-hairy, weakly flexuous, thick, sul- cate, more or less densely lanate, 60-150 cm high, branched above, leafy to top. Leaves almost to base pinnately incised into divergent, linearly lanceolate, unequally 2-parted segments, with shorter apical lobe, narrowed into hard spine, finely and unequally spiny-ciliate along margin, with long, thin, spinous bristles above, grayish-tomentose be- neath, with arachnoid-hairy, prominent veins. Lower leaves up to 37 cm long, elliptical, with narrow segments, tapered to long, hard spine, basally narrowed to spinose-lobate, auriculate petiole, semiamplexicaul; middle cauline leaves sessile, semiamplexicaul, narrowed at apex, gradu- ally reduced upward; apical leaves as long as capitula or 2—3 times surpassing them. Capitula solitary, terminal on stem and few long shoots, 6-8 cm in dia, in corymbose paniculate inflorescence. Involucre more or less densely lanate; involucral bracts linearly lanceolate, with thin and occasional spiny cilia along margin, abruptly broadened above, scarious, fimbriate along margin, obtuse, with thin yellowish spine, somewhat recurved, purple; inner bracts linearly lanceolate, gradually narrowed into long, arachnoid-hairy tip. Corolla red, 34 mm long, tube narrow, to 22 mm long; limb incised to middle. Pappus sordid-white, to 25 mm long, with elongate, toothed tips of inner hairs; achenes to 6 mm long, 2.3 mm wide, light reddish-brown, with black stripes. Flowering July to September. Roadsides, scrubs, fields, mountain slopes.—European Part: Up- per Volga, Volga-Kama, Upper Dnieper, Middle Dnieper, Volga-Don, Trans-Volga, Lower Volga. General distribution: Central Europe. De- scribed from vicinity of Tula. Type in Leningrad. Note. It differs from C. decussatum Janka by having slightly ob- tuse, oblong-ovate and oblong-lanceolate appendages of the involucral bracts, which abruptly end in a short spine; in C. decussatum Janka, the involucral bracts are gradually narrowed and attenuated into a spine at the apex. C. polonicum (Petrak) Iljin is distributed in the eastern part of the range of the subsection Eriophora Petrak and replaces C. decussatum Janka, distributed in the Ciscarpathian part of Eastern Europe. Subsection 6. Ciliata Petrak in Bibl. Bot. 78 (1912) 5 p. p.— Lophiolepis Cass. in Dict. Sc. nat. XLI (1826) 313, pro sect.; DC. Prodr. VI, 634 p. max. p.—Subsect. Ciliata 1. Serrulata Petrak op. cit. 5 p. min. p., 2. Ciliata Petrak, op. cit. 5, 3. Fimbriata Petrak, op. cit. 5 p. min. p.—Capitula large, globose. Involucre ovate-roundish, gla- brous or weakly hairy; apical leaves completely or partly surrounding capitulum. Involucral bracts imbricate, appressed, linearly lanceolate, spinescent, more or less incised, spiny-ciliate or entire; outer bracts upward-spreading, outer deflexed. 129 127 Type of subsection: C. ciliatum (Murr.) Moench. I have excluded Caucasian species with nodding capitula from this subsection, which I place in the separate section Caucasigena Charadze. Some species of [Series] 1. Serrulata Petrak have been included in the subsection Armata Charadze. 48. C. serrulatum (MB.) MB. FI. taur.-cauc. III (1819) 557 p. p. excl. var.; DC. Prodr. VI, 638; Ldb. Fl. Ross. II, 725; Schmalh. FI. II, 101; Fedtsch. and Fler. Fl. Evrop. Ross. 1010, excl. syn.; Nevski in Sorn. Rast. SSSR, 4, 289; Pavlov, Fl. Tsentr. Kazakhst. III, 311; Klok. in Vizn. Rosl. URSR, 568 p. p.—Cnicus serrulatus MB. FI. taur.-cauc. II (1808) 275. Biennial. Stem 100-150 cm high, hard, ascending, corymbosely branched above, leafy to top, sulcate, sparsely crisped-hairy. Leaves green above, more or less densely covered with fine spiny bristles, grayish-tomentose beneath, finely spiny-ciliate along margin; lower leaves oblong-elliptical, 18—25(30) cm long, narrowed into petiole, pinnately incised almost to base into deltoid-lanceolate, 2-parted seg- ments, with long, broadly or narrowly lanceolate, acuminate, at apex somewhat subobtuse lobes; lobes and teeth long tapered to hard spines 4—6 mm long; middle and upper cauline leaves semiamplexicaul, au- riculate, ovate or ovate-lanceolate, deeply two-parted into unequal, lanceolate or linearly lanceolate lobes, long tapered to thin spines 7— 9 mm long; apical leaves to 1.5—2 cm long, occasionally smaller, ovate, shallowly-pinnately lobed, usually reaching base of capitula. Capitula on short, thick peduncles, erect, solitary, terminal on stem and branches, often 5—6 clustered in corymbose inflorescence. Involucre weakly cu- neate at base, 2.5—-3.5 cm in dia, more or less oblong, 3—4 cm long, somewhat expanded at end of flowering; outer involucral bracts strongly reduced, more or less deflexed, lanceolate, long-tapered to erect, hard, short spine, with fine, upward spreading spiny cilia along margin above; middle and inner involucral bracts elongate, lanceolate to linearly lan- ceolate, outer ones gradually narrowed into erect, stiff spines, inner ones into scarious tip, thinly spiny-ciliate along margin with upward- spreading and appressed cilia, stramineous, carinate above, densely thin appressed-hairy along margin. Florets red, 27-30 mm long; co- rolla tube 14-16 mm long; limb cut up to 1/3. Pappus to 19 mm long, sordid-white, yellowish below, with dark, elongate and toothed tips of inner hairs; achenes 4 mm long. Flowering July to September. On solonetzes in forest-steppe zone; meadows, roadsides, irriga- tion channels.—European Part: Volga-Don, Trans-Volga, Bessarabia, Black Sea Region, Crimea, Lower Don, Lower Volga; Western Sibe- ria: Upper Tobol (cited for the former Aktyuba and Irgiz districts). Endemic. Described from Crimea. Type in Leningrad. 13 =) 128 Note. The species varies in the degree of incision of the leaves, width of their segments, and to some extent, size of the capitula. The most typical forms from Crimea are distinguished by very small, nar- row lower involucral bracts. In Ukraine, forms with somewhat larger capitula are found, which are transitional to C. ciliatum (Murr.) Moench, whose northern limit of distribution overlaps the range of C. serrulatum. An especially large number of intermediate forms is observed at the eastern limit of distribution of the species in the Lower Volga and Trans-Volga regions. Authentic specimens of C. ciliatum are known from these same places. C. ukranicum Bess., which has been cited as a synonym of C. ciliatum, differs by having long, deflexed lower in- volucral bracts similar to those of C. ciliatum. Possibly C. ukranicum Bess. is one of the intermediate forms, and whether it is a good species can be resolved finally only by careful study in nature. I am refraining from including it as synonym of C. serrulatum. 49. C. ukranicum Bess. in DC. Prodr. VI (1837) 635.—C. ciliatum hybridum Fisch. Cat. Hort. Gorenk. (1812) 35, nomen.—C. ciliato- affine Bess. Enum. pl. Volhyn. (1822) 75, nomen.—C. serrulatum MB. Fl. taur.-cauc. III (1819) 557 p. p.; Klok. in Vizn. Rosl. URSR, 568 p. min. p. Biennial. Stem ascending, 40-60 cm high, weakly flexuous and branched above, more or less densely crisped-hairy, slightly ribbed, brownish, densely leafy to tip. Leaves divergent, oblong-lanceolate; middle cauline leaves 9-11 cm long, up to 3 cm wide, sinuately pin- nately parted into unequally 2-parted, remote, lateral segments, lobes of segments lanceolate, subacute, strongly spinescent; lower leaves narrowed into short petioles, others basally auriculate, semiamplexicaul, accuminate; apical leaves reduced, lanceolate, upward spreading, often reaching upper margin of involucre; all leaves green above, with small, appressed, spinose bristles, densely grayish-tomentose beneath, with prominent midrib and lateral veins, usually stiff and strongly tapered to spines 4-8 mm long. Capitula solitary, globose-ovate, terminal on stem and branches, usually short-pedunculate, 3—4 clustered in corym- bose inflorescence. Involucre glabrous, 2.5—3.5 cm in dia; involucral bracts from outer narrowly lanceolate to inner linearly-lanceolate, elon- gate, gradually narrowed into thin, long spines, finely setose-ciliate along margin; outer and middle bracts carinate, with longer cilia, outer ones usually recurved, dorsally along margin thinly appressed-hairy, scabrous; inner bracts dark violet above, narrowly linearly lanceolate, narrowed into thin, long, weakly bent cusp. Florets purple, to 27 mm long; corolla tube narrow, to 16 mm long; limb unequally 5-parted to deeper than middle. Pappus sordid-white, with scabrous, elongated 132 129 tips of inner hairs; mature achenes unknown. Flowering July to Sep- tember. Steppe slopes and pastures.—European Part: Bessarabia, Upper Dniester. General distribution: Central Europe (Transylvania). De- scribed from Podolia. Type in Kiev (?). Note. In the pubescence of the floral bracts, it is closer to C. serrulatum (MB.) MB., differing from it by having larger capitula and longer lower involucral bracts, which are ciliate along margin, re- curved, and gradually narrowed to spines. From C. ciliatum (Murr.) Moench it is well differentiated by the form of the narrower involucral bracts. Apparently this species is cited by T. Geidemann (Opred. Rast. Mold. [1954], p. 289) for the flora of Moldavia under the name C. ciliatum (Murr.) Moench. 50. C. ciliatum (Murr.) Moench, Meth. Suppl. (1802) 227; MB. Fl. taur.-cauc. III, 556; Spreng. Syst. veg. 3, 372; DC. Prodr. VI, 635; Ldb. Fl. Ross. II, 725; Boiss. Fl. or. III, 527; Schmalh. Fl. II, 102; Grossh. Fl. Kavk. IV, 179; Charadze in Fl. Gruzii, VIII, 509; in FI. Azerb. VIII, 394.—C. ciliatum B. microcephalum Ldb. FI. Ross. II (1846) 735.—Carduus ciliatus Murr. Comm. Gétt. (1784) 35.—Cnicus ciliatus Willd. Sp. pl. III, 3 (1803) 1667; MB. FI. taur.-cauc. II, 275.— Lophiolepis calocephala Cass. in Dict. Sc. nat. XXVII (1823) 181.— Ic.: Murr. Comm. Gott. tab. 5. Biennial. Stem erect, (50)100—150 cm high, thickened, branched above, slightly ribbed and sulcate, with scattered, crisped, and finely arachnoid hairs. Leaves green and finely appressed-spiny-setose above, grayish and appressed arachnoid-hairy beneath, with prominent thin venation, oblong-elliptical, more or less deeply sinuately pinnately in- cised into oblong-broadly lanceolate, 2—3-parted segments; their lobes tapered to thin, hard spines 3-7 mm long, thinly spiny-ciliate along margin; lower cauline leaves petiolate, elongated, lobed, semi- amplexicaul, with long apical segment incised almost to base into deeply 2—3-parted, oblong lobes, to 35-40 cm long, 18 cm wide, segments usually with 2 almost equal lobes and 1 strongly reduced, ovate lobe, strongly spinescent; other cauline leaves sessile, auriculate, more or less semiamplexicaul; middle leaves 25 cm long, 6—7 cm wide, sinuately pinnately parted into unequal 2-parted segments, with lanceolate, strongly spinescent lobes; upper leaves gradually reduced; apical leaves usually 1—2, linearly lanceolate, more or less recurved, usually as long as capitula. Capitula solitary, terminal, subsessile, globose-ovate, (2)3— 4 cm in dia. Involucre subglabrous, weakly arachnoid-hairy; outer in- volucral bracts oblong-lanceolate, greenish, usually recurved, ciliate above, cilia long, thin, with yellowish spines to 7 mm long; middle 130 Plate VI. 131 1—Cirsium agrillosum V. Petr. ex Charadze; 2—C. sosnowskyi Charadze. 133 131 involucral bracts divergent, linearly lanceolate, stramineous, weakly carinate, with purple spot above, and finely ciliate; inner bracts scari- ous, stramineous, spinescent, along margin smooth. Florets violet-red; corolla 25-26 mm long, limb 11-13 mm long, 5-parted. Pappus sor- did-white, of numerous soft hairs, middle hairs with long filiform tips; achenes 5.5 mm long, yellowish-brown. Flowering August to Septem- ber. Roadsides, logged forests, forest meadows, limestone slopes.— European Part: Volga-Don, Black Sea Region, Lower Don, Lower Volga, Trans-Volga; Western Siberia: Upper Tobol (watersheds of Greater and Lesser Uzen); Caucasus: Ciscaucasia, Dagestan, Eastern and Western Transcaucasia. Endemic. Described from specimens grown in Gottingen Botanical Garden from seeds of an unknown collector. Note. Apparently, we must consider the Volga Region as_ the locus classicus of this species, which was attributed by 18th-century western authors to Siberia. In Trans-Volga this species is found fre- quently; farther west, it is reported sporadically and to the west (Hun- gary) it probably is replaced by C. furiens Griseb. and Schrenk. It is widespread throughout the Caucasus, growing as weed along roads. Among the South European forms a transition to C. serrulatum (MB.) MB. is observed, and in Southern Transcaucasia, to C. szowitsii (C. Koch) Boiss. 51. C. szowitsii (C. Koch) Boiss. Fl. or. III (1875) 527; Grossh. Fl. Kavk. IV, 179; Charadze in Fl. Gruzii, VIII, 510; in Fl. Azerb. VIII, 395.—C. ciliatum Szowits in schedis, non Moench.—C. ciliatum ssp. szowitsii Petrak in Izv. Kavk. Muz. VIII, 1-2 (1914) 40.—C. ciliatum var. xantholepis Petrak in Tr. Tifl. Bot. Sada, XII, 1 (1912) 2.—Epitrachys szowitsii C. Koch in Linnaea, XXIV (1851) 397.— Exs.: Petrak, Cirsioth. Univ. Nos. 85, 86, 139. , Biennial. Stem up to 100 cm high, geniculately ascending, branched above, weakly ribbed, sulcate, subglabrous, with scattered arachnoid hairs, or more or less arachnoid-hairy, densely leafy. Leaves stiff, coriaceous, yellowish-green and more or less densely appressed-spiny- setose above, involute, spiny-ciliate, whitish-arachnoid-hairy beneath or subglabrous and only along veins arachnoid-hairy; veins yellowish- white, prominent, thick, cauline leaves oblong or ovate-lanceolate, middle ones to 15—20 cm long, sinuately pinnately parted into un- equally 2-parted segments; lower cauline leaves longer, with oblong- lanceolate lobes, terminal lobes of segments strongly reduced, deltoid- ovate, tapered to often 3-toothed spines sometimes longer than them; upper leaves reduced, shallowly pinnate into deltoid-lanceolate or deltoid-ovate lobes; all cauline leaves sessile, with toothed, semi- 134 132 amplexicaul auricles, apically with long narrow segment. Capitulum globose-ovate, (2.5) 4-5 cm in dia, solitary, terminal on stem and leafy branches, subsessile, with short, thick peduncles, in more or less lax corymbose-paniculate common inflorescence. Involucral bracts coriaceous, glabrous, ovate below, stramineous, narrow above, more or less long, broadly lanceolate; outer and middle bracts purple above, stramineous along margin, with incised spiny-ciliate margin, long- tapered into erect, long, hard spines; usually outer and some middle bracts recurved; inner bracts carinate, narrowly linearly lanceolate, purple above, along margin sparsely spiny-ciliate, strongly tapered to slightly curved, hard spine. Florets purple; corolla to 32-35 mm long; corolla tube narrow, 20-22 mm long, limb unevenly incised to middle. Pappus with numerous yellowish-white hairs, inner hairs filiform with scabrous tips; achenes 5 mm long, grayish-brown with black stripes. Flowering July to September. Mid-montane zone; dry grassy slopes, forest edges, old fields, stony talus, river, gravels—Caucasus: Dagestan, Southern Trans- caucasia. General distribution: Armenia and Kurdistan, Iran. Described from Karabakh. Type in Berlin; isotype in Leningrad. Note. It differs from C. ciliatum (Murr.) Moench by having broader involucral bracts and very hard spines. Parsa (FI. Iran, III [1943] 563) reports it for Iran both as C. ciliatum and C. szowitsii; the occurrence of the former in Iran seems doubtful to us. Hybrid forms have been observed in the Caucasus. 1. C. ciliatum (Murr.) Moench x C. caucasicum (Adams) Petrak (C. x Petrakii Kozl. and Woron. in schedis): Grossh. Fl. Kavk. IV (1934) 189; Charadze in Fl. Gruzii, VIII, 494. Caucasus: Southern Transcaucasia (Borzhomi, Bakuriani). It grows among the parental forms. A series of transitions to C. ciliatum s. |. and C. caucasicum is observed. Apparently, C. szowitsii, as a species close to C. ciliatum, should be recognized as one of the parental species, which the broad purple involucral bracts of the hy- brid would indicate. From C. caucasicum the hybrid forms inherit long, often thin spines and weak but arachnoid-floccose pubescence of the involucre, nodding capitula, and dense white tomentum on the lower surface of the leaves. These forms are related to C. szowitsii (C. Koch) Boiss. by leaf form, broadly lanceolate, more or less spiny- ciliate, attenuate-spinescent involucral bracts, and almost straight, up- ward-directed capitula. In habit, there are some forms closer to C. szowitsii (C. Koch) Boiss. and others closer to C. caucasicum (Adams) Petrak. 135 133 2. C. aduncum Fisch. and Mey. ex DC. x C. ciliatum (Murr.) Moench.—C. x xenogenum Petrak in Fedde, Repert. XLIII (1938) 299. Caucasus: Southern Transcaucasia (Novo-Bayazet). This hybrid was grown by Petrak in Moravia from seeds collected in Novo-Bayazet by Grossheim and Edelmeyer. Some plants differed from C. aduncum by having leaves slightly narrowed at the base into petioles, with loosely grayish-arachnoid-tomentose pubescence on both sides, especially marked larger globose-ovate capitula that are not tapered to the base; and broader involucral bracts tapered to carinate, hard spines and with avery finely ciliate-spiny margin at the base. Petrak presumes that plants of C. ciliatum would have had to grow near C. aduncum. Apparently, in this case these could be plants of C. szowitsii, which are more widespread in southern Transcaucasia. Subsection 7. Lanceolata Petrak in Bibl. Bot. 78 (1912).—Leaves short- or long-decurrent. Capitula medium- or large-sized, globose- ovate or ovate. Involucre scatteredly arachnoid-hairy or subglabrous; involucral bracts oblong-lanceolate, gradually attenuated into thin spines, usually upward directed or outer bracts falcately decurved. Based on the form of the involucral bracts, the species of this subsection stand the closest of all to the subsection Ciliata, understood by us in a more restricted sense than by Petrak. Series Alata (Petrak) Charadze comb. nova.—Subsect. Lanceolata Petrak 2. Alata Petrak in Bibl. Bot. 78 (1912) 7—Stem winged. Leaves long-decurrent. Capitula somewhat projected above apical leaves. 52. C. vulgare (Savi) Ten. Fl. Nap. V (1835-1836) 209; Arénes in Bull. Jard. Bruxelles, XXIV, 4, 255; Charadze in Fl. Azerb. VIII, 394.—C. vulgare (Savi) Airy-Shaw in Fedde, Repert, XLIII (1938) 304; Grossh. Opred. Rast. Kavk. 481; Kirpichn. in Majevski, FI. Ed. 8, 603.—C. vulgare (Savi) Petrak, Cirsioth. Univ. IV (1912) 33, nom. provis. in adnot.—C. vulgare ssp. vulgare Arénes op. cit. 257.—C. vulgare ssp. silvaticum (Tausch) Arénes, op. cit. 259.—C. lanceolatum (L.) Scop. Fl. Carn. ed. 2, II (1772) 130, non Hill (1769).—C. lanceolatum auct. non Hill: DC. Prodr. VI (1837) 636; Ldb. FI. alt. IV, 4; Fl. Ross. II, 726; Schmalh. Fl. II, 102; O. and B. Fedtsch. Perech. Rast. Turk. IV, 283; Grossh. Fl. Kavk. IV, 178; Kryl. Fl. Zap. Sib. XI, 2886; Klok. in Vizn. Rosl. URSR, 569.—C. lanceolatum B. hypoleucum DC. Prodr. (1837) 636; Grossh. Fl. Kavk. IV, 178.—C. lanceolatum var. vulgare Naegeli ex Koch, Syn. ed. 2 (1843-1845) 990.—C. lanceolatum var. nemorale Naegeli ex Koch. ibid.—C. lanceolatum ssp. eulanceolatum Beger apud Hegi, Ill. Fl. VI, 2 (1929) 136 134 874.—C. lanceolatum ssp. hypoleucum (DC.) Beger apud Hegi, ibid. 875.—C. lanceolatum ssp. savianum Arénes in Not. syst. Mus. Hist. Nat. Paris, XIII, 1-2 (1947) 60.—C. lanceolatum ssp. silvaticum Arénes, ibid.—C. silvaticum Tausch in Flora XII, 1 (1829) 38.—C. nemorale Rchb. Fl. Germ. excurs. (1830) 286.—Carduus lanceolatum L. Sp. pl. (1753) 821.—Carduus vulgaris Savi, F1. Pis. II (1798) 241.— Ascaiea lanceolata (L.) Hill, Herb. Brit. I (1769) 72, tab. 51, fig. 2.— Lophiolepis dubia Cass. in Dict. Sc. nat. XX VII (1823) 183.—Eriolepis lanceolata (L.) Cass. in Dict. Sc. nat. XLI (1826) 331.—Ie.: Hill, Herb. Brit. I, tab. 51, fig. 2; Rchb. Ic. Fl. Germ. tab. 826, fig. 1, 2; Fedtsch. and Fler. Fl. Evrop. Ross. 1012; Dostal, Kvét. CSR, No. 547, 1; Hei, Ill. Fl. VI, 2, fig. 564, 565, 573.—Exs.: Fl. exs. austro-hung. No. 3769 (sub C. silvatico Tausch); GRF No. 2027. Biennial. Rhizome fusiform; stem (5)50—150 cm high, ascending, branched, sulcate, more or less crisped-hairy, with long and thinly arachnoid hairs, toothed- or spiny-winged. Leaves lanceolate, elongate to linearly lanceolate, densely spiny-setose above, arachnoid-hairy to white-tomentose beneath (var. hypoleucum), sinuately pinnately parted; middle cauline leaves 7-10 cm long, 3—4 cm wide, with long-decur- rent, toothed wings, their teeth strongly spinescent, spinulose on mar- gin; segments of leaves remote, usually deeply 2-parted into ovate- deltoid or lanceolate-deltoid lobes, long-tapered to thin, yellowish spines, 5—10 mm long, occasionally anterior lobe almost reduced; basal leaves with broadly ovate lobes, usually withering early; upper cauline leaves gradually narrowed; apical leaves reaching base of capitula. Capitula solitary on reduced peduncles, ovate, (2.5)3—4(5) cm in dia. Involucre roundish-ovate, with or without small bracts at base; involu- cral bracts lanceolately subulate, usually yellowish-green, blackish- brown above, weakly carinate, tapered to thin, yellowish spine, usually shorter than bracts; outer bracts falcately recurved from middle, floc- cose, subglabrous, inner ones linearly lanceolate, tapered to slightly bent spines. Florets red, (25)30-35 mm long, with very narrow and long corolla tube, 2 times as long as limb; latter lobed almost to middle. Pappus sordid-white, with thin lustrous hairs, long-plumose almost to tip; achenes 3.5—4 mm long, grayish. Flowering July to September. Lowlands to mid-montane zone; roadsides, abandoned fields, waste- lands, and stony places.—European Part: All regions; Caucasus: All regions; Western Siberia: Ob Region, Upper Tobol Region, Irtysh; Soviet Central Asia: Aralo-Caspian Region, Lake Balkhash Region, mountainous Turkmenia, Pamiro-Alai Region, Tien Shan. General dis- tribution: Europe (except Arctic), Mediterranean Region, Balkans-Asia Minor, Armenia and Kurdistan, Iran and Afghanistan, Dzhungaria- Kashgaria, North America (introduced). Described from Italy. Type in Italy. 137 135 Section 3. Lamyropsis Charadze in Addenda XXVII, 603.—Ca- pitula on short peduncles, projected above leaves. Outer involucral bracts narrowed to deltoid-lanceolate scarious-tip, spinescent, usually spines 2-3 times surpassing lower broader part of bracts; inner bracts shorter or almost as long as outer. Achenes compressed, with oblong tubercle at apex. Pappus of thin fragile hairs, inner hairs plumose almost up to apex, with filiform tips. Perennial plants with more or less coriaceous leaves or glabrous, or subglabrous above, white- tomentose beneath; lateral segments of leaves reduced and strongly tapered to hard spines. Type of section: C. sinuatum (Trautv.) Boiss. Note. This section is represented by three species, which replace each other in parts of the southern Balkans, the Caucasus, and Soviet Central Asia. In their habit, species of the section Lamyropsis ap- proach members of the genus Lamyra Cass. but differ from them by the structure of achenes and pappus. They show some similarity to species of the section Echenais, differing from them in the glabrous peduncles, capitula lacking apical leaves, form of the involucral bracts, and also coriaceous leaves that are snow-white beneath. The southern Balkan species C. cynaroides (Lam.) Spreng. was referred by de Candolle to the section Platyraphium of the genus Chamaepeuce; and recently it has been included by Tamamschian in the genus Lamyra Cass. as L. cynaroides (Lam.) Tamamsch. (in Zam. po Sist. i Geogr. Rast. Tbil., No. 22, p. 81). However, even Boissier noted that this species is closer to Cirsium in the structure of achenes. I consider it best to assign this species to section Lamyropsis of the genus Cirsium. 53. C. sinuatum (Trautv.) Boiss. Fl. or. III (1875) 546; Grossh. Fl. Kavk. IV, 187; Opred. Rast. Kavk. 485; Charadze in Fl. Gruzii, VIII, 514; in Fl. Azerb. VIII, 402; Papava in Vestn. Gos. Muz. Gruzii, XVA, 32.—C. cynaroides MB. FI. taur.-cauc. III (1819) 559, non Spreng.—Cnicus cynaroides MB. FI. taur.-cauc. II (1808) 280, non Willd —Chamaepeuce sinuata Trautv. in Tr. Peterb. Bot. Sada, II (1873) 477.—C. cynaroides DC. Prodr. VI (1837) 659 p. p.; Ldb. FI. Ross. II, 745 p. p.—Exs.: Herb. Fl. cauc. No. 394. Perennial. Stems basally woody, ascending, more or less divaricately branched, densely leafy up to apex, 30-40 cm high, ap- pressed-white-tomentose. Leaves coriaceous, lower ones oblong-ellip- tical, usually 8-12 cm long, 3.5 cm wide upper ovate-lanceolate, gradu- ally reduced, all sessile, basally narrowed, acute, lustrous and green above, with prominent, whitish lateral veins, glabrous or along veins weakly arachnoid-hairy, snow-white or densely appressed-tomentose 138 136 beneath, on margin broadly sinuately pinnately lobed; lobes usually 2- parted, acuminate, with thick veins, strongly tapered to hard, yellowish spines 4—5 mm long. Capitula solitary, terminal on slightly curved stems and long axillary branches on short peduncles, somewhat pro- jected above small apical leaves. Involucre ovate, 3—4 cm long, 2-3 cm in dia, weakly arachnoid-hairy with scattered hairs; outer involu- cral bracts ovate, narrowed from middle to subulate, spiny, falcately recurved tip; middle bracts oblong-ovate, glabrous, carinate, scarious, stramineous, with tuberculate purple spots, along margin floccose white- arachnoid, narrowed to lanceolate subulate, strongly spinescent tips longer than them, with divergent or upward spreading spines; inner bracts broad, linearly lanceolate, scarious, with narrow, scaly, more or less short point, usually shorter than outer bracts. Corolla dark-pink, to 22 mm long; limb unequally 5-partite almost to middle; anthers 10 mm long, with short filaments, covered with fine papillate hairs. Achenes dark brown, to 5.5 mm long; pappus of thin, fragile, sordid- white or yellowish hairs; inner pappus hairs somewhat longer than outer, plumose almost up to tip, with filiform tips. Flowering July to September. (Plate VII, Fig. 1.) Rubbly slopes of ravines, from mid-montane to subalpine zone, mostly in clay shales Caucasus: Main Range (near Shtulu Pass, ra- vine of Arm-Khi River; near Shatil, Kuba District), Dagestan, Eastern Transcaucasia (former Nukhin District), Southern Transcaucasia (Tedzam ravine). Endemic. Described from Bekker’s specimens from Dagestan (between Kurakh and Kro). Type in Leningrad; isotype in Tbilisi. Note. This species differs from the southern Balkan species C. cynaroides (Lam.) Spreng. by the weakly incised leaves with bifid lobes and the shorter cusps of the outer involucral bracts. 54. C. lamyroides Tamamsch. in Zam. po Sist. 1 Geogr. Rast. Tbil. 22 (1961) 81.—Chamaepeuce macrantha Schrenk in Bull. Acad. Sc. Pétersb. X (1842) 354; Fisch. and Mey. Enum. pl. nov. II, 38; Ldb. Fl. Ross. II, 745. Perennial. Stem ascending, large, up to 40 cm high, branched above, with solitary capitula at apices of stem and long axillary branches, leafy, sulcate, more or less thinly grayish-tomentose. Cauline leaves linear-lanceolate, sessile, semiamplexicaul, with long auricles, acumi- nate, glabrous above, green, with yellowish-white venation, weakly arachnoid-hairy along veins, grayish-tomentose beneath; middle cauline leaves to 20 cm long, 2.5 cm wide, sinuately pinnately parted into deltoid-lanceolate, 2—3-parted segments, their lobes strongly tapered to stiff, yellowish spine 7 mm long, sparsely spiny-ciliate; lower cauline 139 137 leaves broadly lanceolately-elliptical, narrowed into short petiole, sub- acute, 28 cm long, to 7 cm wide, pinnately parted into approximate, roundish-oval, sharp-toothed usually indistinctly 5-parted lobes, with hard, short spines at tips of teeth; lamina lustrous with veins ligneous, thicked at leaf margin above, occasionally sparsely floccose, beneath densely white-tomentose; apical leaves strongly reduced, narrowly lin- early lanceolate, coarsely pinnately toothed with long spines, usually 1-2, uppermost leaves surrounding base of capitula, not surpassing capitula, recurved. Capitula solitary, in corymbose panicles, globose- ovate, with few divaricate, long, yellow cusps‘, of which lower usually deflexed downward. Involucre 2.5-3 cm in dia, 3—3.5 cm long, scatteredly floccose, grayish-arachnoid-hairy above; involucral bracts from outer broadly ovate to middle oblong-ovate, 5-10 mm long, abruptly narrowed into linearly deltoid-lanceolate tip, dorsally cari- nate, incurved spinescent cusp usually longer than broader part of bracts, in middle bracts almost 2 times as long, membranous below, yellowish, glabrous, with dark-brown spot above, carinate, long, floc- cose-arachnoid-hairy below cusp; inner bracts linearly lanceolate, to 2 cm long, narrowed to short cups. Corolla pink, to 22 mm long; limb scarcely shorter than narrow corolla tube, unequally 5-parted almost to middle. Achenes 5 mm long, stramineous, with long obtuse tubercle; pappus with few, thin, fragile hairs, sordid-white, inner hairs long, with denticulate tips. Flowering July to August. Rubbly slopes and river valleys.—Soviet Central Asia: Dzhungaria- Tarbagatai. General distribution: May be found in Dzhungaria- Kashgaria. Described from Dzhungarian Alatau. Type in Leningrad. Note. By many authors this species is described as a member of the genus Chamaepeuce allied with the southern Balkan species Chamaepeuce cynaroides (Lam.) Spreng., from which, however, it is well distinguished by the form of leaves. Section 4. Echenais (Cass.) Petrak in Beih. Bot. Centralbl. XXXV, II, 2-3 (1917) 252.—Echenais Cass. in Bull. Soc. Philom Paris (1818) 33; id. op. cit. (1820) 4; id. in Dict. Sc. nat. XIV (1819) 170, XXV (1822) 226; DC. Prodr. VI, 660.—Leaves narrow, oblong-lanceolate, weakly crisped-arachnoid-hairy, or densely so beneath. Capitula ovate or globose-ovate, medium-sized, basally with tuft of undeveloped long- spinose bracts. Involucral bracts glabrous, or scarious to tip, more or less fimbriate-ciliate along margin or outer ones leafy, greenish, along margin scaly, or more or less cristate-spiny-ciliate, somewhat longer than inner, apically with long, scarious, erose-toothed or fimbriate, appendage, or narrowed into long membranous point, often consider- ably longer than florets. Corolla tube shorter or almost as long as limb. Achenes compressed, narrow, not longer than 4.5 mm. 140 138 Type of section: C. echinus (MB.) Hand.-Mazz. Note. The genus Echenais Cass. was erected from the Caucasian species originally described as Carlina echinus MB. Cassini, after describing the genus Echenais, gives this species a new name, E. carlinoides Cass. Petrak broadened the circumscription of Echenais and treated it as a section, of the genus Cirsium, adding to it, besides the Asiatic ones, the North American species with scarious involucral bract, appendages. Leaving Echenais as an element of Cirsium, I con- sider it possible to refer to this section the group of Soviet Central Asian species of the C. semenovii Rgl. and Schm. alliance. Unlike typical Echenais, these species are distinguished by having the outer involucral bracts narrowed to long cusps. Besides the ones in Soviet Central Asia, species of this alliance with several characters transi- tional to typical Echenais are distributed in Afghanistan and the west- ern Himalayas. Thus, Soviet Central Asia with adjoining parts of West- ern Asia may be considered one of the centers of origin of the section. In the western part of Western Asia, Asia Minor, and the Caucasus, only one member of the section—C. echinus (MB.) Hand.-Mazz.—is found. 55. C. semenovii Rgl. and Schm. in Bull. Soc. Nat. Mosc. XL, 3 (1867) 161; O. and B. Fedtsch. Perech. Rast. Turk., IV, 285; Petrak in Oest. Bot. Zeitschr. LXI, 322 p. p.—Cnicus semenowii Winkl. in Tr. Peterb. Bot. Sada, IX, 2 (1886) 523—Chamaepeuce macrantha B. bracteata Rupr.-in Mém. Acad. Sc. Pétersb. XIV, 4 (1869) 56.—Ie.: Petrak, op. cit. 322, Abb. 2. Perennial. Rhizome thick, vertical, with numerous dark brown fi- brous roots; stems to 55—60 cm high, ribbed weakly sulcate, purple- dark brown below, yellowish above, weakly branched, leafy to tip, with apically clustered, more or less numerous, capitula. Leaves green above, more or less glabrous or with scattered, crisped, arachnoid hairs, glaucesent beneath, along midrib more densely crisped hairy, with scattered hairs along veins, oblong-linearly lanceolate, sinuately pinnately parted into deltoid oval, inequidentate lobes, with apical teeth strongly spinescent, bearing yellowish spines 3—6 mm long; basal and lower cauline leaves 15-17 cm long, 2—3.5 cm wide, narrowed into toothed and finely spinose petiole up to 8 mm long; other cauline leaves oblong, divergent, narrowed at base, sessile, with weakly decur- rent, narrow, semiamplexicaul auricles with spines 12 mm long; apical leaves reduced, not surpassing capitula; apical leaves at base of ca- pitula sometimes with reduced lamina, long-spinescent, shorter than involucre. Capitula numerous, on short peduncles, subsessile, in lax, short recemes, almost capitate inflorescence at stem tip, solitary at 142 139 apices of axillary branches, not surpassing leaves. Involucre ovate- roundish, at beginning of flowering 2—2.5 cm in dia, dark brown, in- cluding spines somewhat surpassing florets, 3.5 cm long, with recurved cusps at end of flowering, occasionally grayish-arachnoid-hairy; outer involucral bracts ovate, to 5 mm long, yellowish below, glabrous, brown, carinate, linearly lanceolate above with long, pointed yellowish spine, almost 2 times as long as broader part in outer bracts, inner ones gradually reduced, innermost broadly lanceolate, narrowed into short point. Florets pale pink, to 21 mm long; corolla tube narrow, up to 9 mm long. Achenes dark brown, 5 mm long; pappus up to 18 mm long, sordid-white, with thin hairs; inner hairs apically scabrous. Flowering July to August. Juniper to alpine zone, on river banks, screes and moraines.— Soviet Central Asia: Tien Shan. General distribution: Dzhungaria- Kashgaria. Described from Trans-Ili Alatau. Type in Leningrad. 56. C. sairamense (Winkl.) O. and B. Fedtsch. Perech. Rast. Turk. IV (1911) 286.—C. semenovii ssp. sairamense Petrak in Oest. Bot. Zeitschr. LXI (1911) 323.—Cnicus sairamensis Winkl. in Tr. Peterb. Bot. Sada. IX, 2 (1886) 522.—Ic.: Petrak, op. cit. 323, Abb. 3. Perennial. Stem ascending, 40-60 cm high, somewhat ribbed and sulcate, finely white-arachnoid-hairy, branched almost from base or middle, with elongated axillary branches, densely leafy, with solitary large capitula at apices of stem and branches, occasionally clustered in 3s, of which lower considerably smaller than terminal. Leaves grayish- green above, sparsely crisped-hairy, thinly grayish-tomentose-arach- noid-hairy beneath, with somewhat prominent whitish veins, sinuately pinnately parted into deltoid-ovate, 3—5-toothed lobes, teeth tapered to long, thin, whitish spines, up to 10-12 mm long, irregularly ciliate- spinulose on margin; basal and lower cauline leaves long, oblong, almost reaching base of capitula or surpassing them; middle and upper cauline leaves linearly elliptical or linearly lanceolate, strongly re- duced, lobate, leaves at base of capitula long-pinnate-spinose, usually not surpassing involucre. Capitula ovate-globose, terminal, 3.5—4 cm in dia, forming corymbose paniculate or narrowly racemose inflores- cence. Involucre weakly arachnoid-hairy with long, upward directed, subulate spines, considerably surpassing florets; outer bracts broadly lanceolate, greenish-brown, glabrous below, sparsely arachnoid-hairy above, narrowed into long, membranous subulate point, below finely ciliate and with more or less elongated, occasional spines at base; middle bracts ovate or oblong-ovate, to 10 mm long, 3.5 mm wide, glabrous beneath, carinate, sparsely arachnoid-hairy above, abruptly narrowed into linear-subulate, more or less attenuate tip, long-tapered 140 Plate VII. 141 1—Cirsium sinuatum (Trautv.) Boiss.; 2—C. coryletorum Kom. 143 141 into acute, yellowish spine, almost 3—5 times surpassing broader part of bracts; inner bracts linearly lanceolate, with very short, scaly, to 3 mm long point. Florets pale-pink, 22—23 mm long; corolla tube to 9 mm long; limb almost to middle unevenly 5-lobed. Achenes light brown, to 5 mm long, with long tubercle at apex; pappus sordid-white, 17-18 mm long, with inner hairs apically scabrous. Flowering July to August. Mountain meadows.—Soviet Central Asia: Dzhungaria-Tarbagatai, Tien Shan. General distribution: Dzhungaria-Kashgaria. Described from Dzhungaria (southern shores of Lake Sairam). Type in Leningrad. Note. It differs from C. semenovii Rgl. and Schm. by having the stem branched almost from the base, solitary capitula with the long- attenuated, erect cusps of the outer involucral bracts considerably sur- passing the florets, and long cauline leaves that usually surpass the capitula. The type of the species is characterized by shortened, densely leafy stems that are branched almost from the base and bearing numer- ous solitary capitula. The plants from Tien Shan with tall stems branched from the middle also belong to this species. 57. C. glabrifolium (Winkl.) O. and B. Fedtsch. Perech. Rast. Turk. IV (1911) 286.—C. glabrifolium Petrak in Oest. Bot. Zeitlschr. LXI (1911) 324.—C. glabrifolium var. spinosissimum O. and B. Fedtsch. op. cit. (1911) 286.—Cnicus glabrifolius Winkl. in Tr. Peterb. Bot. Sada, IX, 2 (1886) 523.—Cnicus glabrifolius var. spinosissimus Winkl. and var. hirtus Winkl. in schedis.—Ic.: Petrak, 1. c. Abb. 4. Perennial. Rhizome with numerous fibrous roots; stem ribbed, sulcate, ascending, 55—70 cm high, at places purple, branched from middle, sparsely crisped-hairy. Leaves oblong-lanceolate; cauline leaves sessile, auriculate, apically narrowed, long-acuminate, usually glabrous above, sparsely crisped-hairy along veins beneath, with prominent midrib and lateral veins, sinuately pinnately-lobed almost to middle, with longer lower lobes; segments 2—3-parted into deltoid-ovate, crenate-toothed lobes, teeth strongly tapered to hard spines 5-12 mm long; middle cauline leaves 20 cm long, 8 cm wide, reduced upward. Capitula terminal on stem and axillary branches, subsessile, 2—6 clus- tered in corymbose-paniculate inflorescence, 1—2 cm in dia at begin- ning of flowering, enlarged at fruiting, 3.0—3.5 cm in dia, basally with rosette of short, spinescent, pinnately incised, linear-lanceolate leaves, with tips almost as long as involucre or somewhat longer. Involucre subglabrous or weakly arachnoid-hairy; outer involucral bracts dark brown, glabrous, broadly lanceolate, narrowed from middle to scari- ous, more or less spinescent tip, erect or upward spreading, almost as long as inner bracts, ciliolate; middle and inner bracts linearly lan- ceolate, apically narrowed to lanceolate, scarious, yellowish or purple, 144 142 fimbriate, acuminate, slightly bent appendage; in var. spinosissiumum O. and B. Fedtsch., outer bracts abruptly narrowed to long cusp, with 1—2 thin spines on each side at base. Florets purple, to 15 mm long. Corolla unevenly 5-parted almost to middle; limb longer than narrow tube, tube to 9 cm long. Achenes 4 mm long, brown with dark streaks, apically obliquely truncate, with long tubercle; pappus sordid-white, inner hairs with longer, scabrous tips. Flowering July to August. (Plate VIII, Figs. 1, 2). Mountain gorges, on stony riverbanks, as roadside weed; mostly in mid-montane zone.—Soviet Central Asia: Dzhungaria-Tarbagatai, Tien Shan, Pamiro-Alai. General distribution: Dzhungaria-Kashgaria. Described from specimens from Dzhungaria and Zeravshan. Type in Leningrad. Note. Winkler cited two specimens under the name Cnicus glabrifolius Winkl. One of them was collected from the Talka Gorge in the vicinity of Lake Sairam in Dzhungaria and the other from the Zeravshan Basin. The first can be considered the type of C. glabrifolium (Winkl.) O. and B. Fedtsch. It was later described in detail by Petrak, who considered it a hybrid of C. semenovii and C. sieversii. The Zeravshan specimen is closer to C. sieversii (Fisch. and Mey.) Petrak. According to Petrak, it should be referred to the latter, but it differs considerably from C. sieversii in the outer involucral bracts, which are attenuate-cusped and lack the membranous-fimbriate border character- istic of C. sieversii. I consider it possible to treat both specimens cited by Winkler as C. glabrifolium (Winkl.) O. and B. Fedtsch. The hybrid origin of the type of C. glabrifolium seems doubtful to me because similar specimens varying with respect to the fringe of the inner bracts are observed over a large area from Dzhungaria to the Zeravshan Range. 58. C. badakhschanicum Charadze in Addenda XXVII, 604. Perennial. Stem to 1 m high, ascending, ribbed, scatteredly crisped- hairy below, densely lanate above with whitish, long-articulated, crisped hairs, branched above, with more or less elongated axillary branches. Cauline leaves glaucescent, scatteredly hairy above, more of less densely lanate beneath on midrib, middle ones to 10 cm long, lacking spines, to 3 cm wide, strongly reduced on floriferous branches, ovate-lan- ceolate or oblong-lanceolate; all leaves weakly, usually up to middle, sinuately pinnately lobed, with narrow sinuses, and broadly deltoid, 3— 5-parted segments; cauline leaves sessile, long-acuminate, strongly spinescent, auriculate, semiamplexicaul, unevenly lobate-toothed, with numerous approximate, yellowish, hard unequal spines, lobes of seg- ments tapered, to spines 12-13 mm long, sinuses with short, spines 2— 3 mm long. Floriferous branches flexuous, with 2—5 clustered, subsessile 143 145 capitula at apices; bracteal leaves at base of capitula many, small, lanceolate, pinnately toothed, strongly tapered to hard yellowish spines 8-10 mm long, almost as long as Involucre or slightly longer. Involu- cre glabrous, roundish-ovate, 2—3.5 cm in dia, divergent at end of fruiting, with more or less horizontal bracts; outer bracts ovate-lan- ceolate, narrowed from middle to long, thin spines, somewhat shorter than inner, erect, sparsely arachnoid-hairy along margin, divergent, yellowish spines to 4 mm long; middle bracts lanceolate, glabrous, narrowed slightly above middle to thin, spinescent, slightly deflexed cusp, scarious at base along margin, with plumosely divergent thin spines; inner bracts linearly lanceolate, narrowed to slightly broader, ’ scarious, undulate, slightly bent, acuminate tip. Florets pink, 17 mm long; corolla tube narrow, 7.5 mm long, limb lobed almost to middle. Achenes 4 mm long, grayish-white, with small dark streaks, compressed, slightly triquetrous, obliquely truncate apically, with tubercle; pappus yellowish-white, to 13 mm long, with very thin hairs, inner hairs sca- brous at tip. Flowering July to August. Mountain river valleys, in grassy areas, in alpine zone.—Soviet Central Asia: Pamiro-Alai (Badakhshan). General distribution: Prob- ably found in adjoining territory of Afghanistan. Described from Gunts River Valley from collections of O. and B. Fedtschenko. Type in Leningrad. Note. It is distinguished from the Afghan species C. argyracanthum DC. by having the outer involucral bracts attenuated to thin, basally pinnate-aculeate spines and the inner bracts apically scarious. From C. glabrifolium (Winkl.) O. and B. Fedtsch. it differs by the outer bracts having long-pinnate-aculeate margins and the inner hairs having a nar- row-membranous tip, as well as by the form of the leaves, which have long thin spines. Besides the collections of O. and B. Fedtschenko, in recent years, additional material of this species was made available by S. Ikonnikov. 59. C. alberti Rgl. and Schm. in Tr. Peterb. Bot. Sada, V, 2 (1880) 318; O. and B. Fedtsch. Perech. Rast. Turk. IV, 286; Petrak in Oest. Bot. Zeitschr. LXI, 321.—C. semenovii ssp. alberti Petrak in schedis.—Cnicus alberti Winkl. in schedis.—Ic.: Petrak, op. cit. 322, Abb. 2. Perennial. Stem ascending, 60—90 cm high, ribbed, branched from middle or almost from base, more or less densely covered with tiny crisped hairs, sparsely arachnoid-hairy, densely leafy. Leaves green above, with scattered crisped-arachnoid hairs, densely and grayish- arachnoid-hairy beneath; lower leaves oblong-lanceolate, narrowed toward base into winged, spinous-toothed petiole, apically attenuated, 146 147 144 22-27 cm long, 7 cm wide, sinuately-pinnately parted into broadly deltoid-ovate, 3—5-parted lobes, teeth on leaves strongly tapered to hard, thin spines 5-6 mm long, sinuses with distant spines 1-3 mm long; other leaves gradually reduced upward, lanceolate or narrowly lanceolate, sinuately pinnatifid with broadly ovate, sinuate-toothed lobes; middle leaves auriculate, and short-decurrent, winged, long- acuminate, pubescence like lower cauline leaves, with shorter spines at apices of teeth; apical leaves at base of capitula many, small, lan- ceolate, long-plumose, divergent, shorter than involucre. Capitula glo- bose-ovate, 1.5—2.0 cm in dia, 3—5-clustered at tips of stem and long, somewhat flexuous, axillary branches, occasionally solitary, subsessile. Involucre finely arachnoid-hairy, with erect and slightly divergent cusps; outer bracts membranous, ovate-oblong, glabrous, arachnoid-hairy along margin, brown above, carinate, more or less gradually narrowed from middle, attenuate into yellowish, subulate, slightly elongate spine, al- most as long as or shorter than inner bracts; inner bracts oblong-lan- ceolate to linearly lanceolate, membranous, with short cusp, at fruiting with falcately recurved apices. Florets 22 mm long; corolla tube nar- row, 9 mm long; limb lobed to 1/3. Achenes 4 mm long, yellowish, with dark brown streaks; pappus sordid-white, with few hairs, inner hairs not plumose to tip. Flowering July to August. Mid-montane to subalpine zone, in river valleys and along roads.— Soviet Central Asia: Tien Shan, Dzhungaria-Tarbagatai. General dis- tribution: Dzhungaria-Kashgaria. Described from Musart River Valley. Type in Leningrad. Note. In the work of Petrak (1911), the involucral bracts were not illustrated quite accurately. In the type of this species, the outer, es- pecially the middle, bracts are attenuated to long spines, almost as long as the inner involucral bracts. 60. C. sieversii (Fisch. and Mey.) Petrak in Oest. Bot. Zeitschr. LXI (1911) 324.—C. polyacanthum Kar. and Kir. in Bull. Soc. Nat. Mosc. XIV (1841) No. 517.—Echenais sieversii Fisch. and Mey. Enum. pl. nov. I (1841) 44; Kar. and Kir. in Bull. Soc. Nat. Mosc. XV, No. 498; Ldb. Fl. Ross. II, 747; O. and B. Fedtsch. Perech. Rast. Turk. IV, 288.—E. sieversii var. B. Ldb. Fl. Ross. II (1846) 747.—E. carlinoides O. and B. Fedtsch. op. cit. (1911) 288 p. p. non Cass.—Cnicus spinosissimum Sievers in schedis.—Ic.: Petrak op. cit. 325, Abb. 5. Perennial. Rhizome vertical, with ligneous fibrous root. Stem ribbed, to 1 m high, ascending, at places purple, with scattered crisped hairs. Leaves oblong-ovate to oblong-lanceolate, glabrous above or more or less scatteredly crisped-hairy, densely crisped-hairy beneath on veins; lower cauline leaves 21 cm long, 7 cm wide, pinnately parted 148 145 in oblong or oblong-deltoid, unevenly 2—3-parted lobes, teeth strongly tapered to spines 8—12 mm long, sinuses with solitary spines 2-7 mm long, narrowly winged, weakly decurrent; basal leaves 35 cm long, 12 cm wide, narrowed into petiole, pinnately parted almost to base into oblong, unequally pinnately lobed segments; upper cauline leaves broadly lanceolate, to 7 cm long, 3 cm wide, auriculate, sessile, semiamplexicaul, unevenly pinnately toothed, ciliate-spinose from long and short, hard spines. Capitula globose-ovate, 2—2.5 cm in dia, 3—7 clustered at tips of stem and its branches, subsessile; bracteal leaves at base of capitula few, sma!l, pinnately incised, long-spiny. Involucre glabrous; outer involucral bracts oblong-ovate, stramineous or brown- ish below, broadened above and membranous, whitish, fimbriate, spinescent; inner bracts linearly lanceolate, narrowed to membranous, slightly broadened tip, cuspidate. Florets pinkish-lilac; corolla 18.5 mm long, tube to 7 mm long, limb lobed to 1/3. Achenes whitish, to 4 mm long, with thin, longitudinal, dark streaks; pappus to 16 mm long, sordid-white, with few finely plumose hairs, inner hairs with long-scabrous tips. Flowering July to August. (Plate VIII, Fig. 3.) Alpine zone, in rubbly places——Western Siberia: Altai (Narym Range, at sources of Tau-Tykol); Soviet Central Asia: Dzhungaria- Tarbagatai, Tien Shan. General distribution: Dzhungaria-Kashgaria. Described from Tarbagatai. Type in Leningrad. Note. C. sieversii (Fisch. and Mey.) Petrak is allied with several species close to C. semenovii Rgl. and Schm. Among them, C. glabrifolium (Winkl.) O. and B. Fedtsch. stands out, with characters transitional to C. sieversii in the structure of involucral bracts. Petrak considers it possible to refer the specimen of the former species from Zeravshan to C. sieversii. However, the latter species is well distin- guished from the Zeravshan specimen by the outer involucral bracts, which are broadened upward and fimbriate, and also by the form of the leaves. The species Echenais sieversii Fisch. and Mey. and Cirsium polyacanthum Kar. and Kir. were published in 1841, but the Enumeratio of Fischer and Meyer came off the Press on June 15, 1841, whereas the first part of Karelin’s and Kirilov’s list appeared on July 2, and the second part in September (N.V. Pavlov—G.S. Karelin, 1948). 61. C. echinus (MB.) Hand.-Mazz. in Ann. Nat. Hofmus. Wien. XXIII (1909) 197; Petrak in Tr. Tifl. Bot. Sada, XII, 1, 25; in Vestn. Tifl. Bot. Sada, 24, 14; in Izv. Kavk. Muz. VIII, 1-2, 75; Grossh. FI. Kavk. IV, 186; Kolak. Fl. Abkhazii, IV, 274; Charadze in Fl. Gruzii, VIII, 510; in Fl. Azerb. VIII, 406.—C. scleranthum MB. taur.-cauc. III (1819) 559; C.A.M. Verzeichn. 69; Boiss. Fl. or. HI, 540.—C. 149 146 scleranthon Spreng. Syst. veg. 3 (1826) 378.—Carlina echinus MB. Beschreib. Land. Casp. Meer. (1800) 199; Fl. taur.-cauc. II, 283.— Echenais carlinoides Cass. in Dict. Sc. nat. XIV (1819) 171; DC. Prodr. VI, 661; Ldb. Fl. Ross. II, 747.— E. carlinoides B. nutans DC. Prodr. VI (1837) 661.—E. nutans Cass. in Dict. Sc. nat. XXV (1822) 226.—Ic.: Charadze in Fl. Azerb. VIII, Plate XL, 1.—Exs.: Petrak, Cirsioth. Univ. Nos. 93, 135, 136. Biennial. Rhizome ligneous, nodulose; stem 45—60 cm high, ribbed ascending, more or less sparsely branched from middle, somewhat densely arachnoid-hairy, densely leafy to tip, with numerous capitula clustered in paniculate inflorescence. Leaves green above, with yel- lowish midrib, scatteredly whitish-arachnoid-tomentose beneath, deeply sinuately pinnately incised to almost entire, toothed, with yellowish spines 10 mm long; basal leaves up to 12 cm long, 3 cm wide, oblong- lanceolate, sinuately-pinnately incised almost to base into deltoid ovate, 2—3-parted, tooth, long-spinescent lobes and spinulose petiole; cauline leaves oblong-lanceolate, sinuately pinnatifid into deltoid ovate lobes, up to 9 cm long, 2 cm wide, with broad weakly decurrent auricles; lower cauline leaves narrowed toward base, semiamplexicaul, acumi- nate; apical leaves reduced, spinescent, surrounding base of capitula and almost as long as capitula. Capitula numerous, solitary or in 2s, terminal on axillary or lateral branches, in short racemose inflores- cence, corymbose-paniculate at stem apices. Involucre roundish-ovate, 1-2 cm in dia, at base with few strongly reduced, long, plumosely- aculeate, bracteoles shorter than involucre, stramineous or whitish- yellow, glabrous, with numerous scales, attenuate into long, hard spines; outer bracts ovate to oblong-lanceolate, narrowed above, with green spot, transitional to broader, membranous ovate appendage, deeply fimbriate in outer and weakly fimbriate in middle bracts, tapered to long, yellowish, erect spine; inner bracts linearly lanceolate, broader, scarious, with toothed apex narrowed to point. Florets whitish or pale; corolla 19 mm long, tube narrow, to 9 mm long, limb lobed to deeper than middle. Achenes brownish-yellow, 4 mm long, with narrow bor- der and raised apical tubercle; pappus sordid-yellowish-white, 17 mm long, inner hairs with toothed tips. Flowering August to September. Mountain meadows and pastures, roadsides, sometimes as weed in crops and old fields, from foothills to subalpine zone.—Caucasus: throughout Caucasus. General distribution: Central Europe (Czecho- slovakia, introduced), Asia Minor, Armenia and Kurdistan, Iran and Afghanistan. Described from Eastern Caucasus. Type in Leningrad. Note. Only one member of the section Echenais is found in the Caucasus, Asia Minor and West Asia. This species is reported for Europe as an introduced plant. Evidently, it crosses with members of 150 147 the section Cirsium. The following are hybrids described from the Caucasus: 1. C. obvallatum (MB.) MB. xX C. echinus (MB.) Hand.-Mazz.— x C. lojkae Somm. and Lev. in Nuov. Giorn. Bot. Ital. N. S. II, 1 (1895) 15; in Tr. Peterb. Bot. Sada, XVI (1900) 259; Petrak in Tr. Tifl. Bot. Sada, XII, 1 (1912) 25.—Ic.: Somm. and Lev. 1. c. (1900) tab. XX VI; in Dechy, Kaukasus, III (1907) tab. XXIV.—Outer involu- cral bracts apically not broadened, membranous, short-cristate-ciliate with short apical spine; inner bracts with scaly deltoid ovate, erose- toothed, cuspidate appendage. Capitula medium-sized surrounded by narrowly plumosely toothed and spinescent, slightly surpassing apical leaves; florets 17-18 mm long, yellowish. Leaves pale yellow, finely arachnoid-hairy beneath. Subalpine zone.—Caucasus: Ciscaucasia (Ardon River Basin, Tsei Ravine), Eastern Transcaucasia (Gidauri and Georgian Military Road). Described from Central Caucasus from Lojka’s collections (Tsei Ra- vine). Type in Florence. 2. C. subinerme C.A.M. X C. echinus (MB.) Hand.-Mazz.—C. grossheimii Petrak (C. elodes x C. echinus) in Vestn. Tifl. Bot. Sada, Nov. Ser., 2 (1926) 43. In habit characters, especially the strongly branched stem, form of the leaves, and structure of the involucral bracts, this hybrid approaches C. echinus, from which, however, it differs by having broadly long- decurrent leaves and reddish florets, which resemble those of C. subinerme. In the shape of the outer involucral bracts, which are broad- ened above, this hybrid also differs somewhat from C. echinus. Caucasus: Southern Transcaucasia. Described from Gyunei shore of Lake Sevan from Shakhdag Mountain. Note. Petrak considers these plants as hybrids between C. elodes and C. echinus. Caucasus authors mistakenly referred C. subinerme Fisch. and Mey. to C. elodes MB. Section 5. Cirsium.—Sect. Onotrophe (Cass.) DC. Prodr. VI (1837) 644; Nakai in Tokyo Bot. Mag. XXVI (1912) 357; Petrak in Beih. Bot. Centralbl. XXXV (1917) 254; Kitam. in Acta Phytot. et Geobot. III, 1 (1934) 2; Comp. Japon. I (1937) 40.—Onotrophe Cass. in Dict. Sc. nat. XXXVI (1825) 145.—Onotrophe sect. Apalocentron Cass. ibid. 146.—Onotrophe sect. Microcentron Cass. ibid.—Sect. Chamaeleon DC. in Duby, Bot. Gall. I (1828) 287; Koch. Syn. (1837) 393; Boiss. Fl. or. III, 524, 539; Nakai in Tokyo Bot. Mag. XXVI (1912) 357, pro subgen.—Sect. Erysithales DC. in Duby, 1. c. 285.— 15 —_ 148 Sect. Acarna DC. in Duby, 1. c. 286 p. p.—Sect. Orthocentrum DC. Prodr. VI (1837) 641.—Orthocentron Cass. in Dict. Sc. nat. XXXVI (1825) 480, pro gen.—Sect. Apalocentron (Cass.) Kitam. in Acta Phytot. et Geobot. XII, 2 (1943) 102.—Leaves glabrous above, thinly or densely arachnoid-hairy, occasionally scabrous from short hairs. Capitula medium- or small-sized or small, occasionally large, globose-ovate or globose. Involucre more or less imbricate, glabrous or arachnoid-hairy; involucral bracts foliaceous; occasionally coriaceous, undivided or spinose-ciliate, occasionally cristate-puberulent, short- or somewhat long-spinescent, sometimes apically broadened to more or less scari- ous, erose-toothed appendage. Florets red or yellowish, less often whitish; corolla tube narrow, almost as long as or shorter than ligule, latter unevenly 5-parted almost to middle; filaments pubescent or with occasional, short, sometimes papillate hairs. Achenes 3-5 mm long; pappus of few, fine plumose hairs, with scabrous or filiform tips. The species of this section are widely distributed in Europe, Asia, and North America. Type of section: type of genus. Subsection 1. Montana Charadze in Addenda XXVII, 604.— Leaves broad, not decurrent, undivided or pinnately lobed; cauline leaves usually sessile, semiamplexicaul or narrowed into petiole, subglabrous, scatteredly hairy or grayish-tomentose beneath. Involucral bracts gradually longer from outer to inner, outer sometimes almost 1/2 as long as inner, occasionally somewhat broadened in upper part, scarious on margin. Capitula few, more or less clustered at apices of stems, occasionally solitary. Perennial large herbs growing in the montane regions of Eastern Europe, Balkans, Caucasus, and Asia Minor. Type of subsection: C. pseudopersonata Boiss. and Bal. 62. C. kusnetzowianum Somm. and Lev. in Tr. Peterb. Bot. Sada, XII, 1 (1892) 155; in Nuov. Giorn. Bot. Ital. N. S. II, 1 (1895) 12; in Tr. Peterb. Bot. Sada, XVI (1900) 261; Petrak in Tr. Tifl. Bot. Sada, XII, 1, 28; Grossh. Fl. Kavk. IV, 189; Kolak. Fl. Abkhazii, IV, 275.— Charadze in Fl. Gruzii, VIII, 481.—C. kusnetzowianum var. polycephalum Alb. Prodr. Fl. Colch. (1895) 144; Grossh. Fl. Kavk. IV, 189.—C. kusnetzowianum var. oblongifolium Alb. loc. cit. Somm. and Lev. op. cit. (1900) 262; Grossh. Fl. Kavk. IV, 189.—C. pseudopersonata ssp. kusnetzowianum (Somm. and Lev.) Petrak in Mitteil. Thiiring. Bot. Gesellsch. II, 1 (1960) 33.—Ic.: Somm. and Lev. in Tr. Peterb. Bot. Sada, XVI, Plate XXVII—Exs.: Petrak, Cirsioth. Univ. No. 130. 152 149 Perennial. Stem up to 1.5 cm high, ascending, thinly arachnoid- hairy, sparsely crisped-hairy, sulcate-ribbed, leafy, with short branches above. Leaves broad, ovate-elliptical, with short-papillate hairs above, more or less pubescent on veins beneath with crisped arachnoid hairs, unevenly more or less coarsely toothed, teeth short-spinescent, some- times leaves glabrous on both sides, coarsely toothed (var. oblongifolium Alb.), short-acuminate; lower cauline leaves abruptly narrowed in broadly winged, lobate petiole, to 6 cm long, including petiole 30 cm long, 13 cm wide; middle and upper cauline leaves subcordate, sessile, semiamplexicaul, reduced toward stem apes, 7-23 cm long, 4-9 cm wide; apical leaves 2—3, broadly or linearly lanceolate, surpassing ca- pitula. Capitula globose-ovate, to 2 cm in dia, solitary, terminal on stem and short branches, or 3—6 in corymbose inflorescence, occasion- ally capitula numerous (var. polycephalum Alb.). Involucre glabrous or scatteredly arachnoid-hairy; inner involucral bracts elongated-lan- ceolate, tapered to thin, yellowish spine, brownish-green below, dark reddish-brown above, weakly carinate, ciliolate; inner bracts linearly lanceolate, with scarious tip. Florets purple-red; corolla 17-19 mm long with narrow tube to 5—7 mm long, limb parted to middle. Achenes 4 mm long, 1.5 mm wide, yellowish with dark brown streaks; pappus sordid-white, 14 mm long, of few unequal hairs, with apically sca- brous inner hairs. Flowering July to September. Forest and subalpine zones, on river terraces, usually in tall herbs, often along forest edges.—Caucasus: Ciscaucasia (western part), East- ern Transcaucasia (Bakuriani), Western Transcaucasia, Southern Transcaucasia (Abastumani District). General distribution: Asia Mi- nor (former Artvin District). Described from two specimens from Abkhazia (upper reaches of Klych River, Dzhodisvik Range). Type in Florence. Note. Recently, Petrak (1960) treated this species as a subspecies of C. pseudopersonata Boiss. and Bal. from Asia Minor, distinguish- ing the Western Transcaucasian form primarily by the pubescence of the leaves. We consider it better to treat S. kusnetzowianum as a sepa- rate species in the series of species close to C. pseudopersonata. This entire series of species differs by its very broad, nondecurrnt leaves and structure of the involucral bracts from C. uliginosum (MB.) MB.., to which Petrak relates this species. Petrak described the hybrid C. obvallatum (MB.) MB. x C. kusnetzowianum Somm. and Lev.—C. x prativagum Petrak in Vestn. Tifl. Bot. Sada, XIX (1911) 4. Alpine meadows.—Caucasus: Western Transcaucasia (Adzharia). It differs from C. obvallatum by having an almost glabrous stem, broader sinuate-pinnate-toothed leaves with spinulose margins, Ss} 150 solitary, the apical capitula surrounded by a few shorter and less densely short-spinose bracteal leaves. C. kusnetzowianum differs from the hybrid by having broader and less coarsely toothed leaves and 1-3 long apical leaves with spinulose margins. 63. C. sychnosanthum Petrak in Izv. Kavk. Muz. VIII, 1-2 (1914) 63; Grossh. Fl. Kavk. IV, 189; Kolak. Fl. Abkhazii, IV, 274; Charadze in Fl. Gruzi, VIII, 481. Perennial. Stem ascending, 1—2 m high, sulcate-ribbed, finely arach- noid-hairy, dark reddish-brown, leafy, with short cymose branches near stem apex. Cauline leaves subglabrous on both sides, weakly arach- noid-hairy beneath on veins, oblong-ovate or ovate-elliptical, pinnati- fid or pinnately parted into remote broadly deltoid ovate and short- acuminate, sometimes apically bifid, coarsely toothed segments, with teeth strongly spinescent, serrulate-spinose between teeth; lower cauline leaves narrowed to short, broadly-winged, spatulate petiole, including petiole 37 cm long, 17 cm wide; middle and upper cauline leaves cordate or auriculate, narrowed above, 12—22 cm long, 6—12 cm wide; apical leaves 1—3, shorter than or slightly exceeding capitula, broadly lanceolate to lanceolate, pinnately lobed, spinulose. Capitula terminal on stem and branches, solitary or in 2s, slightly projected above apical leaves, usually 3—S in corymbose-paniculate inflorescence, subsessile, globose-ovate, 20-24 mm in dia. Involucre glabrous, brownish-red- dish-brown, imbricate; bracts lanceolate, strongly tapered to short- spines, yellowish below, brownish-reddish-brown above; inner bracts linearly lanceolate, narrowed into thin scarious tip. Florets purple-red, to 20 mm long; corolla tube narrow, to 8 mm long, parted to deeper than middle. Achenes to 4 mm long; pappus 16 mm long, sordid-white, basally yellowish, with scabrous hair tips. Flowering July to August. Forest to alpine zone, along streams often as constituent of tall- herb vegetation, now and then forming dense thickets.—Caucasus: Western Ciscaucasia (basin of Belaya [white] River), Western Transcaucasia (Lower Svanetia-Lentakhi, Abkhazia, Mzymta River basin). Endemic. Described from Abkhazia. Type in Tbilisi. Note. In its broad leaves and form of the involucral bracts this species approaches C. kusnetzowianum Somm. and Lev. Petrak com- pares it with C. svaneticum Somm. and Lev. and C. uliginosum (MB.) MB.; however, C. sychnosanthum has almost nothing in common with them. 64. C. oblongifolium C. Koch in Linnaea, XVII (1843) 41; Boiss. Fl. or. III, 539; Alb. Prodr. Fl. Colch. 143; Petrak in Izv. Kavk. Muz. 151 VIII, 1-2, 74; Grossh.: Fl. Kavk. IV, 186; Kolak. Fl. Abkhazii, IV, 273; Charadze in Fl. Gruzii, VIII, 482.—C. uliginosum B. glabrum Ldb. Fl. Ross. II (1846) 736. Perennial. Stem up to 1.5 m high, ascending, ribbed sulcate, simple or cymosely branched above, scatteredly hairy. Cauline leaves oblong or oblong-ovate, sessile, somewhat cordate, semiamplexicaul, subacute, on both sides glabrous, glaucesent beneath, shallow and sinuately pin- nately toothed with simple or bifid unequal teeth, teeth deltoid ovate, tapered to more or less hard spines 2.5 mm long, sinuses spinulose; lower cauline leaves to 27 cm long, 9 cm wide, internodes short; middle cauline leaves lanceolate, to 25 cm long, 7 cm wide, reduced upward, 12 cm long, 4 cm wide; apical leaves 1—4, small, linearly lanceolate, pinnate-long-spinose. Capitula globose-ovate, 1.5—2.5 cm in dia, 2-9 clustered at stem tip, subsessile, on short branches. Involucre subglabrous or weakly arachnoid-hairy, purple; outer involucral bracts yellowish below, purple above, with scarious, scarcely broadened, finely toothed margin, spinulose; inner bracts linearly lanceolate, with scari- ous, purple, appendicular, slightly bent tip. Florets purple-red, 18 mm long; corolla with narrow tube to 6 mm long; limb parted to middle. Achenes 3 mm long, reddish-brown to brown, basally narrowed; pap- pus 14 mm long, sordid-white, basally reddish-brown with thin hairs, short plumose almost to tip. Flowering July to August. Forest and subalpine zones, in wet meadows, on riverbed gravels and in tall-herb vegetation, often forming dense thickets.—Caucasus: Eastern Transcaucasia (upper reaches of Liakhvi River), Western Transcaucasia (Gornaya Racha, upper reaches of Rioni River, Abkhazia- Agurepsta). Endemic. Described from Racha. Type was in Berlin; isotype in Leningrad. 65. C. waldsteinii Rouy, Fl. Fr. IX (1905) 84; Dostal, Kvet. CSR (1950) 1668; Hermann, FI. Nord. u. Mitt—Eur. 1073.—C. pauciflorum (Waldst. and Kit.) Spreng. Syst. veg. 3 (1826) 375, non Lam. (1778); DC. Prodr. VI, 649; Boiss. Fl. or. III, 545; Koch, Syn. ed. 3, II (1902) 1544; Klok. in Vizn. Rosl. URSR, 569; Hegi, Ill, Fl. VI, 2, 894.— Cnicus pauciflorus Waldst. and Kit. in Willd. Sp. pl. II, 3 (1803) 1677; Waldst. and Kit. Desr. et icon. pl. rar. Hung. II, 175.—Ic.: Waldst. and Kit. 1. c. tab. 161; Rchb. Ic. Fl. Germ. XV, tab. 833; Javorka and Csapody, Iconogr. Fl. Hung. 543, fig. 3883; Hegi, Ill. FI. VI, 2, 894, fig. 596.—Exs.: Herb. norm. No. 2650; Fl. Stir. exs. No. 692. Perennial. Rhizome thick, strongly branched; stem 75—80 cm high, ascending, sulcate, reddish-brown, thinly and scatteredly arachnoid- 154 hairy, leafy, with large, broad leaves, with elongate inflorescence at 155 152 apex comprising 3—4 capitula, densely arachnoid-hairy below capitula. Basal leaves with appressed, scattered, crisped hairs above, lanate along veins beneath, sinuate-crenate-toothed, deeply pinnatisect in lower half, with remote roundish-ovate lobes, abruptly narrowed into long, winged petiole, petiole a half as long as lamina, including petiole 55 cm long, 15 cm wide, ciliate from thin, yellow spines to 3 mm long;- cauline leaves usually thinly grayish-arachnoid beneath, appressed and scatteredly crisped hairy above, roundish-ovate, weakly parted into deltoid-ovate subacute lobes, toward base more or less weakly nar- rowed, auriculate, semiamplexicaul, lower ones to 26 cm long, 18-20 cm wide; middle cauline leaves pinnately lobed, with ovate, subacute segments having sinuate-pinnately lobed, spinulose-ciliolate, deltoid- ovate, subacute lobes, abruptly narrowed into broadly winged short petiole, with large auricles, including petiole 20-27 cm long, 14-16 cm wide; upper leaves roundish-ovate or ovate-lanceolate, unevenly pinnately toothed, latter narrowed into short spines, acuminate, sessile, semiamplexicaul, arachnoid-hairy, almost tomentose below, 6—9.5 cm long, 3.5—4.5 cm wide. Capitula basally with one apical lanceolate, declinate, almost as long as inflorescence or slightly shorter; capitula usually 3—4, clustered at stem tip, subsessile or on short peduncles, apical capitula 2—3 cm in dia, lower ones usually smaller than upper. Outer involucral bracts coriaceous, very tiny appressed-hairy (seen under powerful hand lens), reddish-brown to brown, dark brown along midrib, ovate to ovate-lanceolate, 7-10 mm long, gradually narrowed to thin, short spine; inner bracts membranous, light brown, lustrous, lanceolate, apically narrowed into scarious, slightly reddish-brown to brown tip. Florets dark purple, to 19 mm long, with narrow corolla tube to 7 mm long; limb unequally parted. Achenes dark-reddish-brown, with apical tubercle; pappus light reddish-brown, to 18 mm long, apically with slightly racemelike broadened, toothed, darker reddish- brown inner hairs. Flowering July to September. Subalpine zone, among scrubs, on talus.—European Part: Upper Dniester, Bessarabia. General distribution: Central Europe (Eastern Alps and Eastern Carpathians), Balkans. Described from Carpathians. Type in Budapest. 66. C. hypoleucum DC. Prodr. VI (1837) 645; Boiss. Fl. or. III, 544; Petrak in Tr. Tifl. Bot. Sada, XII, 1, 28, 42; in Vestn. Tifl. Bot. Sada, 24, 12; Grossh. Fl. Kavk. IV, 187; Charadze in Fl. Gruzii, VIII, 483.—C. hypoleucum 8. lazicum C. Koch in Linnaea, XXIV (1851) 409.—C. hypoleucum var. ponticum Somm. and Lev. in Nuov. Giorn. Bot. Ital. N. S. II, 1 (1895) 18; in Tr. Peterb. Bot. Sada, XVI, 264; Grossh. Fl. Kavk. IV, 187.—C. hypoleucum ssp. drymeium Petrak in 156 153 Tr. Tifl. Bot. Sada, XII, 1 (1912) 43; Grossh. Fl. Kavk. IV, 187.—Ie.: Dmitrieva, Opred. Rast. Adzh. Fig. 32.—Exs.: Petrak, Cirsioth. Univ. No. 94. Perennial. Rhizome thick, woody; stem ascending, 1.5 cm high, leafy, sulcate, purple, branched almost from middle, with more or less leafless long branches, scatteredly arachnoid-hairy, densely below capitula. Leaves more or less densely covered above with small papil- liform and. crisped segmented hairs, white-tomentose beneath, soft, pinnately parted or sinuately pinnately toothed, spinulose; lower cauline leaves large, oblong-ovate, deeply pinnately incised into ovate or oblong-ovate, spinose-unequally toothed, obtuse lobes, basally more or less gradually narrowed into broadly winged, sinuate, spinose, spatu- late, more or less short petiole, including petiole 22 cm long, 8 cm wide, auriculate, subacute; middle cauline leaves oblong-ovate, weakly lobate, with roundish, sharp-toothed, long and thinly spinose lobes, auriculate, semiamplexicaul, 16—18 cm long, 6—9 cm wide; upper leaves on long branches 2—3.5 cm long, lanceolate, acute, spinose-toothed, remote; apical leaf somewhat remote from capitula, linear, bent, shorter than involucre. Capitula on long, almost leafless branches, globose- ovate, solitary terminal, 1.5—2.5 cm in dia, occasionally 2 or many, clustered (ssp. drymenium Petrak). Involucre purple, finely arachnoid- hairy; outer bracts appressed, deltoid-ovate, tapered to yellowish, short- spine, brownish-green, purple above; inner bracts linearly lanceolate, narrowed to purple, scarious tip, almost 2 times as long as outer, bent at flowering. Florets purple, 18 mm long, somewhat surpassing inner involucral bracts; corolla tube narrow, 5 mm long; limb parted up to 1/3. Achenes stramineous, 4 mm long; pappus 13 mm long, whitish, of few thin hairs with scabrous, scarcely thickened tips. Flowering July to August. Forest and subalpine zones, often among spruce-fir forests and as component of tall-herb vegetation —Caucasus: Eastern Transcaucasia (Borzhomo-Bakuriani District), Western Transcaucasia (Adzharo- Imeretian Range), Southern Transcaucasia (Abastumani). General dis- tribution: Balkans-Asia Minor. Described from Bithynian Olympus in Asia Minor. Type in Geneva; Topotype in Leningrad. Note. The species varies in the size of the spines, form of the involucral bracts, and texture of the leaves. The type of the species (described from the collections of Osh) is characterized by more co- riaceous leaves with hard spines along the margin. Perhaps the Caucasus-Lazistan form, recognized by Sommier and Levier as var. ponticum Somm. and Lev., should be treated as a separate species that replaces C. hypoleucum DC. in the east. 154 Hybrids have been described from the Caucasus and adjoining regions of Asia Minor between the following species: C. hypoleucum DC., C. obvallatum (MB.) MB., and C. echinus (MB.) Hand.-Mazz. 1. C. hypoleucum DC. x C. obvallatum (MB.) MB. f. woronowii Petrak in /zv. Kavk. Muz. VIII, 1-2 (1914) 77.— Cirsium x woronowii Petrak in Vestn. Tifl. Bot. Sada, 19 (1911) 19.—Exs.: Petrak, Cirsioth. Univ. No. 133; Herb. FI. cauc. No. 548. Described from Adzharia from Yu.N. Voronov’s collections. Later reported for Bakuriani. It differs from C. hypoleucum in the following characters: basally narrowed and deeply sinuate-pinnately incised leaves, loosely white- tomentose beneath; solitary, subsessile capitula terminal on elongate branches, in axils of linear-lanceolate, pectinately ciliate-spiny apical leaves; outer involucral bracts with scattered arachnoid-hairs and acumi- nate spines to 2 mm long; yellowish florets with pink anthers. C. obvallatum differs from the hybrid by more sinuately and deeply in- cised large leaves, which reach stem tip; capitula in clusters of several, very densely ciliate-spiny floral leaves and involucral bracts with dark- colored resinous band. 2. C. hypoleucum DC. xX C. obvallatum (MB.) MB. f. subhypoleucum Petrak in Izv. Kavk. Muz. VIII, 1-2 (1914) 78. Described from Kokhta Mountain in the vicinity of Bakuriani where it grows together with related species and f. woronowii Petrak. The latter is closer in habit to C. obvallatum, but f. subhypoleucum Petrak is closer to C. hypoleucum, differing by deeply sinuate-repand pinnate leaves and outer involucral bracts. 3. C. hypoleucum DC. x C. echinus (MB.) Hand.-Mazz.—C. x rollowii Petrak and Woron. ex Petrak in Vestn. Tifl. Bot. Sada, 24 (1912) 12. Described from Eastern Anatolia from the former Artvin District (between Salalet and Khinzort). It may be found in Adzharia. It is similar to C. hypoleucum DC. in the broad middle cauline leaves, tomentose hairs beneath, and form of the capitula and florets, but differs by having upper cauline leaves with teeth attenuated into long, thin, yellowish spines, long-spiny, small floral leaves, and outer involucral bracts, with spiny-ciliate margins and ending in yellowish spines to 2 mm long. The color of the corolla varies from yellowish to dirty-purple. Subsection 2. Sinocirsium Kitam. in Acta Phytot. et Geobot. III, 1 (1934) 3; Comp. Japon. I (1937) 61.—Rhizome with numerous fi- 157 brous roots. Basal leaves rosulate, persisting after flowering; cauline 15 [e.<) is3 leaves semiamplexicaul, not decurrent. Capitula medium-sized, erect or nodding; involucral bracts narrow, imbricate, straight, smooth, dor- sally with more or less bulged, stramineous dark bend, short-spinescent. Florets with narrow corolla tube, almost as long as limb, sometimes shorter or slightly longer. Achenes 3—4 mm long. Type of subsection: C. japonicum DC. Note. An Eastern Asian subsection, whose species are distributed in northeastern China, the Soviet Far East, Korea, and the islands of Japan. Kitamura segregated this group of species as a subsection, subordinate to the boreal section Onotrophe (Cass.) DC. (= Cirsium). The Okhotsk-Beringian species group of the subsection Borealicola Kitam. s. str. are closest to the subsection Sinocirsium in the structure of the achenes and florets. Both subsections include old, relatively mesophilous series of species, whose members are constituents of forest cenoses, coastal meadow vegetation, and large herb communities. Series 1. Japonica Kitam. in Acta Phytot. et Geobot. III, 1 (1934) 4; Comp. Japon. I (1937) 62.—Capitula at flowering erect; outer in- volucral bracts long, densely covered outside with dark glandular hairs. 67. C. maackii Maxim. Prim. fl. amur. (1859) 172; Trautv. in Tr. Peterb. Bot. Sada, VIII, 2, 503; Kom. Fl. Man’chzh. III, 750; Kom. and Alis. Opred. Rast. Dal’nevost. Kr. II, 1085; Kitam. Comp. Japon. I, 62.—C. littorale var. ussuriense Rgl. Tent. Fl. Ussur. (1861) 102; Herder in Bull: Soc. Nat. Mosc. XLIII, 1, 85——C. japonicum ssp. maackii (Maxim.) Nakai in Tokyo Bot. Mag. XXV (1911) 61.—C. japonicum var. amurense Kitam. Cirs. Nov. Orient.-Asiat. (1931) 12.— Cnicus japonicus €. maackii Maxim. in Bull. Acad. Sc. Pétersb. XIX (1874) 503 and in Mél. Biol. IX, 326; Palib. Consp. Fl. Cor. I, 119.— Ic.: Kitam. Comp. Japon. I (1937) pl. XII, fig. 3. ; Perennial. Stem ascending, 50-100 cm high, more or less pubes- cent, especially below, hairs crisped, dark brown, more or less scatteredly floccose-arachnoid-hairy above with dark brown hairs. Basal leaves oblong, narrowed to winged, spiny-ciliate petiole, petiole shorter than lamina, together 19-20 cm long; lamina 4—6 cm wide, sinuately pinnately parted, with broadly-ovate, often 2-parted, unevenly repand- toothed segments with teeth abruptly narrowed to small spines, vena- tion conspicuous, more or less crisped-hairy beneath along midrib, more or less thinly arachnoid-hairy, subglabrous above; cauline leaves oblong elliptical, to 17 cm long, 8 cm wide, incised almost to base into elongate-ovate sinuate, bifid segments, retusely pinnately toothed, with lanceolate or deltoid-ovate subacute lobes, teeth tapered to thin spines 2—4 mm long; lower leaves apically attenuate, basally narrowed into long, narrow-winged spinose petiole; middle and upper cauline leaves 156 sessile, auriculate; all leaves with appressed spines along margin, with teeth tapered to spines to 3 mm long; upper leaves basally broadened, semiamplexicaul, 9-10 cm long, shallowly pinnatifid, more or less grayish-arachnoid-hairy beneath; apical leaves on more less short, arach- noid-hairy petioles, linear, uppermost to 2 cm long, long-ciliate-spiny, reaching to base or middle of capitulum. Capitula solitary or 2, on long, slightly curved, tomentose-hairy peduncles, terminal or axillary, few, erect at flowering, 2—3 cm in dia. Involucre dark colored, more or less weakly arachnoid-hairy; involucral bracts lax, lanceolate, spinescent, later weakly curved; outer bracts dorsally bulged with black- purple band, densely covered with blackish glandular hairs; inner bracts narrow, linearly lanceolate, with scarious tips. Florets violet-purple, 21-23 mm long; corolla tube narrow, 12 mm long; limb lobed to 2/3. Achenes 3.5 mm long, light greenish, weakly triquetrous, with filiform elongate disk, basally slightly narrowed; pappus 18.5—25 mm long, whitish, basally sordid-white, with pointed tips of inner hairs. Flower- ing July to August. Dry meadows and forested mountain slopes, sometimes in old fields.—Far East: Uda River area, Ussuri. General distribution: Ko- rean Peninsula, China (northern). Described from mouth of Sungari River. Type in Leningrad. Note. Apparently, it is a polytypic species for which different varieties were described in Korea and Japan, mainly by Nakai (1912*). In Ohwi’s Manual (F7. Jap., 1953) these varieties are referred to C. japonicum DC. C. maackii was combined with the latter by Maximowicz in his later work (1874) as Cnicus japonicus €. maackii Maxim. Ac- cording to Kitamura (1934,** 1937), both species can be treated in one series of Manchurian-Japanese species. C. Maackii differs from C. japonicus DC. by having softer leaves that blacken on drying, and densely glandular outer involucral bracts. Series 2. Schantarensia Kitam. in Acta Phytot. et Geobot. III, 1 (1934) 4; Comp. Japon. I (1937) 71.—Capitula nodding; involucral bracts glandular. Species of this series are widely distributed in Eastern Asia; only one species grows in the Soviet Union. 68. C. schantarense Trautv. and Mey. Fl. Ochot. I, 2 (1856) 58; Kom. Fl. Man’chzh. III, 751; Nakai, Fl. Koreana II, 47; Kom. and 159 Alis. Opred. Rast. Dal’nevost. Kr. II, 1085; Kitam. Comp. Japon. I, *1911 cited in synonymy above.—Sci. Editors. **1931 cited in synonymy above.—Sci. Editors. 16 i) 157 76.—C. pendulum B. oligocephalum Rgl. and Till. Fl. Ajan. (1859) 107.—C. littorale Maxim. Prim. fl. amur. (1859) 173; Herder in Bull. Soc. Nat. Mosc. XLIII, 1, 85, excl. €. ussuriense Rgl.—C. littorale 6. nudum Rgl. Tent. Fl. Ussur. (1861) 95.—Cnicus japonicus €. schantarensis (Trautv. and Mey.) Maxim. in Bull. Acad. Sc. Pétersb. XIX (1874) 498 and in Mel. Biol. IX, 326.—Ic.: Kitam. Comp. Japon. I (1937) pl. XVI, fig. 2. Perennial. Rhizome short, oblique, thick, with numerous ligneous fibrous roots; stem bases with dark reddish-brown remnants of old leaves. Stems yellowish, ascending, to 70 cm high, sparsely crisped- hairy, with articulate hairs or weakly arachnoid-hairy, sulcate. Leaves in rosettes, on long, weakly winged petioles, broadly ovate, 13-25 cm long, 5.5—9.5 cm wide, obtuse, or acute, basally narrowed to long petiole not longer than lamina, subglabrous above with scattered simple hairs, more or less sparsely arachnoid-hairy beneath, with articulate, crisped hairs along veins, shallow repandly pectinately-toothed, with distant, fine, and tiny spine-like cilia; cauline leaves few, deflected; lower and upper cauline leaves identical in shape, oblong-ovate, deeply pinnately incised to in oblong and broadly deltoid-lanceolate segments, usually 15-18 cm long, 7.5—10.5 cm wide, lower narrowed to short petiole, others semiamplexicaul, more or less glabrous above, scatteredly pilose to finely grayish-arachnoid-hairy beneath, segments sharp- toothed, with spinescent teeth and small, appressed spine-like cilia on margin; apical leaves ovate or ovate-lanceolate, gradually narrowed above to broadly-oblong-lanceolate tip 6-7 cm long. Capitula 1—4, on long tomentose peduncles, nodding from base, with 1—2 very small, more or less linear, apical leaves. Involucre wide open, with more or less horizontal, lower involucral bracts, imbricate, broadly lanceolate, gradually becoming longer from outer to inner ones, outer ones tapered to dark spine, dorsally reddish-brown, thinly hairy, thinly arachnoid- hairy along margin, with broad blackish-purple band above, usually glandular along midrib; inner bracts lanceolate, carinate, with scarious bent tip. Florets dark purple-violet, to 21 mm long; corolla tube to 7 mm long; limb 2 times as long as tube, lobed to 1/3. Achenes yellowish, to 4 mm long, longitudinally weakly striate, with fine narrow tubercle at apex; pappus to 20 mm long, light reddish-brown, tips of inner hairs not plumose. Flowering July to September. Mixed and coniferous montane forests, also in deciduous forests, along riverbanks.—Far East: Uda River area (Bolshoi Shantar and Feklistov islands, Udskoe, Uda Range), Ussuri (basin of Suifun River, in vicinity of Vladivostok), Sakhalin. General distribution: China (northeastern). Described from Shantar Island. Type and paratype in Leningrad. 158 Note. The authors based the description on two of Middendorf’s specimens. One of them was collected on Shantar Island and the other in the vicinity of Udskoe. The same species was later described by Maximowicz as C. littorale Maxim. but in a broad sense. The variant of C. pendulum Fisch. described by Regel and Tilling from Ayan was rightly combined by Herder with C. schantarense Trautv. Mey. but, interpreting the species broadly, Herder, in his monographic treatment, created a complex system of subordinate units in C. littorale Maxim., combining under this name C. maackii Maxim. and C. schantarense Trautv. and Mey. Apparently, these varieties belong to the latter spe- cies: &. genuinum Maxim. B. asperum Maxim., y. schrenkii Maxim., 6. nudum Rgl. and €. setosum Herd. Maximowicz (1874), in his monographic treatment of the genus Cnicus auct. (Cirsium), included the name Cirsium littorale among the synonyms of Cnicus japonicus (DC.) Maxim., considering C. schantarense as part of the latter species. V.L. Komarov noted considerable differences of C. schantarense from C. japonicum. Kitamura (1934), having paid attention to the capitula of C. schantarense that nod from the base, placed it in the separate series Schantarensia Kitam. as distinct from C. maackii Maxim. and C. japonicum DC., which were referred to the series Japonica Kitam. According to Kitamura, the two series constitute the subsection Sinocirsium of the boreal section Onotrophe (= Cirsium). C. schantarense varies considerably in pubescence, incision, and texture of the leaves. For C. littorale, Maximowicz named forms differing mainly in pubescence: a. genuinum—with leaves grayish-pubescent beneath; 8. asperum—with leaves scabrous above and tomentose be- neath; y. schrenkii—with leaves green, shallowly incised, almost gla- brous. Apparently, the less incised’ and weakly pubescent specimens are the shade plants of forest formations. The type of the species is characterized by more or less deeply incised leaves, which are grayish- tomentose beneath. From C. maackii, this species is distinguished pri- marily by having capitula that nod from the base and by the form of the leaves. Elucidation of the taxonomic significance of the separate forms that have been proposed for C. schantarense Trautv. and Mey. is not yet possible and will require more thorough investigation of extensive material. Kitamura reports for Korea the hybrid C. x heterocanthum Kitam. C. pendulum Fisch. x C. schantarense Trautv. and Mey. (Kitam. in Acta Phytol. et Geobot., Ill, 1 [1934] 8). For the flora of the USSR this hybrid has not yet been reported, but finding it is possible. Subsection 3. Apalocentron (Cass.) Charadze comb. nova.— Onotrophe sect. Apalocentron Cass. in Dict. Sc. nat. XXXVI (1825) 16 — 159 146.—Cirsium sect. Apalocentron (Cass.) Kitam. in Acta Phytot. et Geobot. XII, 2 (1943) 102 p. min. p.—Leaves thin, more or less broad, usually almost undivided or more or less deeply pinnately incised, with approximate segments. Capitula surrounded by broad apical leaves, or 1-3, small, narrow, bracteal leaves at base of capitula. Outer in- volucral bracts with more or less bulged, resinous, dark band or weakly carinate, acuminate. Florets yellowish or red, occasionally white. Pe- rennial plants mostly confined to woodland regions of Central Europe. A few species reach the European Part of the USSR, and only one extends to Western Siberia. Type of subsection: C. oleraceum (L.) Scop. Note. Apparently, it is a collective species, comparatively recent in origin and readily hybridizing with species of the section Cirsium. Individual species tend toward members of the subsections Sinocirsium, Cirsium, and Montana. 69. C. oleraceum (L.) Scop. Fl. Carn. ed. 2, II (1772) 124; Ldb. Fl. alt. IV, 12; DC. Prodr. VI, 647; Ldb. Fl. Ross. II, 738; Schmalh. Fl. II, 105; Koch, Syn. ed. 3, II (1902) 1550; Fedtsch. and Fler. FI. Evrop. Ross. 1012; Schischk. in Fl. Zap. Sib. XI, 2893; Majevski, FI. ed. 8, 602; Arénes in Bull. Jard. Bruxelles, XXIV, 264; Hegi, Ill. FI. VI, 2, 898.—C. pratense Rupr. FI. Ingr. (1860) 532.—Cnicus oleraceus L. Sp. pl. (1753) 826.—C. pratensis Lam. FI. Fr. II (1778) 14.—C. bracteatus Gilib. Fl. Lithuan. III (1781) 184.—Carduus oleraceus Gorter, Fl. Ingr. (1761) 129; Vill. His. pl. Dauph. III, 21.—C. acanthifolius Lam. Encycl. méth. I (1783) 703.—Ie.: Rehb. Ic. FI. Germ. XV, tab. 834; Syreistsch. Ill. Fl. Mosk. Gub. III, 299; Javorka and Csapody, Iconogr. Fl. Hung. 542, fig. 3880; Fiori, Ic. Fl. Ital. ed. 3, No. 3725; Hegi, Ill. Fl. VI, 2, 898.—Exs.: Pl. polon. exs. No. 746; Pl. Suec. exs. No. 1585. Perennial. Rhizome horizontal, with funiform fibrous roots; stem ascending, more or less branched from middle, 60-150 cm high, sul- cate, subglabrous or scatteredly arachnoid-hairy. Leaves elliptical or ovate, green and glabrous above, glaucescent beneath, somewhat arach- noid-hairy on veins; lower cauline leaves 20—40 cm long, 10-30 cm wide, narrowed to winged, more or less elongate petiole, sinuate-pin- nately parted into ovate, unevenly toothed segments, teeth tapered to thin spines 2—3 mm long; middle cauline leaves more or less deeply pinnatisect into lanceolate, spinescent approximate lobes, sessile, amplexicaul, usually 15 cm long, 5 cm wide; upper cauline leaves ovate or oblong-ovate, more or less pinnately toothed or almost entire, with spinelike cilia, long-acuminate, usually 10 cm long, 4 cm wide. Capitula globose-ovate, 2—3 cm in dia, in clusters of 2—5 at apices of stem and few, long branches, basally surrounded by yellowish-green 16 i) 160 and broadly ovate, in upper capitula more or less lanceolate, apical leaves; capitula more or less nodding. Involucre finely arachnoid-hairy; outer involucral bracts lanceolate, weakly arachnoid-hairy, narrowed into deflexed yellowish spine, with dark green band above, weakly carinate; inner bracts linearly-lanceolate, glabrous, narrowed into long scarious tip. Florets greenish-yellow, 22 mm long; corolla tube nar- row, 8 mm long; limb lobed almost to middle. Achenes 4.5 mm long, pale yellowish-gray; pappus sordid-white, to 19 mm long, hairs with long filiform tips. Flowering July to September. Commonly in forest zone, occasionally in forest steppe, from time to time in steppe regions; in coniferous and birch-aspen forests, pine forests, along forest edges, among scrubs, in wet meadows, often on edges of swamps.—European Part: Dvina-Pechora, Baltic Region, Ladoga-Ilmen, Upper Volga, Volga-Kama, Upper Dnieper, Volga-Don, Trans-Volga, Upper Dniester; Western Siberia: Upper Tobol, Irtysh (Baraba, Zmeinogorsk). General distribution: Scandinavia, Central Europe, Atlantic Europe, Western Mediterranean Region (northern Italy), Balkans (northern). Described from Southern Europe. Type in London. Note. It often hybridizes with species of the section Cirsium. Among them, the hybrids between C. oleraceum and C. palustre that have been reported from the European Part of the USSR are closer in appearance to C. oleraceum. 1. C. oleraceum (L.) Scop. x C. palustre (L.) Scop.—C. lacteum auct.: Iljin in Fl. Yugo-Vost. VI (1936) 408.—Ie.: Syreistsch. JI]. FI. Mosk. Gub. III (1910) 300. Specimens from the Leningrad Region differ from C. oleraceum by having more deeply incised, somewhat decurrent leaves. Some speci- mens (Perm District) have narrower leaves. Plants collected from the Pskov District are closer to C. palustre. For specimens from Syzran, M.M. Iljin notes the following characters: Stem branched above; cauline leaves pinnately divided, lower broadly decurrent, upper short-decur- rent. Capitula medium-sized, with yellow or reddish florets. 70. C. rivulare (Jacq.) All. Auct. ad Fl. Pedem. (1789) 10, excl. descr. sec. Mansfeld in Fedde, Repert. 52, 2, 175; Link, Enum. pl. II, 301; Koch, Syn. (1837) 397; Schmalh. FI. II, 105; Klok. in Vizn. Rosl. URSR, 569; Majevski, Fl. ed. 8, 605.—Carduus rivularis Jacq. FI. Austr. I (1773) 57, tab. 91.—Cnicus rivularis Willd. Sp. pl. II, 3 (1803) 1676.—Cnicus salisburgensis Willd. ibid. 1675.—Ic.: Rchb. Ic. Fl. Germ. XV, tab. 835, fig. 1; Javorka and Csapody Iconogr. FI. Hung. 542, fig. 3882.—Exs.: Fl. exs. austro-hung. No. 1768; Pl. polon. exs. No. 449; GRF No. 1070. 163 161 Perennial. Rhizome with numerous thin-funiform roots; stem 75— 100 cm high, yellowish, ascending, sulcate, scatteredly crisped-hairy below, thinly arachnoid-hairy above, simple, with long few-leaved pe- duncle bearing solitary terminal capitula. Leaves subglabrous above and green or sparsely crisped-hairy above, grayish-green and more or less densely crisped-hairy beneath, especially along veins; basal leaves 15-25 long, 5.5—9.5 cm wide, obovate or ovate-lanceolate, narrowed at both ends basally narrowed into more or less short winged petiole, with large remote lobes, tapered to long, thin spines, usually unequally pinnately incised into approximate, deltoid-ovate, broadly deltoid-lan- ceolate or more or less digitately elongated segments; each segment with subacute deltoid teeth, with dense, short (2-3 mm long) spinelike cilia; apical segments acuminate; lower cauline leaves narrowed at base, auriculate, broadly ovate, very unequally, more or less deeply pinnatisect into oblong, toward middle of lamina almost digitately elongated segments, 9-18 cm long, 6.5—8.5 cm wide, occasionally al- most undivided; upper leaves small, lanceolate, 1-2 on common pe- duncles. Capitula globose-ovate, apical, solitary or in clusters of 2-3, 1.3—2.3 cm in dia, on short peduncles. Involucre imbricate, dark purple, scatteredly arachnoid-hairy; outer involucral bracts ovate-lanceolate, tiny ciliate on margins, spinescent; inner bracts linearly-lanceolate, acuminate. Florets red, to 19 mm long; corolla tube narrow, 8 mm long; limb lobed to middle. Achenes yellowish, 4.5 mm long; pappus sordid-white, with inner hairs toothed at tips. Flowering June to July. Wet, marshy meadows, coniferous forests in logged areas, black alder forests, and along ditches.—European Part: Baltic Region, Ladogo-Ilmen, Upper Volga, Upper Dnieper, Middle Dnieper, Volga- Don, Upper Dniester. General distribution: Central Europe. Described from Europe. Type in London. 71. C. erisithales (Jacq.) Scop. Fl. Carn. ed. 2, II (1772) 125; Koch, Syn. (1837) 395; Schmalh. Fl. II, 105; Fedtsch. and Fler. Fl. Evrop. Ross. 1012.—C. glutinosum Lam. Fl. Fr. I (1778) 27; DC. Prodr. VI, 649.—C. ochroleucum auct. non All.: DC. in DC. and Lam. Fl. Fr. ed. 3, IV (1805) 115; DC. Prodr. VI, 648.—Carduus erisithales Jacq. Enum. Stirp. Vind. (1762) 279.—Cnicus erisithales L. Sp. pl. ed. 2 (1763) 1157; Willd. Sp. pl. III, 3, 1679.—Ic.: Rchb. Ic. Fl. Germ. XV, tab. 837; Javorka and Csapody, Iconogr. Fl. Hung. 542, fig. 3881; Fiori, Ic. Fl. Ital. ed. 3, 468, No. 3722; Dostal. Kvet. CSR, 1671, tab. 553; Hegi, Ill. Fl. VI, 2, 897, fig. 598 and 898, fig. 599. (foto).—Exs:.: Fl. exs. austr.-hung. No. 1768; Fl. exs. reip. Bohem.-Slov. No. 633. Perennial. Rhizome cylindrical, nodulose, oblique, with numerous slender fibrous roots; stem ascending, weakly sulcate, yellowish or 164 162 reddish-brown, at apex drooping, branched above, branches few, more or less long, less often simple, with tiny articulate hairs below, finely arachnoid hairy and glutinous above, densely leafy up to middle, al- most leafless above. Cauline leaves dark green, subglabrous above or with scattered short papillate hairs, somewhat pale green beneath, with flexuous and articulate hairs along veins, with short papillate hairs throughout, oblong or ovate, often pinnately incised almost to base, into 8-12 oblong, acuminate, finely spiny-ciliate segments on each side, with 3-5 parallel veins; basal leaves to 30 cm long, 15 cm wide, narrowed in more or less short, winged petiole; lower cauline leaves with shorter petiole, to 25 cm long, middle leaves sessile, with cordate amplexicaul base, to 12 cm long; upper leaves usually 2 on elongate stem, strongly reduced, 5 cm long, 2.5 cm wide, pinnately parted or only pinnately toothed; apical leaves at base of capitula or inflores- cence, linearly lanceolate, spiny-ciliate. Capitula solitary or 2—5 clus- tered at apices of stem and branches, nodding. Involucre more or less ovate, about 2.5 cm in dia; involucral bracts lanceolate, spinescent, dorsally and along margin scabrous, very tiny appressed-hairy, with dark resinous stripe outside; outer bracts at flowering appressed, all divergent at fruiting; inner bracts linearly lanceolate, with scarious tip, with fine, resinous dark stripe above, finely hairy outside. Florets usually lemon-yellow, occasionally purple, to 18 mm long; limb longer than narrow corolla tube, lobed almost to middle. Achenes 5 mm long; pappus somewhat shorter than florets sordid-white, with fine, rarely plumose hairs with filiform tips. Flowering July to September. Grows singly or in groups or thickets in thin forests and scrubs, on stony slopes, along forest edges, in ravines, near springs, in mountain meadows and pastures.—European Part: Upper Dnieper, Middle Dnieper (left bank), Upper Dniester, Bessarabia. General distribution: Central Europe, Western Mediterranean (Maritime Alps), Balkans (northern). Described from Austria. Type in London. Subsection 4. Cirsium.—Stem ascending, simple or somewhat branched, less leafy above or with almost leafless floriferous branches. Leaves somewhat coriaceous, or thin, glabrous above, finely or arach- noid-hairy beneath, more or less grayish or whitish-tomentose, undi- vided or unequally pinnately parted, with few lobes. Capitula medium, solitary on leafless floriferous branches, occasionally in small clusters. Involucre subglabrous; involucral bracts purple above. Florets red, very rarely white. Perennials, widely distributed in boreal region of Northern Hemi- sphere and in mountainous areas of Eurasia. Type of subsection: type of genus and section. 165 163 Series 1. Heterophylla Charadze.—Cauline leaves sessile. 72. C. heterophyllum (L.) Hill. Hort. Kew. (1768) 64; Ldb. RI. Ross. II, 739 p. p.; Schmalh. Fl. II, 104, excl. var.; Fedtsch. and Fler. Fl. Evrop. Ross. 1011; Schischk. in Fl. Zap. Sib. XI, 2891 p. max. p.; Klok. in Vizn. Rosl. URSR, 569; mishkin, Fl. Khibin, 69; Hegi, Ill. FI. VI, 2, 885.—C. heterophyllum a. helenioides DC. and B. incisum DC. in Duby, Bot. Gall. I (1828) 288.—C. heterophyllum a. indivisum DC. p. p. and B. incisum DC. Prodr. VI (1837) 653; Iljin in Fl. Yugo-Vost. VI, 406; Kryl. Fl. zap. Sib. XI, 2891.—Carduus heterophyllus L. Sp. pl. (1753) 824.—Carduus helenioides L. Sp. pl. (1753) 825, excl. pl. sibir.—Cnicus heterophyllus (L.) Retz. Fl. Scand. Prodr. ed. 2 (1795) 191; Willd. Sp. pl. III, 3, 1673.—Cnicus helenioides (L.) Retz. loc. cit.—Ic.: Rchb. Ic. Fl. Germ. XV, tab. 838; Syreistsch. Ill. Fl. Mosk. Gub. III, 301; Javorka and Csapody, Iconogr. Fl. Hung. 542, fig. 3879; Hegi, Ill. Fl. VI, 2, 885, fig. 587, 886, fig. 588. Perennial. Rhizome thick, oblique, with funiform fibrous roots and long, slender underground branches; stem 50—150 cm high, ascending, sulcate-ribbed, more or less thinly arachnoid-hairy, white-tomentose below capitula, dark purple, simple or with 1—2 branches above, with solitary terminal capitula. Basal leaves 9-12 cm long, 3.5—4.0 cm wide, lanceolate, narrowed into long petiole; cauline leaves green above, glabrous, appressedly grayish-arachnoid-hairy to white-tomentose be- neath, entire or finely toothed and spiny-ciliate; lower leaves 10-30 cm long, 4-8 cm wide, ovate or oblong-lanceolate, narrowed to long, winged petiole, acuminate; middle cauline leaves broadly lanceolate, basally slightly lyrate, with broad amplexicaul auricles; all leaves undivided, in var. indivisum DC. or var. incisum DC. lower leaves unequally pinnately or somewhat palmately incised into oblong seg- ments; upper leaves entire, lanceolate, semiamplexicaul, long-acumi- nate, to 12 cm long, 3 cm wide; apical leaves lanceolate or linearly lanceolate, small, 1-2 bracteal leaves at base of capitula. Capitula 3.5—5.0 cm in dia, globose-ovate, basally concave, solitary on long peduncles and axillary branches. Involucral bracts imbricate, outer ones oblong-ovate, yellowish-brownish-green, weakly carinate, scabrous on upper margin, spinulescent; inner ones linearly lanceolate, gradually narrowed into purple, scarious tip, curved at base. Florets 25-30 mm long; corolla tube narrow, to 11 mm long; limb lobed to 1/3, purple- red, rarely whitish. Achenes stramineous, 4 mm long; pappus 21 mm long, sordid-white or brownish-white with few hairs. Flowering July to August. Thin mixed and larch forests, edges and in forest and alpine 166 meadows; usually in forest zone, occasionally in forest-steppe zone.— 164 European Part: Karelia-Lapland, Dvina-Pechora, Baltic Region, Ladoga-Ilmen, Upper Volga, Volga-Kama, Upper Dnieper, Middle Dnieper (left bank), Trans-Volga, Lower Volga; Western Siberia: Ob Region, Upper Tobol, Irtysh. General distribution: Central Europe, Atlantic Europe. Described from western Europe. Type in London. Note. The variety with deeply incised cauline leaves predominates in the Baltic Region and in the north of the European part of the USSR, and in the other regions both varieties are found; var. indivisum DC. noticeably predominates in Western Siberia. C. heterophyllum (L.) Hill is replaced in Eastern Siberia, the Altai, and the mountainous regions of Kazakhstan by the closely related species—C. helenioides (L.) Hill, which is distinguished by its entire, broad, coarsely toothed leaves, and aggregated, smaller capitula. The following hybrids have been reported from the European part of the USSR: ‘1. C. heterophyllum (L.) Hill x C. palustre (L.) Scop.—C. x wankelii Reichardt (cf. [jin in Fl. Yugo-Vost. VI, 409) with small, decurrent, pinnately divided leaves, somewhat smaller capitula in clus- ters of 1-3, and purple florets. 2. C. oleraceum (L.) Scop. X C. heterophyllum (L.) Hill.—C. affine Tausch (see Iljin in Fl. Yugo-Vost. V1, 408). Forms that ap- proach C. heterophyllum in habit and are distinguished by yellowish florets and broader leaves have been found in the Leningrad Region (Luga, Tikhvinka) and along the northern limit of the range of C. oleraceum (Ust-Sysolsk). 73. C. helenioides (L.) Hill, Hort. Kew (1768) 64 p. p.—C. heterophyllum auct. non Hill: Ldb. FI. Ross. II (1846) 739 p. p. quoad pl. Siber., Alt. and Baikal; Turcz. Fl. baic.-dahur. II, 1, 129; Kryl. FI. Zap. Sib. XI, 2891 p. p—C. heterophyllum ssp. angarense M. Pop. Fl. Sr. Sib. II (1959) 753.—C. heterophylloides Pavl. Fl. Tsentr. Kazakhst. III (1938) 313; Botschantzev in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX (1959) 628.—Carduus helenioides L. Sp. pl. (1753) 825 p. p. quoad pl. e “Sibiria” excl. syn europ.—Cnicus helenioides Willd. Sp. pl. III, 3 (1803) 1674. Perennial. Rhizome large, horizontal, dark brown, with long woody, funiform fibrous root. Stem 40—130 cm high, ascending, simple, leafy, more or less densely crisped-hairy to lanate, usually tomentose below capitula, finely sulcate. Leaves with scattered flexuous hairs above, at places finely floccose-arachnoid-hairy, more or less densely grayish- tomentose beneath, with more or less distinctly raised thin veins, deeply 167 165 biserrate-dentate, with recurved, broadly deltoid teeth, tapered into spines to 2 mm long, finely appressed-spinose between teeth; leaves on short branches at base of stem oblong-lanceolate, weakly toothed, 10—20 cm long, 2—4 cm wide, narrowed into petiole almost as long as lamina, spiny-ciliate; lower cauline leaves broadly oblong-ovate to broadly lanceolate, acuminate, lowermost narrowed into winged peti- ole, others auriculate, semiamplexicaul, 28—35(60) cm long, 6—13(25) cm wide, coarsely toothed; upper leaves up to 12 cm long, 2.5 cm wide, lanceolate, amplexicaul, weakly toothed and finely spiny-ciliate; apical leaves small, linear, spinose on margins. Capitula globose, 1.3— 2.2(3) cm in dia, 2—5 at apices of stem, occasionally solitary, on longer peduncles in corymbose common inflorescence. Involucre subglabrous, dark reddish-brown to brown, imbricate; outer bracts lanceolate, gla- brous, ciliolate, dark reddish-brown with almost black midrib, puberu- lent above, spinescent; inner bracts membranous, linearly lanceolate, with scarious, purple cusp. Florets red, to 22 mm long; corolla tube narrow, about 8-10 mm long; limb lobed to more than 1/2. Achenes curved, yellowish, with brown streaks, to 4 mm long; pappus brown- ish-sordid-white, to 16 mm long, with inner hairs weakly toothed at apices. Flowering July to August. Mountain herb slopes, river valleys, deciduous forests, forest edges and lake shores.—Western Siberia: Altai; Eastern Siberia: Angara- Sayans, Dauria; Far East: Zeya-Bureya; Central Asia: Lake Balkhash Region. General distribution: northern Mongolia. Described from Si- beria. Type in London. Note. In the Sayans this species replaces the boreal Euro-Siberian C. heterophyllum (L.) Hill. For Carduus helenioides L., Linnaeus cited the following localities: “Anglia, Siberia.” Apparently, Linnaeus used this name for the entire-leaved forms of C. heterophyllum (L.) Hill from Siberia. Smith (Smith in Engl. Bot. III [1794] 177; X [1799] 675), who studied the specimen of Carduus helenioides in the Lin- naean Herbarium, notes that specimens matching the type are not found in Europe and, possibly, the name Cirsium helenioides should be ex- cluded from the flora of Europe. Later, Airy-Shaw, having combined the European forms that have been treated as Cirsium heterophyllum and Cirsium helenioides, restored the name C. helenioides (L.) Hill as the earliest combination. Following Smith, he notes that the Linnaean Herbarium contains only one specimen, apparently, from Siberia, which is different from Carduus heterophyllus, described from Europe. We think that the name C. helenioides (L.) Hill sensu stricto should be retained for the Eastern Siberian mountain species, and in C. heterophyllum (L.) Hill, distributed in the boreal regions of Europe and Western Siberia, one can recognize the varieties, segregated 168 166 already by De Candolle, var. incisum DC. and var. indivisum DC., which, evidently, do not have any special taxonomic significance. It is also necessary to treat C. heterophylloides Pavl., described from Kazakhstan by N.V. Paviov as a synonym of C. helenioides (L.) Hill s. str. Its characters agree fully with those of the specimen in the Linnaean Herbarium. We were able to verify this while examining the microfilm of the Linnaean Herbarium in the library of the Botanical Institute of the Academy of Sciences of the USSR. Later, M.G. Popov split off from C. heterophyllum (L.) Hill, as a subspecies, the plant from Central Siberia, proposing the name ssp. angarense M. Pop for it. The latter should also be referred to C. helenioides (L.) Hill. M.G. Popov indicates that typical C. heterophyllum (L.) Hill reaches the Yenisei River in the east and on the south side of the Western Sayans, it is replaced eastward by a separate subspecies. The boundaries of both species in the USSR still need to be worked out. 74. C. dealbatum MB. FI. taur.-cauc. III (1819) 560; C.A.M. verzeichn. 70; DC. Prodr. VI, 653; Ldb. Fl. Ross. II, 740; Boiss. FI. or. III, 539; Petrak in Tr. Tifl. Bot. Sada, XII, 1, 28; in Izv. Kavk. Muz. VIII, 1-2, 63; Grossh. Fl. Kavk. IV, 186.—C. heterophyllum var. dealbatum Schmalh. Fl. II (1897) 104.—C. heterophyllum B. dealbatum Lipsky, Fl. Kavk. (1899) 358. Perennial. Stem ascending, hard, 70-100 cm high, weakly ribbed and sulcate, brownish-purple, more or less densely floccose-arachnoid- hairy, with large leaves in lower part, usually in upper part with 1—5 elongate, almost leafless branches bearing solitary capitula. Cauline leaves sessile, amplexicaul, middle ones 25—28 cm long; 7.5—9.5 cm wide, broadly ovate, abruptly narrowed from middle into lanceolate, long-acuminate tip, with almost lanceolate point, green above, with scattered arachnoid hairs, at places floccose, more or less densely grayish- or whitish-tomentose beneath, undivided, unequally small- toothed, finely spiny-ciliate, teeth tapered to spines 2 mm long. upper cauline leaves at base of peduncles reduced, auriculate, gradually narrowed above, 9-12 cm long, 3.5—4.5 cm wide; apical leaves (1—2) 40 mm long. Capitula globose-ovate, erect, solitary, 2.5—3.5 cm in dia, with slightly concave base. Involucre dark brown to purple, sparsely arachnoid-hairy, more or less compactly imbricate; outer bracts mem- branous, ovate-lanceolate, greenish-yellow below, blackish-purple above, tapered to scarcely, noticeable spine, entire below, very finely fimbriate above, about 7 mm long, gradually longer toward inner bracts; inner bracts oblong-lanceolate to linearly-lanceolate, weakly broad- ened above, transitional to scarious, purple, acuminate, deflexed tips. 169 Florets purple-red, 22-25 mm long; corolla tube 11-12 mm long; limb 170 167 lobed to 1/3. Achenes yellowish, to 4 mm long; pappus sordid-white, 19 mm long, with few hairs, inner hairs with slightly longer toothed tips. Flowering July to August. Moist places, among forests, from mid-montane to subalpine zone.—Caucasus: Ciscaucasia (basin of Kuban River and vicinity of Kislovodsk). Endemic. Described from Kislovodsk. Type in Leningrad. Note. It differs from C. heterophyllum (L.) Hill by having broader, always entire cauline leaves, abruptly narrowed from the middle, and strongly elongate, leafless flowering branches bearing larger solitary capitula. It differs from the central Siberian species C. helenioides (L.) Hill by having well spaced, larger capitula and acuminate cauline leaves. Moreover, for C. dealbatum MB. outer and inner involucral bracts that are somewhat broadened above are characteristic. Series 2. Cana Charadze.—Cauline leaves auriculate, more or less decurrent. 75. C. canum (L.) All. Fl. Pedem. I (1785) 151; Moench, Meth. 556; MB. FI. taur.-cauc. III, 556 p. p. excl. pl. cauc.; Ldb. Fl. Ross. II, 741 p. p.; Schmalh. Fl. I, 103; Fedtsch. and Fler. Fl. Evrop. Ross. 1011; Schischk. in Fl. Zap. Sib. XI, 2888; Klok. in Vizn. Rosl. URSR, 569; Hegi, Ill. Fl. VI, 2, 883.—Carduus canus L. Mant. pl. (1767) 105; L. Syst. veg. ed. 10 (1784) 25; 1 Jacq. Fl. Austr. I, 27.—Cnicus canus Willd. Sp. pl. III, 3 (1803) 1664; MB. FI. taur.-cauc. II, 274 p. p.—lIe.: Jacq. Fl. Austr. I (1773) 27, tab. 82 and 83; Rchb. Ic. FI. Germ. XV, tab. 828; Javorka and Csapody, Iconogr. Fl. Hung. 541, fig. 3877; Dostal, Kvet. CSR, 1663, tab. 549, 3; Hegi, Ill: Powis; 884, fig. 586.—Exs.: Petrak, Cirsioth. Univ. No. 35. Perennial. Rhizome short, vertical, with fusiform, thick, long roots from near root neck. Stem to 120 cm high, erect, weakly branched, deeply sulcate, purple in lower part, scatteredly arachnoid-hairy at base of capitula almost tomentose, more leafy in lower part with large, short-decurrent leaves, distant, gradually reducing leaves in upper part; basal leaves oblong-lanceolate, with winged petiole, 25—40 cm long, 4—7 cm wide long-acuminate, on both sides finely arachnoid-hairy or floccose, undivided, occasionally pinnately divided, more or less bicristate-toothed, densely setose, with thin, long and short spiny bristles; lower cauline leaves lanceolate and oblong-lanceolate, 18—20 cm long, 3—5 cm wide, approximate, with broad and short petiole narrowing downward decurrent on stem; middle cauline leaves reduced above, from 7-8 to 4 cm long, elongate to linearly-lanceolate, sessile, with slightly decurrent roundish auricles; all cauline leaves usually undivided or pinnate, subglabrous or thinly arachnoid-hairy, densely 17 — 168 spiny-setose; apical leaves on elongate peduncles, undivided, linear, arachnoid-hairy, with occasional cilia on margins, at base of capitula 0.5—1.0 cm long. Capitula solitary or 2—3 at apices of stem and axil- lary branches, (1.5) 2—2.5(3) cm in dia, at end of flowering wide open, somewhat nodding. Involucral bracts small, imbricate, outer from ovate- oblong to ovate-lanceolate, tapered into short, purple spines 1.0—1.5 mm long, with dark purple band above, scatteredly arachnoid-hairy on margin, inner ones glabrous, lustrous, purple above, narrowed to slightly scarious, acuminate cusp. Florets red, to 20 mm long; corolla tube to 11 mm long; limb 5-parted to middle. Achenes lustrous, stramineous, 3.5—-4.0 mm long, 1.5 mm wide, with narrow edge and conical tu- bercle; pappus to 14 mm long, sordid-white, with thin, sparsely plu- mose hairs, inner hairs apically scabrous. Flowering July to August. Steppe and forest-steppe zones, in alkaline and wet meadows.— European Part: Upper Dnieper, Middle Dnieper, Volga-Don, Trans- Volga, Upper Dniester, Bessarabia, Black Sea Region, Lower Don; Western Siberia: Upper Tobol, Irtysh. General distribution: Scandinavia (southern Norway), Central Europe, Mediterranean Region (western), Balkans. Described from Austria. Type in London. Note. This species has not been studied adequately. The form with entire, or more rarely, pinnately incised cauline leaves, more common in Western Europe, predominates in the European Part of the USSR. The hybrid C. oleraceum (L.) Scop. x C. canum (L.) All—C. tataricum auct. Iljin in Fl. Yugo-Vost. VI (1936) 408—has been re- ported. In habit it resembles C. canum, differing from it by having yellow- ish florets and clustered capitula (specimens from Kursk Region and Buguruslan District in Trans-Volga); given reddish florets and the form of the cauline leaves it is closer to C. canum (specimen from Ulyanovsk). 76. C. biebersteinii Charadze in Fl. Azerb. VIII (1916) 409, descr. ross.; in Zam. po Sist. i Georg. Rast. Tbil. 23 (1963) 112, diagn.—C. canum MB. FI. taur.-cauc. III (1819) 556 p. p.; DC. Prodr. VI, 644; Ldb. Fl. Ross. II, 741 p. p.; Boiss. Fl. or. III, 541 p. p.; Grossh. FI. Kavk. IV, 187, non All.—Cnicus canus MB. FI. taur.-cauc. II (1808) 274 p. p. non Willd. Perennial. Rhizome short, with fasicle of thick fusiform roots; stems ascending, 50-100 cm high, thick, deeply sulcate, floccose, white- tomentose-arachnoid, simple [or] branched above, with 3—4(6) long flowering branches. Leaves very large, crowded in lower part of stem, gradually reduced toward stem apex, on both sides more or less gray- ish, at places floccose-arachnoid-hairy; basal and lower cauline leaves 172 169 broadly-obovate or broadly elliptical, short acuminate, narrowed in more or less short winged petiole, deltoid-sinuate-toothed, finely spiny- setose, spines on teeth to 4 mm long; basal leaves to 40 cm long and 10 cm wide; lower cauline leaves 18-25 cm long, 8-10 cm wide, middle ones broadly lanceolate, to 15 cm long and 4 cm wide, with short, usually unequal, basally obtuse, rounded, short-decurrent au- ticles. Capitula solitary, terminal on stem and axillary branches, latter on strongly elongate peduncles often projected above stem apex, form- ing depauperate corymbose-paniculate inflorescence; peduncles below capitula tomentose, more or less projected above leaves, after flower- ing weakly falcately recurved. Involucre 2—2.5 cm in dia; involucral bracts imbricate, ovate-lanceolate to oblong-lanceolate, 5—7 mm long, coriaceous, yellowish-green, with broad dark-purple band above, sparsely arachnoid-hairy, gradually narrowed and weakly recurved, spinulescent; inner bracts oblong-lanceolate, membranous, with lan- ceolate, usually purple deflexed tip. Florets to 23 mm long; corolla tube 12 mm long; limb lobed to 1/2 into unequal linear lobes. Achenes stramineous, 4 mm long; pappus sordid-white, with occasional, sparsely plumose hairs, inner hairs short-plumose with weakly toothed tip. Flow- ering August to September. Lower to middle-montane zone, along irrigation ditches, in flood- plain forests, scrubs.—Caucasus: Ciscaucasia Dagestan, Southern Transcaucasia. General distribution: Armenia and Kurdistan. Described from Ciscaucasia. Type in Leningrad. Note. C. biebersteini Charadze replaces C. canum (L.) All. in the regions of Eastern and Central Ciscaucasia, and also in Southern Transcaucasia, differing from the latter by broader, more coarsely toothed lower leaves, densely grayish-arachnoid-hairy on both sides, and by narrower involucral bracts. 77. C. pannonicum (L. fil.) Link Enum. pl. II (1822) 299; DC. Prodr. VI, 645; Ldb. Fl. Ross. II, 741; Fedtsch. and Fler Fl. Evrop. Ross. 1011; Klok. in Vizn. Rosl. URSR, 568; Hegi, Ill. Fl. VI, 2, 881.—C. canum y. pannonicum Schmalh. F1. II (1879) 104.—Carduus pannonicus L. fill. Suppl. (1781) 348; Willd. Sp. pl. III, 3, 1658.—Ie.: Jacq. Fl. Austr. IT (1774) tab. 127; Rchb. Ic. Fl. Germ. XV (1852) tab. 829; Syreistsch. Ill. Fl. Mosk. (1910) 303; Javorka and Csapody, Iconogr. Fl. Hung. (1933) 541, fig. 3878; Hegi Ill. Fl. VI (1929) 882, fig. 584.—Exs.: Fl. exs. austro-hung. No. 212; Fl. exs. reip. Bohem.— Slov. No. 966; FI. Ital. exs. No. 692; Petrak, Cirsioth. Univ. No. 15; GRF No. 1071. Perennial. Rhizome horizontal, basally with fibrous remnants of old leaves. Stem to 110 cm high, ascending, ribbed, finely arachnoid- 173 170 hairy, at places floccose, usually stramineous or greenish, below densely- and above sparsely leafy, with long, glabrous peduncles, weakly branched, bearing 1—3 terminal capitula. Leaves oblong-lanceolate or lanceolate-oblong, densely pubescent above, at base hairs tuberculate, flexuous, more or less finely grayish arachnoid-hairy beneath, densely spinose-setose from long, thin, and shorter bristles. Basal leaves entire or toothed, narrowed into short-winged petiole, 13-17 cm long, 3- 3.3 cm wide; lower cauline leaves oblong, to 25 cm long, 4.5 cm wide, basally narrowed, short-decurrent with narrow wing, subacute; middle cauline leaves reduced, linearly-lanceolate, usually to 7 cm long, short- decurrent with broad wing; peduncles considerably projected above leaves. Capitula solitary, terminal on stem and 2-3 axillary branches, erect, at end of flowering slightly nodding, 1.5—2.2 cm in dia. Involu- cre imbricate, scatteredly arachnoid-hairy; outer bracts small, to 3 mm long, gradually longer inward, ovate-lanceolate to oblong-lanceolate, glabrous and lustrous beneath, with blackish-purple band above, spinescent, dorsally bulged, with scattered crisped or arachnoid hairs; inner bracts linearly-lanceolate, with scarious tip, dorsally with thin dark band. Corolla purple-red, 19-22 mm long with narrow tube. 7— 9 mm long. Achenes stramineous, with sharp edge and conical tubercle at tip, obliquely truncate, curved; pappus yellowish-whitish with few scabrous inner hairs at apex. Flowering July to August. Scrubs and forest slopes.—European Part: Upper Volga, Middle Dnieper, Volga-Don. General distribution: Central Europe, Western Mediterranean Region, Balkans (north). Described from Western Eu- rope. Type in London. Note. It differs from C. canum (L.) All. with which it is often confused, by its narrower and smaller leaves, pubescence, long leaf- less peduncles, and smaller involucral bracts. A hybrid has been re- ported from the Tambov and Voronezh regions.—C. heterophyllum (L.) Hill x C. pannonicum (L. fil.) Link. Subsection 5. Spanioptilon (Less.) Kitam. in Acta Phytot. et Geobot. III, 1 (1934) 10; Less. Syn. Comp. (1832) 10, pro gen.; DC. Prodr. VI, 621, pro gen.—Sect. Onotrophe DC. Prodr. VI (1837) 644 p. min. p.—Subsect. Borealicola Kitam. op. cit. 7, p. min. p.— Capitula medium-sized, distant from or approximated to apical leaves; involucral bracts imbricate, usually spinescent, entire, in upper part sometimes membranous and erosely toothed. Corolla tube narrow, al- most as long as limb. Perennials with simple or weakly branched stem, not winged, occasionally narrow winged below and usually undivided or somewhat lobate narrow leaves; basal leaves like cauline, usually withering early or persistent. 174 171 Type of subsection: C. lineare (Thunb.) Sch. Bip. The species of the subsection are distributed in Eastern Asia, in- cluding the Japanese islands, Siberia, and Mongolia. We are broaden- ing the circumscription of this subsection somewhat, by including in it species of the C. serratuloides (L.) Hill alliance. Series 1. Vlassoviana Charadze.—Leaves sessile, grayish-arach- noid-hairy; basal leaves withering early; apical leaves few, narrow, at base of capitula. 78. C. vlassovianum Fisch. in DC. Prodr. VI (1837) 653; Ldb. FI. Ross. II, 741; Maxim. Prim. fl. amur. 175; Kom. Fl. Man’chzh. III, 753; Nakai in Tokyo Bot. Mag. XXVI, 372; Kom. and Alis. Opred. Rast. Dal’nevost. Kr. II, 1085; Kitam. Comp. Japon. I, 87 p. p.; Popov, Fl. Sr. Sib. I, 754.—C. vlassovianum a. genuinum Herder in Bull. Soc. Nat. Mosc. XLIII, 1 (1870) 8.—C. vlassovianum B. bracteatum Ldb. FI. Ross. II (1846) 741; Herder, op. cit. 9; Nakai in Tokyo Bot. Mag. XXVI, 372.—C. vlassovianum y. laciniatum Herder, op. cit. 9.—Cnicus vlassovianus (Fisch.) Maxim. in Bull. Acad. Sc. Pétersb. XIX (1874) 509 and in Mel. Biol. IX, 329. Perennial. Rhizome thick, with few, more or less slender, fibrous roots. Stem dark reddish-brown, purple, or greenish, basally covered with remnants of old basal leaves, simple or weakly branched only above, 60—100 cm high, sulcate, more or less scatteredly crisped-hairy, with whitish articulate hairs. Basal and lower cauline leaves lanceolate, 8-20 cm long, 2.8—3 cm wide, narrowed into short petiole, apically attenuated, green above with scattered crisped appressed hairs, weakly scabrous, grayish-arachnoid-hairy to grayish-tomentose beneath, serru- late-dentate, spinous-setose with erect, long and short spines; cauline leaves gradually reduced toward stem apex, 2—9 cm long, sessile; middle ones lanceolately narrowed from more or less broad, almost amplexicaul base; apical leaves lanceolate, few at base of capitula, as long as ca- pitula or shorter. Capitula solitary, terminal on stem and short axillary branches, usually 1.8—2.5 cm in dia. Involucre weakly arachnoid-hairy; outer bracts imbricate, lanceolate, from outer (to 7 cm long) to inner gradually longer, dark green, lustrous, dark purple above, somewhat deflexcted, more or less strongly spinescent; inner bracts linearly lan- ceolate, membranous, with scarious long point. Florets purple, to 20 mm long; corolla tube to 9 mm long; limb lobed to middle. Achenes slightly compressed, to 3.5 mm long, with long narrow tubercle at apex, stramineous, with scattered purple bands; pappus with few bristles to 15 mm long, inner hairs apically toothed. Flowering July to August. Meadows, glades, forest edges, mainly on sandy soils.—Eastern 176 172 Siberia: Dauria; Far East: Zeya-Bureya, Ussuri. General distribution: Mongolia, northern China, Korean Peninsula. Described from Dauria. Type in Geneva. Note. The Siberian specimens are marked by narrower, entire leaves. Var. B. bracteatum Ldb., described by Ledebour, is character- ized by having the apical leaves surrounding the capitula and surpass- ing them. In this character, it approaches C. coryletorum Kom. Herder (1. c.) recognizes the following forms for C. vlassovianum: genuinum Herd., bracteatum Ldb., and laciniatum Herd., evidently without taxo- nomic significance. 79. C. coryletorum Kom. in Izv. Glavn. Bot. Sada, XVI (1916) 179; Kom. and Alis. Opred. Rast. Dal’nevost. Kr. II, 1085.—C. vlassovianum Kitam. Comp. Japon. I (1937) 87 p. p. non Fisch. Perennial. Rhizome short, with thick, woody, fibrous roots. Stem 50-100 cm high, usually branched from middle, with distant and arcu- ate -branches, sulcate, weakly ribbed, often purple on ribs, densely leafy, scatteredly hairy or glabrous. Leaves oblong-lanceolate, nar- rowed from broad base, almost deltoid-lanceolately pointed above; lower cauline leaves sessile, acuminate, unequally toothed, oblong- lanceolate, to 20 cm long, 5 cm wide; middle cauline leaves sessile, 4(8.5)—10 cm long, (1)2.5—3.5 cm wide, with obtuse, semiamplexicaul auricles; all leaves sharply sinuately toothed, teeth bearing short, hard, apical spine; lamina finely spinose with appressed, squarrose, spines, glabrous and dull green above, more or less finely arachnoid-hairy, almost grayish-tomentose beneath; leaves on flowering branches usu- ally smaller, upper of them lanceolate, to 2.5 cm long, 0.7 cm wide, almost reaching base of capitula. Capitula solitary, terminal on stem and branches, in corymbose-paniculate common inflorescence. Involu- cre (1.5)2—3 cm in dia, weakly grayish-arachnoid-hairy, with basal, unequal, 6-20, linearly lanceolate or linear involucral bracts, usually surpassing capitulum or almost as long; outer bracts firmly appressed, shorter than or almost as long as inner ones, lanceolately-linear, very slightly broadened above, very weakly spinescent, dorsally arachnoid- hairy; inner bracts membranous, lustrous, with purple acute scarious point. Corolla to 17 cm long; corolla tube to 7 mm long; limb deeply 5-parted to middle. Florets whitish, usually with purple staminal tube. Achenes stramineous, apically with long, thin tubercle up to 4 mm long; pappus yellowish-brown, to 19-20 mm long, with apically long-plumose inner hairs. Flowering August to September. (Plate VII, Fig. 2.) On hummocks in scrubs, in forest meadows, usually in filbert thickets, occasionally in old fields —Far East: Ussuri. Endemic. Described from region of Gulf of America. Type and paratype in Leningrad. 174 13 a7 bay AN BEE. i iy Wee SY, >. ZANT Plate VIII. 1, 2—Cirsium glabrifolium (Winkl.) O. and B. Fedtsch. 3—C. sieversii (Fisch. and Mey.) Petrak. 177 174 Note. A very distinct species, which approaches C. vlassovianum Fisch. but is well distinguished from it by the color of the corolla, form of the larger cauline leaves and the small leaves on the inflores- cence branches, as well as by the long apical leaves closely subtending the capitulum like an involucre. The few later collections of this spe- cies also are from the Pacific Coastal Region (Olga Bay, Posjet Bay and others). Kitamura (1. c.) includes this species as a synonym of C. vlassovianum Fisch. to which it undoubtedly is very close, but from which it is well distinguished by the characters indicated. Series 2. Asiatica Charadze.—Subsect. Borealicola ser. Erecta Kitam. in Acta Phytot. et Geobot. III, 1 (1934) 10 p. p.—Plants subglabrous with more or less decurrent lower cauline leaves. 80. C. serratuloides (L.) Hill, Hort. Kew. (1768) 64 p. p. non Scop. (1772); Ldb. Fl. alt. IV, 7; DC. Prodr. VI, 652; Ldb. Fl. Ross. II, 742; Kryl. Fl. Alt. Ill, 693.—C. asiaticum Schisckh. in Fl. Zap. Sib. XI (1949) 2890; Grubov, Konsp. Fl. Mong. No. 1781; Popov, FI. Sr: Sib. II, 753.—Carduus serratuloides L. Sp. pl. (1753) 825, excl. syn. europ.—Cnicus serratuloides Roth. Tent. Fl. Germ. I (1788) 346 p. p.; Willd. Sp. pl. Il, 3, 1674, excl. syn. europ.—Iec.: Gmel. FI. sibirica, II (1749) tab. 23, fig. 1; Ldb. Ic. pl. Fl. Ross. IV, tab. 375.— Exs.: Petrak, Cirsioth. Univ. No. 100. Perennial. Rhizome thick, covered with numerous fibrous roots. Stem ribbed, with scattered flexuous hairs or subglabrous, to 1 m high, simple or branched above, brownish-purple, leafy to tip. Cauline leaves lanceolate, long, lower ones semiamplexicaul, somewhat decurrent, middle ones sessile, gradually reduced toward stem apex, cordate, semiamplexicaul, short-acuminate, usually undivided, occasionally toothed, tiny spiny-setose, green and usually glabrous above, occa- sionally with scattered flexuous hairs, somewhat glaucous beneath, glabrous or on veins arachnoid-hairy, 5-14 cm long, 1-3 cm wide. Capitula solitary, terminal on stem and long, sparsely leafy branches, few, 23.5 cm in dia. Involucre weakly arachnoid-hairy or glabrous; involucral bracts lanceolate, attenuate into short, slightly deflected spine, dark green above with purple band, sparsely arachnoid-hairy; inner bracts linearly lanceolate, acuminate into scarious, colored tip. Corolla lilac-purple, 22-23 mm long; corolla tube 12 mm long; limb lobed to middle. Achenes reddish-brown, to 3.5—-4 mm long; pappus silvery- white, 16 mm long, inner hairs with somewhat pointed tips. Flowering July to August. Thin coniferous, mixed, and deciduous forests, forest edges, river banks, tall herb meadows. In mountains it reaches to lower belt of 178 175 alpine zone.—Western Siberia: Ob Region, Upper Tobol, Irtysh, Altai; Eastern Siberia: Yenisei, Lena-Kolyma, Angara-Sayans, Dauria; Far East: Zeya-Bureya. General distribution: Mongolia. Described from Siberia. Type in London. Note. We refer C. serratuloides (L.) Hill to the subsection Spanioptilon, bringing this species close, on the one hand, to tie east- ern Asiatic C. vlassovianum in the leaf form and, to some extent, the pubescence of flexuous hairs, which are weakly developed in this species. On the other hand, C. serratuloides is related to C. komarovii through C. schischkinii. The latter species shows some resemblance to C. mongolicum, sharply differing from it in leaf form and more winged stems. In C. serratuloides variability is manifest in the form of the leaves and the pubescence. Plants from the Yenisei region have entire leaves and larger capitula. In Eastern Siberia, there are forms with toothed and longer spiny leaf margins; in the nature of pubescence they approach C. vlassovianum. Some of them may be considered hybrids between C. serratuloides and C. vlassovianum. M.G. Popov (FI. Sr. Sib. Il, 759) describes the hybrid C. serratuloides x C. heterophyllum from the Kuta River with the following characters: leaves, on the average, 10—20 cm long, 5 cm wide, grayish- tomentose beneath, spinose-ciliate, amplexicaul. Capitula solitary. 81. C. schischkinii serg. in Sistemat. Zam. Gerb. Tomsk. Univ. 2 (1941) 5; Schishk. in Fl. Zap. Sib. XI, 2899.—C. komarovii Schischk. in Fl. Zap. Sib. XI (1949) 2899 p. min. p. Perennial. Stem 35—80 cm high, branched above, weakly ribbed, with scattered, articulate and flexuous hairs; densely leafy. Leaves oblong or linearly-lanceolate, glabrous above, weakly arachnoid-hairy on midrib, glabrous or scatteredly arachnoid-hairy beneath, with squar- rose glandular hairs; middle cauline leaves 14-17 cm long, 2—3 cm wide, shallowly sinuately-pinnately lobed, basally narrowed, with nar- row short-decurrent wing, apically attenuated, acuminate; leaf lobes broadly deltoid, distant, sinuate-toothed, teeth strongly tapered to hard, yellowish spines 2—4 mm long, finely spinose-ciliate; upper leaves reduced, lanceolate, sessile, with short wing weakly decurrent, more or less long attenuate, 3.5—-8.0 cm long, 0.5—1.5 cm wide; apical leaves reaching base of capitula or distant. Capitula solitary, terminal on stem, or 3—4 on long leafy branches, in corymbose-paniculate general inflo- rescence, projected above apical leaves. Involucre 2—3.5 cm in dia, weakly arachnoid-hairy; involucral bracts gradually elongated from outer to inner, subglabrous, weakly arachnoid-hairy on margin; outer bracts basally pale yellow, dark green above, 7-12 mm long, long WS 176 acuminate into short, weakly deflected spine; middle bracts broadly ovate-lanceolate, purple above, attenuate into short spine, inner ones linear-lanceolate, with purple scarious tip. Corolla dark pink, to 22 mm long; limb lobed to 1/3. Achenes yellowish, 3.5—4 mm long, apically with long tubercle; pappus sordid-white, 18-19 mm long, inner hairs with slightly longer thickened tips. Flowering August. _ River valleys in wet meadows.—Western Siberia: Altai; Eastern Siberia: Angara-Sayans. Endemic. Described from Chulyshman Valley and also from southern slope of Lake Teletskoe. Type in Tomsk; paratype in Leningrad. Note. It is distinguished from C. serratuloides (L.) Hill by its pinnately incised, long-spinescent leaves, and from C. komarovii Schisckh. by the less winged stem and oblong leaf lobes. The plants collected near the confluence of the Chedygem and Argut rivers and identified by B.K. Schischkin as C. komarovii Schischk., apparently are close to C. schischkinii Serg. Pubescent plants of this species are reported by L. Sergievskaya as C. schischkinii var. villosum Serg. (Khemchik River Valley in Tuva Province). 82. C. komarovii Schischk. in Bot. Zhurn. SSSR, XXIV, 5-6 (1939) 420 p. max. p.; in Fl. Zap. Sib. XI (1949) 2899 p. max. p.— Ic.: Schischk. op. cit. (1939) 421. Perennial. Rhizome horizontal, thick, nodulose, with funiform, fibrous roots. Stems basally covered with blackish-brown remnants of early-withering basal leaves, ascending, 50-100 cm high, simple or with 2—3 short branches in upper part, densely leafy, more or less scatteredly arachnoid-hairy, especially in lower part, winged from decurrent leaves, wings narrow, almost undivided, toothed, with thin and long-spinescent teeth; cauline leaves broadly linear or linearly lanceolate, repandly and retusely pinnately toothed, finely spinose- ciliate, teeth strongly tapered into hard, yellowish spines, 3-5 mm long, with scattered flexuous hairs above, more or less thinly arach- noid-hairy beneath; middle cauline leaves usually 10-12 cm long, 1— 1.5 cm wide, subacute, with more of less long wings decurrent on internodes, leaves gradually reduced toward stem tip, short decurrent, to 5 cm long, narrowly-lanceolate; apical leaf approximate to base of capitula. Capitula solitary, terminal on stem and few axillary branches, short-pedunculate, somewhat projected above subtending leaves, 1.5— 2 cm in dia, weakly arachnoid-hairy. Involucral bracts ovate-lanceolate, dark green, sparsely arachnoid-hairy along margin, strongly spinescent, inner ones linear-lanceolate, dark purple above, with scarious, weakly elongate tip. Corolla lilac-purple, to 22 mm long; corolla tube to 10 mm long; limb lobed to 1/3. Achenes unknown; pappus sordid-white, 180 177 to 17 mm long, inner hairs with long, scabrous, slightly thickened tips. Flowering July to August. Stony slopes and screes, reaching to subalpine zone.—Western Siberia: Altai (basin of Argut River, Kukurei and Karakem river val- leys, Narym Range, Bukhtarma). Endemic. Described from Argut River. Type in Leningrad. Note. B.K, Schischkin allies C. komarovii Schischk. with C. mongolicum Petrak, from which it differs by having entire, weakly pubescent leaves. Specimens from the Argut River collected near the Chedygem estuary are barely distinguisable from C. schischkinii Serg. and are referred by me to the latter. Subsection 6. Borealicola Kitam. in Acta Phytot. et Geobot. III, 1 (1934) 7; Comp. Japon. I (1937) 41.—Rhizome with numerous scales. Basal leaves persisting after flowering, as long as middle cauline leaves or shorter; only middle cauline or all leaves decurrent. Capitula soli- tary, nodding. Involucral bracts long, outer somewhat shorter than inner, more or less deflexed, with short cusp, entire, or bracts imbricate, small, acute. The members of the subsection Borealicola are distributed in Sakhalin, Kamchatka, Hokkaido Island, and the Aleutian Islands. One species—C. palustre (L.) Scop.—is distributed in the boreal region of Eurasia. Type of subsection: C. kamtschaticum Ldb. Series 1. Nutantia Kitam. in Acta Phytot. et Geobot. III, 1 (1934) 7; Comp. Japon. I (1937) 41.—Capitula nodding, on long peduncles. 83. C. kamtschaticum Ldb. ex DC. Prodr. VI (1837) 644; Ldb. Fl. Ross. II, 736; Nakai in Tokyo Bot. Mag. XXVI, 360; Hultén, FI. Kamtsch. III, 214; Kom. Fl. Kamch. III, 182; Kitam. Comp. Japon. I, 42 p. p.; Hultén, Fl. Aleut. Isl. 334; Vasiliev, Fl. i Pal. Komand. Ostr. 193.—C. kamtschaticum a. genuinum Herder in Bull. Soc. Nat. Mosc. XLII, 1 (1870) 84.—Cnicus kamtschaticus a. genuinus Maxim. in Bull. Acad. Sc. Pétersb. XIX, (1874) 495 and in Mel. Biol. IX, 310.— C. kamtschaticum var. alata Trautv. in Tr. Peterb. Bot. Sada, IX, 2 (1886) 478.—Ic.: Matsumura, Shokubutsumeii, No. 883; Gmel. Fl. sibirica, II, tab. 24. Perennial. Rhizome short, woody. Stem more or less sulcate, finely arachnoid-hairy, almost tomentose above, usually few-branched with 1—2 short branches in leaf axils, 50-200 cm high, leafy to tip. Leaves broadly ovate or ovate-lanceolate, subglabrous above, with more or less scattered, simple hairs, weakly arachnoid-hairy beneath, on veins, 181 178 venation prominent, coarsely pinnately-toothed to pinnately lobed, with tiny appressed spines on margin, toothed with tiny, erect spines; basal and lower cauline leaves 23-25 cm long, petiolate, including petiole 35 cm long and to 13 cm wide; pinnately incised to deeper than middle into antrorse, large, ovate, weakly toothed lobes, 4—5 on each side, basally narrowed into winged or long spiny-toothed petiole to 20 cm long, short-decurrent; middle cauline leaves basally narrowed, some- what decurrent, with more or less auriculate or long and broad wings, 15—18(38) cm long, (7)10—13(26) cm wide, pinnately divided to less than 1/2 into ovate lobes, 7 on each side; upper leaves reduced, semiamplexicaul, shallowly pinnately toothed or almost undivided, gla- brous or finely arachnoid-hairy; apical leaves 1—2, almost as long as capitula or somewhat longer, usually broadly lanceolate. Capitula soli- tary or 2—3, terminal on stem and long axillary branches, nodding at end of flowering, on tomentose, slightly drooping peduncles. Involu- cre more or less floccose-arachnoid-hairy, 1.5—2.5 cm in dia, at fruit- ing expanding to 5 cm, wide open; involucral bracts linear above, slightly deflected, finely spinescent, beneath finely, above weakly arach- noid-hairy, outer ones shorter than inner, about 17 mm long, middle ones usually brownish-reddish-brown or dark purple. Florets dark vio- let, rapidly turning brown, usually with white staminal tubes to 18 mm long; corolla tube 7 mm long; limb long, lobed to 2/3. Achenes red- dish-brown, weakly triquetrous, with thin apical tubercle, to 5 mm long; pappus to 15 mm long, brownish to reddish-brown, inner hairs apically long-scabrous, weakly papillate. Flowering July to September. Alpine shrub belt, birch forests, coastal meadows, as component of tall-herb community.—Far East: Okhotsk, Kamchatka (and Commander Islands), Sakhalin (and Kuril Islands). General distribution: Beringia (Aleutian Islands). Described from Kamchatka. Type in Leningrad. Note. The species has been understood broadly. It has included C. weyrichii Maxim. as a variety. Vassiliev refers this species to the Okhotsk-Kamchatka type of distribution area, considering it to be in the group of forest species. Evidently it is variable in the winged nature of the stem. Vassiliev reported var. alatum Trautv. for the Commander Islands. A white corolla has been reported for C. kamtschaticum Ldb. ex DC., V.L. Komarov (1930) reports for Kamchatka a dark purple co- rolla in this species, in contrast to the pure white staminal tubes that apparently impart a whitish cast to the florets in the capitulum. 84. C. weyrichii Maxim. Prim. fl. amur. (1859) 174; Makino in Tokyo Bot. Mag. XVIII, 299; Nakai in Tokyo Bot. Mag. XXVI, 361.— 182 179 C. kamtschaticum Kitam. Comp. Japon. I (1937) 41 p. p. non Ldb.— C. kamtschaticum 8. weyrichii Herder in Bull. Soc. Nat. Mosc. XLII, 1 (1870) 84.—Cnicus weyrichii Maxim. in Bull. Acad. Sc. Pétersb. XIX (1874) 495 and in Mél. Biol. IX, 310.—Ic.: Tolmatchev. O. FI. o. Sakhalina (1959) 56, Fig. 32. Perennial. Rhizome thick, with brownish scales and long, ligne- ous, fibrous roots. Stem 1—2 m high, hard, ascending, somewhat flexu- ous above, sulcate, purple, more or less sparsely hairy, leafy. Basal leaves sparsely crisped-hairy above, thinly arachnoid-hairy beneath, grayish-green; lower cauline leaves to 35 cm long, 18 cm wide, deeply sinuate to pinnately incised into elongate, broadly-lanceolate segments, remotely spinose-serrate-dentate, narrowed into short auriculate, spiny- ciliate petiole, decurrent as narrow wings; middle cauline leaves to 25 cm long, cordately narrowed below, pinnately incised to middle into oblong-lanceolate, acuminate segments, usually 4 on each side; upper ones oblong-lanceolate, narrowed below and incised or almost undi- vided, long-acuminate, subsessile; all leaves very finely appressed- spiny-ciliate, at apices of teeth and segments with spines 3 m long. Capitula 2—3.5 cm in dia, solitary, terminal at tips of stem and long branches, usually on tomentose peduncles, in paniculate-corymbose general inflorescence. Outer involucral bracts linearly narrowed from broad base, brown imbricate, scarcely arachnoid-hairy, outer and middle ones with narrowly filiform tip, short-spinescent, almost as long as inner; inner bracts linearly-lanceolate, narrowed into scarious tip. Corolla dark purple, 17—21 mm long; corolla tube to 7.5 mm long; limb 5-parted to 1/3. Achenes dark reddish-brown, 3.5—4.0 mm long, compressed, weakly triquetrous, basally narrowed, apically slightly ob- liquely truncate, with long, thin tubercle at apex; pappus pale reddish- brown, to 19 mm long, inner hairs longer than outer, with toothed, slightly thickened tips. Flowering July to September. (Plate IX.) One of main components of Sakhalin tall-herb community, which grows in zone of coastal and valley forests, as also in marshy places and scrubs.—Far East: Sakhalin (mostly southern part of island). General distribution: Japan (Nippon, Yezo). Described from vicinity of Due Port. Type in Leningrad. Note. The Sakhalin-Hokkaido element of the flora (according to Tolmatchev) has been combined by a number of authors with C. kamtschaticum Ldb. ex DC., from which it is well distinguished by the form of the leaves, involucral bracts, and lax inflorescence. 85. C. pectinellum A. Gray, On the Bot. Jap. (1859) 395; Nakai in Tokyo Bot. Mag. XXVI, 361; Kitam. Comp. Japon. I, 44.—C. pectinellum o. typicum Nakai and B. modestum Nakai in Tokyo Bot. 183 180 Mag. XXVI (1912) 361.—C. pectinellum var. mamiyanum Miyabe and Kudo, Fl. Saghal. (1915) 273.—C. mamiyanum Koidz. pl. Sachal. (1910) 124; Nakai in Tokyo Bot. Mag. XXVI, 321.—Cnicus pectinellus (A. Gray) Maxim. in Bull. Acad. Sc. Pétersb. XIX (1874) 491 and in Mel. Biol. IX, 358.—Ic.: Kitam. Comp. Japon. I (1937) pl. XV, fig. 2. Perennial. Rhizome horizontal, nodulose, thick, with ligneous, long, fibrous roots. Stem 50-150 cm high, ascending, sulcate, crisped-hairy, narrow-winged or sharply carinate, throughout or only below with long and fine yellowish spines on edges, leafy, with long internodes. Leaves deeply pinnatisect into linear segments, narrowed into more or less winged, thinly spinose petiole, decurrent as narrow wing or long spines; middle cauline leaves to 17 cm long, upper ones basally narrowed, subsessile, reduced, with short-decurrent, as narrow spinose wing, pinnatisect, acuminate; apical leaves almost undivided, linearly-lan- ceolate; all leaves glabrous above, more or less thinly grayish-arachniod- hairy beneath (especially upper leaves), very tiny spinose-ciliate or longer scattered spines; apical leaves linear, approximate to base of capitula, as long as them, or somewhat longer. Capitula nodding from base, solitary or in 2’s on short tomentose peduncles, forming corym- bose general inflorescence. Involucre at fruiting divergent, at flower- ing 1.5—2.0 cm in dia, more or less whitish-tomentose; involucral bracts linearly lanceolate, outer ones somewhat shorter than inner, narrowed into long, thin spinescent tip, deflexed in upper part, more or less floccose-arachnoid. Florets pink; corolla to 16 mm long; corolla tube to 6 mm long; limbs unevenly lobed up to middle. Achenes reddish- brown to brown, 4 mm long, with thin apically extended disk; pappus to 13 mm long, light reddish-brown, darker tips of inner modified to short-toothed bristles. Flowering July to August. Bogs, swampy birch forests, boggy meadows, forest edges, in foothills and intermontane valleys.—Far East: Sakhalin (southern part, Kuril Islands). General distribution: Japan (Nippon, Yezo). Described from Japan. Note. C. pectinellum A. Gray, with its finely incised leaves, spi- nose-winged stem, tomentose pubescence, and smaller capitula is well distinguished from C. weyrichii Maxim., which is widespread in the southern part of Sakhalin. A relatively large amount of material of this species was collected by M.G. Popov, among which there are speci- mens with finely incised leaves or broader leaf segments, which ap- proach C. weyrichii. The possibility is not ruled out that on Sakhalin these species hybridize with each other. Such specimens were col- lected by M.G. Popov near the villages of Klyuch and Kochuma. C. pectinellum A. Gray is known from Sakhalin on the basis of a small number of specimens from its southern part and needs further inves- 184 181 tigation. The specimen collected by E.G. Pobedimova and G. Konovalova on the Kuril Islands differs somewhat from the typical specimens and on closer study may prove to be a new species close to C. pectinellum A. Gray. Series 2. Erecta Kitam. in Acta Phytot. et Geobot. III, 1 (1934) 7 p. max. p.; Comp. Japon. I (1937) 46.—Capitula erect, aggregated at stem apices. Species of this series are distributed in Japan (Hokkaido, Hondo), in the north of Eastern Asia, Europe, and Beringia. From this series we exclude C. serratuloides (L.) Hill, which is close to species of the subsection Spanioptilon Kitam. 86. C. palustre (L.) Scop. Fl. Carn. ed. 2, II (1772) 128; Ldb. FI. alt. IV, 6; DC. Prodr. VI, 645; Ldb. Fl. Ross. II, 733; Fedtsch. and Fler. Fl. Evrop. Ross. 1010; Iljin in Fl. Yugo-Vost. VI, 403; Schischk. in Fl. Zap. Sib. XI, 2887; Klok. in Vizn. Rosl. URSR, 569; Hegi, Ill. Fl. VI, 2, 878.—C. palustre B. umbrosum Kauffm. Mosk. Fl. Ed. 2 (1889) 274; Fedtsch. and Fler. Fl. Evrop. Ross. 1011.—Carduus palustris L. Sp. pl. (1753) 822.—C. altissimus Gilib. Fl. Lithuan. III (1781) 186.—Cnicus palustris Willd. Fl. Berol. Prodr. (1787) 260; Sp. pl. Ill, 3, 1662.—Onotrophe palustris Cass. in Dict. Sc. nat. XXXVI (1825) 147.—Ic.: Gmel. Fl. sibirica, II (1749) tab. 23, fig. 2; Rchb. Ic. Fl. Germ. XV, tab. 831; Syreistsch. Ill. Fl. Mosk. Gub. III, 304; Fedtsch. and Fler. loc. cit. Fig. 1013; Javorka and Csapody, Iconogr. Fl. Hung. 541. fig. 3875; Hegi, 1. c. Taf. 271, fig. 4—Exs.: GRF No. 1671; FI. Gall. et Germ. exs. No. 677; Herb. norm. No. 84; Pl. Suec. exs. No. 1582. Perennial. Rhizome with funiform roots. Stem erect, 60-150 cm high, with sinuate-toothed wings, dark purple above, finely arachnoid- hairy, leafy, near apex usually with smaller leaves, occasionally branched from base to apex, with long leafy branches (var. umbrosum Kauffm.). Leaves linearly-lanceolate, 4-15 cm long, 0.5—3.0 cm wide, pinnately divided or pinnately incised into rhombic, lobate-toothed segments, scatteredly hairy above, more densely so beneath along veins with long, crisped hairs, thinly spiny-ciliate along margin. Rosulate leaves narrowed into short petiole; Cauline leaves narrowly sinuate, thinly spiny-ciliate decurrent wings, wings in upper part lacerated and long-spinose. Capitula 9-15 mm in dia, aggregated at apices of stem and branches, into corymbose-paniculate common inflorescence. In- volucral bracts scatteredly arachnoid-hairy along margin, dark purple above, outer ones ovate, to 2.5 mm long, spinescent, inner ones lin- early lanceolate, scarious at apex, colored. Florets to 14 mm long, 185 182 lilac-purple, corolla limb 8 mm long, lobed to 1/3 into narrowly linear lobes. Achenes smooth, stramineous or purple (f. erythrocarpum I\jin), 2.5—-3.5 mm long; pappus sordid-white, hairs thin, to 11 m long. Flow- ering April to September. Mostly in coniferous and deciduous forest zone, occasionally in forest-steppe; wet meadows, swamps and their edges, sometimes in scrubs.—European Part: Dvina-Pechora, Baltic Region, Ladoga-IImen, Upper Volga, Volga-Kama, Upper Dnieper, Middle Dnieper, Volga- Don, Trans-Volga, Upper Dniester, Bessarabia, Lower Don; Western Siberia: Ob Region, Upper Tobol, Irtysh; Eastern Siberia: Angara- Sayans (reported for Krasnodar Territory). General distribution: Scandinavia (to 67°50' N. Lat.), Central Europe, Atlantic Europe (north- ern part), Mediterranean Region (western), North America (introduced). Described from western Europe. Type in London. Note. Readily forms hybrids with species of the section Cirsium. From the European Part of the USSR, the following hybrids have been reported: 1. C. heterophyllum x C. palustre: Trautv. in Tr. Peterb. Bot. Sada, VIII, 2 (1883), Nos. 3105, 3117. 2. C. oleraceum X C. palustre: Trautv. ibid., Nos. 3116, 3118. 3. C. palustre x C. rivulare: Trautv., ibid., No. 3120. Subsection 7. Orthocentrum (Cass.) Charadze comb. nova; DC. Prodr. VI (1837) 641, pro sect.—Orthocentron Cass. in Dict. Sc. nat. XXVII (1823) 184, pro subgen., XXV (1825) 173 and XXXVI (1825) 480, pro gen.—Capitula ovate. Involucral bracts oblong-ovate, soft and scatteredly pilose or glabrous, imbricate, subulate or abruptly narrowed into thin, straight or slightly deflexed, more or less short spine. Florets red, very rarely whitish; corolla tube usually shorter than limb or longer. Anther filaments with papillose or fine hairs. Stem winged, with leaves decurrent on internodes, occasionally leaves sessile or scarcely decurrent, glabrous, weakly arachnoid-hairy or mostly grayish-tomentose, more or less deeply pinnately incised, occasionally weakly pinnately toothed. Type of subsection: C. siculum Spreng. Note. This subsection includes species distributed in West Asia, the Mediterranean Region, the Caucasus, and in the steppe regions of Eurasia. In the species of the subsection Orthocentrum (Cass.) Charadze, the filaments are finely papillate (visible under a strong magnifying glass) and not glabrous, as reported by Cassini. Evidently, on this 186 183 basis C. libanoticum DC. was referred by De Candolle to the section Onotrophe (Cass.) DC. (=Cirsium), in which finely hairy filaments are characteristic. In not attaching great significance to the nature of the pubescence of the filaments in the present case, I believe that the circumscription of the subsection should be broadened to include the species of the section Onotrophe (Cass.) DC. (= Cirsium) that are close in habit, with more or less winged stems and relatively shallowly incised leaves. Series 1. Pubigera Charadze.—Leaves more or less glabrous, pin- nately toothed or pinnately lobed; upper cauline leaves sessile, lower ones weakly decurrent with narrow wing. Capitula solitary or in clus- ters of 2—3, 15-30 mm in dia. This series includes Asia Minor species. One grows in Southern Transcaucasia, and the other is distributed in the Greater Caucasus. 87. C. depilatum Boiss. and Bal. in Boiss. Fl. or. III (1875) 543; Somm. and Lev. in Nuov. Giorn. Bot. Ital. N. S. II, 1, 18; Grossh. FI. Kavk. IV, 189; Charadze in Fl. Gruzii, VIII, 482.—C. pubigerum var. pubigerum Petrak in Mitteil. Thtiring. Bot. Gesellsch. I, 1 (1960) 26 p. p.—Exs.: Petrak, Cirsioth. Univ. No. 90. Perennial. Stem ascending, to 2 m tall, sulcate, sparsely arach- noid-hairy, branched above. Cauline leaves semiamplexicaul, with auricles weakly decurrent, oblong-ovate or ovate, apically narrowed, erose-toothed or lobed with spinescent teeth, scatteredly fine hairy on both sides or glabrous, sometimes sparsely floccose-arachnoid-hairy along veins; middle cauline leaves approximate, 16—23 cm long, 4—5 cm wide; apical leaves 3—6 cm long, distant. Capitula solitary, 15—30 mm in dia, at apices of stem and branches, projected above subtending leaves, sometimes forming lax paniculate general inflorescence. In- volucre subglabrous, with occasional, thin, arachnoid hairs; involucral bracts imbricate, outer ones 5-6 mm long lanceolate to ovate-lan- ceolate, spinescent, with scarcely deflexed spine, dorsally with black- ish-purple band, inner ones linear-lanceolate, narrowed into thin scari- ous tip. Florets pink; corolla 19-22 mm long, limb 12—14 mm long. Achenes reddish-brown, 3.5 mm long; pappus whitish, to 16 mm long, its inner hairs thin, somewhat long. Flowering August. Near streams, in depressions and on gravel beds, often forming thickets, in mid-montane and subalpine zones.—Caucasus: Southern Transcaucasia, Eastern Transcaucasia (former Borzhomi District, South- ern Ossetia). General distribution: Asia Minor. Described from Dzhimil in Lazistan. Type in Geneva. Note. Petrak (1960) considers C. depilatum Boiss. and Bal. iden- tical with C. pubigerum DC. He treats the latter species broadly and 187 184 includes a number of varieties, of which, for the Caucasus, he cites var. pubigerum, combining the original specimens of C. pubigerum DC., C. depilatum Boiss. and Bal., and C. elbrusense Somm. and Lev. However, in the Flora Orientalis Boissier distinguished C. pubigerum from C. depilatum by the smaller capitula and involucral bracts. The considerable material from the former Artvin and Oltu dis- tricts makes it possible to distinguish two species, one of which, with narrower, more or less decurrent or subsessile leaves and relatively small capitula, I consider to be C. pubigerum DC. In the Caucasus, I think it is possible to distinguish C. depilatum Boiss. and Bal. and C. elbrusense Somm. and Lev. Moreover, to C. pubigerum DC. we also refer the specimen from the former Oltu District cited by Petrak as C. pubigerum var. elbrusense (Somm. and Lev.) Petrak and mistakenly identified by him with the type of C. elbrusense Somm. and Lev. This same specimen was described by A.V. Fomin (/zv. Kavk. Mus. III, 1902, 281) as C. oltense Fom. 88. C. hygrophiloides Charadze in Zam. po Sist. i Geogr. Rast. Tbil. 22 (1961) 76.—C. uliginosum Grossh. Fl. Kavk. IV (1934) 187 p. p. non MB.—C. elbrusense Charadze in Fl. Gruzii, VIII (1952) 485 p. p. quoad pl. ex Achalchevi, non Somm. and Lev. Perennial. Stem hard, ascending, to 1.5 m high, sulcate, with oc- casional arachnoid hairs, branched. Leaves oblong, more or less deeply sinuate-pinnately lobed with deltoid, bifid lobes, strongly tapered into yellow, hard spines to 11 mm long, finely spinose, glabrous or sparsely arachnoid-hairy beneath; basal leaves oblong-elliptical, 30-33 cm long, 10-12 cm wide, acuminate, narrowed into short-winged petiole, on both sides, especially beneath, with scattered, thin, lanate hairs, deeply pinnately parted into elongate, distant and coarsely toothed, obtuse segments, with apical spines to 6 mm long; cauline leaves oblong-lanceolate, spinescent, more or less narrowed downward and with wings short-decurrent, 20-25 cm long, 7-9 mm wide, upper leaves reduced, somewhat surpassing capitula. Capitula 2—3 cm in dia, soli- tary or in clusters of 2—3, terminal on stem and branches, in corym- bose paniculate general inflorescence. Involucre with occasional arach- noid hairs; outer bracts broadly ovate to oblong-ovate, greenish, with broad blackish-purple band above, 7-9 mm long, subglabrous or with occasional arachnoid hairs, spinescent, inner lanceolate, narrowed into acute, scarious tip. Florets purple; corolla 21-22 mm long; limb to 12 mm long, lobed to 1/3. Achenes brownish, with light yellow bands, to 4 mm long; pappus 16-17 mm long, whitish, its inner hairs scarcely spreading racemosely at apex, somewhat longer than outer. Flowering July to August. 185 River terraces, near springs in wet places.—Caucasus: Ciscaucasia (mountains of Ingushetia, Northern Ossetia), Eastern Transcaucasia (upper reaches of Aragvi River). Described from eastern Georgia (Mlet drainage). Endemic. Type in Tbilisi. Note. It is distinguished from C. depilatum Boiss. and Bal. by more deeply incised leaves, short-spinescent involucral bracts, and darker-colored florets. From C. pubigerum DC. s. 1., it is distinguished by broader leaves and larger capitula. Series 2. Hygrophila Charadze.—Stem winged; leaves glabrous, decurrent, more or less deeply pinnately lobed. Capitula clustered at apices of stems, occasionally solitary, 15-25 mm in dia. 89. C. hygrophilum Boiss. Diagn. pl. or. ser. I, 10 (1849) 89; FI. or. III, 549; Charadze in Fl. Azerb. VIII, 410; Petrak in Mitteil. Thiiring. Bot. Gesellsch. II, 1, 24. Perennial. Stem ascending, branched, densely leafy, winged, with scattered occasional hairs. Leaves green above, glaucesent beneath, glabrous, with scattered simple hairs; cauline leaves linearly oblong, deltoid-sinuate and pinnately lobed, with bisected, toothed, approxi- mate lobes, 11 mm long, teeth long tapered to yellowish spine; apical leaves small, approximate to capitula, equalling surpassing them. Ca- pitula terminal or axillary, solitary, to 20 mm in dia, clustered at api- ces of stem and branches. Involucral bracts green, with purple band above, glabrous, sparsely floccose-pubescent; outer bracts oblong, with straight or somewhat deflexed, yellowish, spine 1.5 mm long, inner linearly lanceolate, with elongate, scarious cusp. Florets purple, 19 mm long; corolla limb to 11 mm long. Achenes to 4 mm long; pappus sordid white. Flowering July to August. Subalpine zone, in wet meadows and along mountain streams.— Caucasus: Southern Transcaucasia (Nakhichevan ASSR). General dis- tribution: Armenia and Kurdistan, Iran. Described from northern Iran. Type in Geneva. Note. This species was mistakenly cited by Petrak and later by A.A. Grossheim for the upper reaches of the Kuban River. The Desulavi specimen cited by Petrak (/zv. Kavk. Muz. VIII, 1-2, 69) from the upper reaches of the Kuban (Uchkulan) is referred to C. elbrusense Somm. and Lev., which is distinguished from C. hygrophilum by more vigorous growth, strongly branched stem in the upper part, capitula usually clustered at the stem tips and rising above the apical leaves. Later, Petrak (in Mitteil. Thiiring. Bot. Gesellsch. Il, 1, 24) allies C. hygrophilum Boiss. with C. pubigerum var. paphlagonicum Petrak; 188 this species is distinguished from the latter by broadly winged stems 186 that are densely leafy to the tip, long-spinose leaves, the upper of which surround the capitula and somewhat surpass them. 90. C. elbrusense Somm. and Lev. in Nuov. Giorn. Bot. Ital. N. S. II, 1 (1895) 16; in Tr. Peterb. Bot. Sada, XVI, 263; Grossh. FI. Kavk. IV, 188; Opred. Rast. Kavk. 486, quoad pl. kubanensem.—C. hygrophylum auct. non Boiss.: Petrak in Izv. Kavk. Muz. VII, 1-2 (1914) 69; Grossh. Fl. Kavk. IV, 185.—C. pubigerum var. elbrusense (Somm. and Lev.) Petrak in Mitteil. Thiiring. Bot. Gesellsch. II, 1 (1960) 20 p. p. quoad pl. e Kiikiirtli. Perennial. Stem winged, hard, ascending, sulcate, weakly branched above, densely leafy, glabrous or with occasional arachnoid hairs, to 1.5 m high, wings sinuate-lobed, spiny-toothed. Lower leaves to 23 cm long, narrowed into short petiole, pinnately lobed, with broad, toothed and spinescent lobes; middle cauline leaves oblong-lanceolate or ellip- tical, coriaceous, pinnately parted with 2-, 3-, occasionally 5-sect, elon- gate segments, lobes tapered to hard spines 5—7 mm long, glabrous on both sides, occasionally arachnoid-hairy, long-acuminate, auriculate, more or less long-decurrent, sinuate-toothed, more or less coarsely spiny-ciliate; upper leaves to 15 cm long, 6 cm wide. Capitula 1.5—2.5 cm in dia, solitary or 2—4 on short branches, at stem tips, subsessile in axils of long leaves; forming racemose-paniculate general inflores- cence; apical leaves at bases of capitula small, finely incised. Involu- cral bracts spinescent, outer ones ovate, to 5 mm long, scatteredly arachnoid-hairy in upper part, with blackish purple band; inner bracts linearly lanceolate, acuminate, with scarious tip. Florets purple; co- rolla 18-19 mm long; limb 11-12 mm long. Achenes to 3—4 mm long, light reddish-brown; pappus silvery-white, to 15 mm long. Flowering July to August. Mid-montane and subalpine zones, as small thickets, occasionally as solitary plants, in depressions in wet places and near springs.— Caucasus: Ciscaucasia (upper reaches of Kuban). Endemic. Described from Kyukyurtal in basin of Kuban River. Type in Florence. Note. This species was described from incomplete specimens from the Kyukyurtal Gorge. The absence of subsequent collections and the somewhat inaccurate description have created confusion in the inter- pretation of this species. For example, Petrak referred to it the Oltu specimen from the C. pubigerum DC. s. 1. complex; apparently, typi- cal specimens of C. elbrusense Somm. and Lev. from the Kuban were referred by the same author to C. hygrophilum Boiss., because of their absence in the region of the Main Range. New collections from the upper reaches of Kuban have confirmed the specific status of C. elbrusense Somm. and Lev., which is among a number of Elbrus 187 189 endemics. The Oltu specimen is well distinguished by its less incised, slightly decurrent leaves, and entirely glabrous, wingless internodes. Later, Petrak broadened the circumscription of C. pubigerum DC. by including C. elbrusense Somm. and Lev. as a variety, with which I cannot agree. 91. C. albowianum Somm. and Lev. in Tr. Peterb. Bot. Sada, XII, 1 (1892) 153, XVI (1900) 264; Petrak in Mitteil. Thiiring. Bot. Gesellsch. II, 1, 22; Grossh. Fl. Kavk. IV, 185.—C. svaneticum var. albowianum (Somm. and Lev.) Charadze in Fl. Gruzii, VII, (1952) 484. Perennial. Stem ascending, to 1 m high, sulcate, sparsely arach- noid-hairy, usually densely leafy, with spinose wings along edges, short branches above bearing 1—3 terminal capitula. Lower cauline leaves usually linearly lanceolate, narrowed at both ends, glabrous, very finely, scatteredly hairy only on midrib above, amplexicaul, decurrent, broad wings, usually sinuately-pinnately parted into deltoid-ovate, unequally 3—4—5S-toothed segments, their lobes basally deltoid or deltoid-lan- ceolate, spinescent, unevenly spiny-ciliate; middle and upper cauline leaves gradually reduced, like lower. Capitula at tips of stem and branches solitary or 2, of which 1 subsessile, undeveloped, on strongly reduced peduncles, globose-ovate, to 15 mm in dia, with 1 short-de- current, linearly-lanceolate, distant, spinose toothed apical leaf. In- volucral bracts glabrous or only basally scatteredly arachnoid-hairy, outer ones ovate-oblong, abruptly tapered to straight or horizontally deflexed, thin spine, like middle bracts, dorsally with blackish-purple glandular band above; inner bracts elongate, gradually narrowed into long scarious tip. Florets reddish-violet, 20-25 mm long; limb 5-lobed to middle, 2 times as long as corolla tube. Achenes unknown; pappus somewhat shorter than florets, sordid-white, hairs plumose, apically tiny setose and somewhat thickened. Flowering August. Upper forest zone, at about 2,000 m.—Caucasus: Western Transcaucasia (Upper Svanetia). Endemic. Described from Svanetia. Type in Florence. Note. Known only from the collections of Sommier and Levier. Judging from the diagnosis and detailed description of this species given by Petrak, it stands closest to C. elbrusense Somm. and Lev., differing only by solitary and not clustered, terminal capitula, as in the latter. The species needs to be investigated further. Series 3. Elodea Charadze.—Stem winged. Leaves glabrous or finely arachnoid hairy. Capitula to 15 mm in dia. 190 188 92. C. apiculatum DC. Prodr. VI (1837) 642; Bornm. in Beih. Bot. Centralbl. XX, Abt. II, 2, 162; Charadze in Fl. Turkm. VII, 244.— C. apiculatum ssp. glaberrimum Petrak in Oest. Bot. Zeitschr. LX, 2 (1910) 460.—C. libanoticum Boiss. Fl. or. HI (1875) 548 p. p. non DC.—Exs.: Sint. It. transcasp.-pers. No. 960. Perennial. Stem ascending, hard, branched above, to 100 cm tall, winged, without or with tiny scattered hairs, wings more or less deeply orbiculate-lobate, with short and long, hard, yellowish spines to 8 mm long. Leaves glabrous, glaucesent, oblong- or linearly lanceolate, pin- natifid to pinnately lobed, with orbicular lobes, acuminate; basal leaves narrowed into long petiole, together 28 cm long; lower cauline leaves to 10 cm long, 2.5 cm wide; all leaves auriculate, decurrent on petiole as lobate wings, acuminate, with short and long yellowish spines. Capitula to 15 mm in dia, solitary at tips of reduced peduncles and 3-5 clustered at tips of stem and long branches, forming racemose- umbellate general inflorescence. Involucral bracts ovate-lanceolate to linearly lanceolate, pale yellow, with dark spot above, tapered to slightly recurved spines 2.5—3 mm long lacking or with occasional arachnoid hairs. Florets pale pink, to 18 mm long. Achenes pale yellow, to 3.5 mm long; pappus with thin, basally scaly hairs, to 15 mm long. Flow- ering July to August. Swampy meadows, along mountain streams.—Soviet Central Asia: mountainous Turkmenia (Karaul, Sulyuki, Kusika). General distribu- tion: Armenia and Kurdistan, Iran. Described from Iran from collec- tion of Belangé. Type unknown. Note. The species is close to C. elodes MB. and is distinguished by stiffer, narrower leaves, spines crowded at the tips of stem and branches, larger capitula, and weakly colored involucral bracts. The specimen of this species collected by Bornmiiller along the border with Iran was referred by him first to C. elodes as var. glaberrium Bornm. and later to C. apiculatum DC. Boissier combined C. apiculatum DC. and C. libanoticum DC. under the latter name, adding to it, as a synonym, C. kotschyanum Boiss., described, like C. libanoticum DC., from Lebanon. The specimen, distributed by Kotschy (Pl. Pers. austr. No. 789) as C. apiculatum DC., has a critical anno- tation by Boissier, which reveals that C. apiculatum DC. is not present in the Herbarium of De Candolle and the distributed specimen was referred to the above species only on the basis of the diagnosis. Later, Boissier (F/. or.) cites this specimen under the name C. libanoticum DC. In the Herbarium of the Botanical Institute of the Academy of Sciences of the USSR, one specimen from Iran (from Bunge’s herbarium) is preserved, which we are inclined to consider identical with the type of C. apiculatum DC. From the specimens collected by 19 — 189 Kotschy (C. kotschyanum Boiss., C. apiculatum DC., and also C. libanoticum DC. from Hamadan), this specimen differs in several characters and is similar to Bornmiiller’s specimen from Sulyukli. The major differences of C. libanoticum DC. are its more or less dense, finely pilose stems and leaves, narrower racemose inflorescence, and others. I am inclined to consider C. apiculatum DC. as a species re- placing C. libanoticum DC. in northern Iran and Turkmenia, the range of which covers the mountains of Lebanon and southern Iran. 93. C. elodes MB. FI. taur.-cauc. III (1819) 555, excl. syn. Gmel.: C.A.M. in Mém. Acad. Sc. Pétersb. VI, sér. sc. nat. VI, 52; Charadze in Fl. Azerb. VIII, 409.—C. desertorum auct. fl. cauc. non Fisch. ex Link: Petrak in Izv. Kavk. Muz. VIII, 1-2 (1914) 73; Grossh. FI. Kavk. IV, 185; Charadze in Fl. Gruzii, VIII, 486.—C. setigerum Grossh. Opred. Rast. Kavk. (1949) 484, non Ldb.—Cnicus palustris var. B. MB. FI. taur.-cauc. II (1808) 274. Perennial. Root funiform or slightly tuberculate. Stem ascending, weakly ribbed, 35-90 cm high, yellowish, glabrous, occasionally scatteredly hairy below, leafy to tip and sinuately broadly winged, from middle with more or less paniculate-cymose, long curved branches, with 3—5(7) small capitula clustered at their tips. Cauline leaves gla- brous, smooth or tuberculate, with thin, hard spines, 3-5—7 mm long, oblong-elliptical; basal leaves 15-16 cm long, 3.5 cm wide, narrowed into short petiole, toothed or shallowly-pinnately lobed, more or less spiny-ciliate; lower cauline leaves like basal leaves, decurrent with broad wing; leaves from middle to upper gradually reduced, oblong- lanceolate to linearly-lanceolate, decurrent; apical leaves deeply pin- nately lobed, with broad, sharply toothed segments, smaller, linear. Capitula 8—-10(12) mm in dia, 12-15 mm long, clustered on short, small-leaved and winged, spinose peduncles, forming corymbose-pan- iculate generai inflorescence. Involucral bracts small, lower ones 3 mm, middle ones to 7 mm long, oblong-lanceolate, lustrous, stramineous, with purple-brownish subapical band, tapered thin, weakly decurved, more or less elongate spines, weakly arachnoid-hairy; inner involucral bracts linearly-lanceolate, narrowed into scarious, weakly recurved tip. Florets red, to 17 mm long; corolla tube to 7 mm long; limb unevenly 5-parted up to middle. Achenes 2.5-3.0 mm long, stramineous; pappus sordid-white with thin, entirely plumose outer hairs and scabrous-tipped inner hairs. Flowering July to August. Wet places, roadsides, in lowlands and foothills, in steppe and semidesert regions.—Caucasus: Ciscaucasia (eastern part), Eastern Transcaucasia, Southern Transcaucasia. General distribution: Armenia and Kurdistan, Iran and Afghanistan (northern). Described from vicin- ity of Shemakha. Type in Leningrad. 192 190 Note. This species replaces C. alatum (S.G. Gmel.) Bobr. in the eastern part of Ciscaucasia and Transcaucasia. The possibility of its occurrence in regions adjoining Southern Transcaucasia is also not ruled out. It is distinguished by entirely glabrous stems and leaves and particularly by the form of the involucral bracts, which are gradually narrowed into thin spines and are lacking the floccose pubescence along the margin characteristic of C. alatum (S.G. Gmel.) Bobr. 94. C. alatum (S.G. Gmel.) Bobr. in Bot. Zhurn. SSSR, XLIII, 11 (1958) 1547.—C. desertorum Fisch. ex Link, Enum. pl. hort. Berol. II (1822) 300; Spreng. Syst. veg. 3, 371; Ldb. FI. Ross. II, 734; C.A.M. in Mém. Acad. Sc. Pétersb. VI, sér., sc. nat. VI, 50; Trautv. in Bull. Soc. Nat. Mosc. XXXIX, 2, 376; in Tr. Péterb. Bot. Sada, VIII, 1, 500; Schischk. in Fl. Zap. Sib. XI, 2888.—C. desertorum var. subintegerrima Trautyv. and var. sinuatolobuta Trautv. in Bull. Soc. Nat. Mosc. XXXIX, 2 (1866) 376.—C. setigerum Ldb. Ic. pl. Fl. Ross. I (1829) 9; Fl. alt. IV, 5; DC. Prodr. VI, 642; Ldb. FI. Ross. II, 734; C.A.M. op. cit. 51; Iljin in Fl. Yugo-Vost. VI, 404.—C. elodes auct. fl. Ross. eur. non MB.: Ldb. op. cit. II (1846) 733 p. p.; O. and B. Fedtsch. Perech. Rast. Turk. IV, 284; Fedtsch. and Fler. Fl. Evrop. Ross. 1011; Klok. in Vizn. Rosl. URSR, 569.—C. elodes B. setigerum Kryl. Fl. Alt. III (1904) 692.—C. elodes var. sinuato-lobatum and var. subintegerrimum O. and B. Fedtsch. op. cit. (1911) 284.—Serratula alata S.G. Gmel. Reise, I (1770) 155.—Carduus desertorum Fisch. ex Steud. Nomencl. bot. ed. 2, I (1841) 283.—Ic.: Rchb. Iconogr. bot. tab. 986, No. 1317; Ldb. Ic. pl. Fl. Ross. tab. XXXV; S.G. Gmel. Reise, I, tab. XXXV. Perennial. Rhizome long-tuberous or with funiform roots. Stem solitary, straight, more or less branched only in upper part, yellowish, sulcate, pubescent, especially below, with more or less scattered crisped hairs, 30-100 cm high, leafy to tip, winged to middle; wings lacerated in upper part, entire or weakly sinuately toothed, lobed in upper part of stem, with hard, tiny and longer yellowish spines. Leaves coria- ceous, ovate-elliptical to oblong-lanceolate, almost undivided (var. subintegerrimum (Trautv.) Charadze) or sinuately-lobed (var. sinuato- lobatum (Trautv.) Charadze), subentire or bicrenate-dentate, ciliate from short, hard, and thin spines up to 3 mm long, verrucose above, scatteredly pilose or more or less glabrous beneath; basal leaves more or less broadly oblong-lanceolate, narrowed into short, narrowly-winged petiole, subacute, weakly toothed, scatteredly pilose or subglabrous, to 30 cm long and 4 cm wide; lower cauline leaves like basal, decurrent on stem by broad wings; middle and upper leaves strongly reduced, oblong, more or less deeply sinuately lobed, long-spinose; apical leaves 193 191 on elongate branches of corymbose inflorescence small, shortly-decur- rent, with lobed wings. Capitula 11-15 mm in dia, solitary or 2-3 at tips of stem and branches, forming corymbose-paniculate general in- florescence. Involucral bracts, from outer ovate and oblong-ovate to inner linearly-lanceolate, greenish-yellow, with purple to reddish-brown band above and more or less floccose-white hairy; outer bracts tapered into hard, straight or slightly deflexed, short spine, compactly imbri- cate; inner bracts narrowed into scarious, colored, thin point. Florets pink; corolla 18—19 mm long, corolla tube 7—8 mm long, limb 5-parted almost to middle. Achenes to 3 mm long, pale yellow, smooth, lus- trous; pappus of fewer hairs, basally white, slightly brownish, 12—13 mm long, inner hairs with slightly thickened, short-setose tips. Flow- ering July to August. Wet alkaline meadows, in lowlands and foothills, in steppe and semidesert zone.—European Part: Volga-Kama, Middle Dnieper, Volga-Don, Trans-Volga, Bessarabia, Lower Don, Lower Volga; West- ern Siberia: Upper Tobol, Irtysh; Soviet Central Asia: Aralo-Caspian Region, Baltic Region, Syr-Darya, Tien Shan. Described from Lower Don (Pyatiiz-benskaya). Type in Leningrad. Note. The name C. alatum (S.G. Gmel.) Bobr. was first proposed by E.G. Bobrov, who restored Gmelin’s name for this plant in a new combination. He excluded this species from the Caucasian species C. elodes MB., which is usually identified with C. alatum under the name C. desertorum Fisch. ex Link and sometimes with C. setigerum Ldb. At the same time, authors of the Russian flora have treated this same plant under the name C. elodes MB., described from the Caucasus. Moreover, Serratula alata S.G. Gmel. was treated by, Marschall- Bieberstein as a synonym of C. elodes MB. and then by Meyer as a synonym of C. setigerum Ldb. At present, it seems better to treat the southern vicariant races of C. alatum as separate species: the Cauca- sian one, C. elodes MB., and the Irano-Turkmenian one, C. apiculatum DC. Despite great variation, these species are distinguished by a suite of small morphological differences imparting to them a distinctive ap- pearance. I agree with E.G. Bobrov that the nature of the thickening of the root fibers from funiform to tuberous, on which an attempt was made to distinguish C. desertorum from C. setigerum, evidently does not have much importance, since it is found to be mixed all over its range. Series 4. Leucopsia Charadze.—Plants with grayish or white arach- noid hairs. Leaves more or less arachnoid-hairy on both sides or only beneath, broadly decurrent. Capitula 18—25 mm in dia. 194 192 95. C. svaneticum Somm. and Lev. in Nuov. Giorn. Bot. Ital. N. S. II, 1 (1895) 20; in Tr. Peterb. Bot. Sada, XVI (1900) 265; Petrak in Tr. Tifl. Bot. Sada, XII, 1, 30; in Izv. Kavk. Muz. VIII, 1-2, 63; Grossh. Fl. Kavk. IV, 185; Charadze in Fl. Gruzii, VIII, 484, excl. var.—C. svaneticum var. ramosum Somm. and Lev. op. cit. (1895) 20; op. cit. (1900) 266; Grossh. op. cit. 185.—C. leucopsis var. caucasicum Somm. and Lev. Nuov. Giorn. Bot. Ital. N. S. II, 1 (1895) 20.—C. leucopsis Petrak in Mitteil. Thiiring. Bot. Gesellsch. II, 1 (1960) 29 p. p. non DC.—Exs.: Petrak, Cirsioth. Univ. No. 91. Perennial. Stem tall, sulcate, many-branched above or from middle (var. ramosum Somm. and Lev.), arachnoid-hairy, to 1.5 m high, lac- erate-winged. Leaves soft, coriaceous, green, floccose, and finely arach- noid hairy above, grayish or whitish-, more or less densely arachnoid- hairy to tomentose beneath, sinuate-toothed or pinnately lobed, more of less long-spinose; lower leaves elongated, broadly-lanceolate or sinuately-lobed almost to middle, with bifid, deltoid ovate lobes, nar- rowed into petiole, decurrent on stem by lacerated wings, to 25 cm long; middle leaves oblong-lanceolate, apiculate, shallowly sinuately- lobed, with broadly deltoid, toothed lobes, spinulose, basally narrowed, decurrent on petiole, 15-18 cm long; apical leaves apically toothed, teeth tapered to more or less long, thin, yellowish spines, linear-lan- ceolate, acuminate. Capitula 18-25 mm in dia, solitary or 2-3 at tips of stem and branches. Involucre subglabrous, basally weakly arachnoid-hairy; outer involucral bracts ovate or ovate-lanceolate, spinescent, subglabrous, sparsely arachnoid-hairy along margin, with black or purple band above; inner bracts lanceolate, narrowed into scarious tip. Florets purple-violet; corolla 15—16 mm long, limb 9 mm long. Achenes dark reddish-brown, 3.5 mm long; pappus to 12 mm long, with whitish hairs, basally brown. Flowering July to August. Upper forest and subalpine zones, in clusters or small thickets near streams, along forest edges and on alluvium.—Caucasus: Ciscaucasia (western part and Elbrus District), Western Transcaucasia (Abkhazia, Svanetia, Gornaya Racha). Endemic. Described from Svanetia. Type in Florence. Note. It varies greatly in the form and pubescence of the leaves; the mid-montane forms have narrower, deeply incised leaves, subglabrous above; the subalpine forms have broad, shallowly incised leaves, grayish-arachnoid-hairy above. Petrak (1960) unites C. svaneticum Somm. and Lev. with the Asia Minor C. leucopsis DC. undoubtedly a very closely related species. However, C. leucopsis DC., is distinguished by smaller capitula (13-17 mm across), pale reddish florets, and a corolla more deeply cut than in C. svaneticum, as well as by coriaceous leaves. The isolated range of our species in the 195 193 secluded gorges of Western Transcaucasia and the Central Caucasus is cut off from the range of the Asia Minor C. leucopsis DC. 96. C. uliginosum (MB.) MB. FI. taur.-cauc. IIT (1819) 555; Boiss. Fl. or III, 544; Somm. and Lev. in Nuov. Giorn. Bot. Ital. N. S. II, 1, 19; Grossh. Fl. Kavk. IV, 187; Petrak in Mitteil. Thtiring. Bot. Gesellsch. II, 1, 29.—Cnicus uliginoisus MB. F1. taur.-cauc. II (1808) 274. Perennial. Stem tall, to 1.5 m high, hard, ascending, sulcate, branched above, usually sparsely and inflorescence more densely arach- noid-hairy, with internodes to middle spinose-winged. Leaves oblong- ovate, to 17 mm long, 8 cm wide, narrowed toward tip, shallowly sinuate to pinnately lobed, narrowed toward tip, shallowly sinuate to pinnately lobed, with subacute, bisected lobes tapered into hard spines 3—5 mm long; leaves glabrous above or weakly arachnoid-hairy, gray- ish-arachnoid-hairy beneath, more or less long decurrent; upper leaves small, linear, deeply pinnately incised, not surpassing capitula. Ca- pitula at tips of elongate stem and branches, in clusters of 2—4(5), occasionally solitary, 12-120 mm in dia. Involucral bracts oblong- lanceolate; outer ones 5—6 mm long, with purple band above, with short weakly deflected, spinous tip, weakly arachnoid-hairy, inner ones linear-lanceolate, narrowed into scarious tip. Florets purple; corolla 16-17 mm long, limb 10-11 mm long. Achenes dark brown, to 3 mm long; pappus silvery-white, to 13 mm long. Flowering August. - Depressions, in wet places near streams in mid-montane and sub- alpine zones.—Caucasus: Ciscaucasia (central part). Endemic. De- scribed from vicinity of Kislovodsk. Type in Leningrad. Note. This species was erroneously cited for several districts of Transcaucasia instead of C. hygrophiloides Charadze or C. oblongifolium C. Koch, from which it is well distinguished by leaves that are more or less grayish-arachnoid-hairy beneath and decurrent on the stem. Petrak (Petrak in Mitteil. Thiiring. Bot. Gesellsch. II, 1 [1960] 29) allies C. uliginosum (MB.) MB. with C. kusnetzowianum Somm. and Lev. and C. sychnosanthum Petrak with which, however, it has very little in common, especially in the shape of involucral bracts. Series 5. Subinermia Charadze.—Stem winged. Leaves more or less grayish-arachnoid-hairy or whitish-tomentose. Capitula to 15 mm in dia. 97. C. subinerme Fisch. and Mey. in Mém. Acad. Sc. Pétersb. VI, sér., sc. nat. VI (1849) 55; Charadze in Fl. Azerb. VIII, 393.—C. duriusculum C. Koch in Linnaea, XXIV (1851) 410.—C. elodes auct. 196 194 non MB.: DC. Prodr. VI (1837) 642; Ldb. Fl. Ross. II, 733; Boiss. FI. or. III, 547; Somm. and Lev. in Nuov. Giorn. Bot. Ital. N. S. II, 1, 15; Grossh. Fl. Kavk. IV, 185.—C. elodes B. indivisum DC. Prodr. VI (1837) 642; Boiss. Fl. or. III, 547; Somm. and Lev. op. cit. 15; Petrak in Vestn. Tifl. Bot. Sada, Nov. Ser. 2, 42; Grossh. Fl. Kavk. IV, 185.—C. leucopsis B. indivisum C.A.M. and y. cinarescens C.A.M. Cirs. Rut. (1849) 54.—C. leucopsis var. indivisa Trautv. in Tr. Peterb. Bot. Sada, II (1873) 556. Perennial. Rhizome with string-like roots. Stem erect, to 75 cm high, branched above, leafy to tip, more or less broadly-winged, gray- ish-arachnoid-hairy. Leaves green above, scatteredly whitish-verrucose, puberulent, occasionally finely arachnoid-hairy, beneath grayish- or more or less densely arachnoid-hairy, almost entire or coarsely toothed to pinnately lobed (var. duriusculum (C. Koch) Charadze), oblong, to 35 cm long, 9 cm wide, narrowed into elongate winged petiole; cauline leaves ovate, oblong-ovate or broadly-lanceolate, 12—25 cm long, 4— 10 cm wide, more or less long-tapered in upper part, acuminate, more or less decurrent on stem with broad, entire, occasionally lacerated, thin, spinulose wings; apical leaves usually not surpassing capitula. Capitula 12-15 mm in dia, in clusters of 3—7 at tips of stem and branches forming corymbose-paniculate general inflorescence. Involu- cral bracts small, outer ones 3-4 mm long, ovate or ovate-oblong, narrowed into short spine, glabrous, with blackish apical macula, finely floccose-arachnoid-hairy, inner bracts linearly-lanceolate, narrowed into scarious, purple tip Corolla purple, to 15 mm long; limb 9 mm long, lobed to more than 1/3. Achenes to 3 mm long; pappus sordid white, to 11 mm long, with thin and sparse hairs, inner hairs surpassing outer, with scabrous tips. Flowering July to September. Mid-montane and subalpine zones, in ravines, occasionally in for- ests, in moist places.—Caucasus: Eastern Transcaucasia (southern part), Southern Transcaucasia. General distribution: Asia Minor, Armenia and Kurdistan. Described from Dilizhan. Type in Leningrad. Note. In the form of the leaves, pubescence, and a number of other characters, this species sharply differs from C. elodes MB. Its range is restricted to the southern part of Transcaucasia and Lazistan and in Asia Minor to the adjoining part of eastern Anatolia. The spe- cies is highly variable in the spinosity and incision of the leaves. The name C. subinerme Fisch. and Mey. must take priority for this species, although under this new name an extreme shade form has been de- scribed, which is almost devoid of spines on the lamina. The form described by C. Koch from eastern Anatolia under the name C. duriusculum C. Koch has a more xeromorphic habit. The Lagov plants preserved in the Herbarium of the Botanical Institute of the Academy of Sciences of the USSR from Dagestan and 197 195 the vicinity of Tbilisi are doubtful and have not been confirmed by later collections. Hybrid forms are known, which have been described by Petrak as hybrids between C. elodes and C. obvallatum from the Gyunei shore of Lake Sevan (Shakhdag Mountain). They are distinguished by sev- eral characters that are intermediate between the parent species. 1. C. subinerme Fisch. and Mey. x C. obvallatum (MB.) MB.—C. elodes X C. obvallatum f. delizhanicum Petrak, f. obvallatum Petrak and f. subelodes Petrak in Vestn. Tifl. Bot. Sada, Nov. Ser. 2 (1926) 45—48. —Leaves broad, shallowly sinuate-toothed, weakly arachnoid- hairy beneath, cuneate and short-decurrent on stem. Capitula at apices of stem and widely deflected branches, in clusters of 2—5, 13-17 mm in dia; involucral bracts spinescent with slightly deflected or recurved spine, surrounded by smaller, pinnately incised apical leaves, surpass- ing capitula. Southern Transcaucasia (Gyunei). 98. C. rhabdotolepis Petrak in Vestn. Tifl. Bot. Sada, Nov. Ser. 2 (1926) 49; Grossh. Fl. Kavk. IV, 186; Opred. Rast. Kavk. 485. Perennial. Stems large, ascending, to 1 m high, sulcate, densely leafy, more of less densely grayish arachnoid-hairy, winged, with sinu- ate-toothed, long-ciliate and thinly spinose, arachnoid-hairy wings, above with reduced axillary branches. Cauline leaves elongately ellip- tical, acuminate, basally narrowed, decurrent on stem, with more or less broad, densely ciliate-spinose wings, spines to 12 mm long; lower and middle cauline leaves 12-15 cm long, 4-6 cm wide, pinnately parted into ovate-deltoid, approximate segments with 2—3-fid, deltoid, spinescent lobes with yellowish spines to 5-7 mm long, coarsely crenate-toothed, ciliate-spinous, grayish-arachnoid-hairy above, white- arachnoid-hairy or loosely tomentose beneath. Capitula clustered at apices of branches and forming racemose general inflorescence; ca- pitula numerous at stem apices, subsessile, in axils of smaller apical leaves, usually not surpassing them, or 3-4, at tips of axillary branches, usually not longer than cauline leaves or somewhat longer; involucre subglabrous, 10-14 mm in dia; involucral bracts small, outer and middle ones ovate to ovate-lanceolate, glabrous, very sparsely arachnoid-hairy, narrowed into short erect spine, with blackish-purple band above; in- ner bracts linearly-lanceolate, narrowed into scarious, purple tip. Flo- rets purple, 16-17 mm long; corolla limb 9-10 mm long, lobed to 1/3. Achenes light-reddish-brown, to 3 mm long; pappus sordid white, with thin hairs, to 12 mm long. Flowering July to August. At altitudes of 2,100—2,200 m, on slopes near springs.—Caucasus: Southern Transcaucasia (vicinity of Nor-Bayazet and Gyunei shore of 198 196 Lake Sevan). Endemic. Described from Nor-Bayazet. Type in Erevan; isotype in Tbilisi. Subsection 8. Aciculata Charadze in Addenda XXVII, 605.— Leaves with thin, short and long spines on margins, pinnately parted or almost entire. Stem simple or weakly branched above, usually leafy to tip. Capitula medium-sized, basally with few pinnate-spinose apical leaves, shorter than capitula or much longer, surrounding capitula like involucre. Involucral bracts ovate to linearly-lanceolate, narrowed into short spine or point. Type of subsection: C. obvallatum (MB.) MB. Note. The species of the subsection are distributed in the Caucasus, Asia Minor, and to some extent in West Asia. The origin of these species is associated with the Caucasian Mountain region from where they could have spread to the south. In the development of the thin ciliate spines on the leaves and the long thin spines of the floral bracts, the species of the subsection approach the Central Asian species of the section Echenais (Cass.) Petrak, but in other characters, especially the structure of involucral bracts, they must be related to the section Cirsium and in habit stand closer to the subsection Acaulia Rouy. 99. C. obvallatum (MB.) MB. FI. taur.-cauc. III (1819) 559; C.A.M. Verzeichn. 70; DC. Prodr. VI, 661; C. Koch in Linnaea, XXIV, 408; Ldb. Fl. Ross. II, 744; Boiss. Fl. or. III, 541; Somm. and Lev. in Nuov. Giorn. Bot. Ital. N. S. II, 1, IJ; Grossh. Fl. Kavk. IV, 186; Charadze in Fl. Gruzii, VIII, 512; in Fl. Azerb. VIII, 412.—C. obvallatum &. pinnatipartitum C. Koch and B. pinnatilobum C. Koch, loc. cit. (1851).—Cnicus obvallatus MB. FI. taur.-cauc. II (1808) 279.— Exs.: Petrak, Cirsioth. Univ. Nos. 92, 134; Herb. Fl. cauc. No. 200. Perennial. Rhizome nodulose, oblique, with numerous long, ligne- ous roots. Stem 100—150 cm high, ascending, with short branches above, more or less densely crisped-hairy, leafy to tip, brownish-purple. Leaves ovate-oblong or elliptical, sinuate to pinnately parted to middle or deeper into ovate, 2-fid or coarsely toothed lobes, more or less long- spinose, with deltoid-ovate, teeth narrowed into yellow, hard spines, 5—7 mm long, with isolated tiny spines between teeth, glabrous and bright-green above, with whitish prominent venation, pale green be- neath, more or less densely crisped-hairy along veins; basal leaves to 30 cm long, 10 cm wide, pinnately incised into approximate, oblong- ovate segments, basally narrowed into more or less long, spatulate petiole; cauline leaves sessile, with broad, amplexicaul auricles, middle leaves pinnately parted, 10-15 cm long, 4-8 cm wide; upper leaves shallowly pinnately lobed, with approximate, long-spinescent teeth; 199 197 apical leaves at base of capitula many, linearly-lanceolate or linear, whitish-green, toothed ciliate, with long and thin spines, surrounding involucre as spathe and usually not surpassing it. Capitula solitary or in clusters of 2—3 at tips of stem and branches. Involucre globose- ovate, to 2—2.5 cm in dia, glabrous; involucral bracts imbricate, from outer ovate-oblong to inner oblong-lanceolate, glabrous, stramineous beneath, weakly carinate above, greenish-brown, spinescent with yel- lowish spine. Florets 18 mm long, whitish or pale yellow; corolla with narrow tube 8.5 mm long, limb lobed almost to 2/3. Achenes 4.5 mm long, with thin, reddish-brown streaks; pappus 16 mm long, slightly brownish- to light reddish-brown or sordid-white, with thin, sparse, plumose hairs, inner hairs with filiform tips. Flowering July to Sep- tember. Subalpine zone, as a component of tall-herb vegetation, in moun- tain pastures as weed.—Caucasus: throughout high-mountain regions. General distribution: Asia Minor, Armenia and Kurdistan, Iran. De- scribed from Kabarda. Type in Leningrad. Note. It is variable in the form of the cauline leaves and the length and number of the flora bracts. Var. iranicum Bornm. (Parsa, Fl. Iran, III, 1943) has been described for Iran. C. obvallatum (MB.) MB. readily hybridizes with species of the sections Cirsium and Echenais. Among them the following merit men- tion. 1. C. subinerme Fisch. and Mey. x C. obvallatum (MB.) MB.—C. elodes x C. obvallatum, 1) f. delizhanicum Petrak in Vestn. Tifl. Bot. Sada, Nov. Ser. 2 (1926) 45, 2) f. obvallatiforme Petrak, ibid. 46, and 3) f. subelodes Petrak ibid. 47. Caucasus: Southern Transcaucasia. All three forms are described from Shakhdag Mountain on the Gyuinei shore of Lake Sevan (collec- tions of Grossheim and Edelmeyer). The three hybrids described by Petrak from Bozdag are interme- diate between C. obvallatum (MB.) MB. and C. subinerme Fisch. and Mey. The following can be considered intermediate characters: more or less decurrent leaves, varying from almost entire to deeply pin- nately divided; spinosity of leaves; nature of their pubescence, from almost glabrous to more of less arachnoid-hairy, yellowish color of the corolla in specimens lacking characteristic, broadened apical leaves; or the presence of apical leaves surpassing capitula. 2. C. simplex C.A.M. x C. obvallatum (MB.) MB.— C. x kozlowskyi Petrak in Izv. Kavk. Muz. VIII, 1-2 (1914) 76; Kolak. Fi. Abkhazii, 1V (1949) 276.—Exs.: Pl. or. exs. No. 174. 200 198 Caucasus: Western Transcaucasia (Abkhazia), Southern Transcaucasia (Bakuriani). The hybrid was described by Petrak from specimens from Bakuriani and later observed by A.A. Kolakovsky in the Katsirkha Range together with the parental species. Kolakovsky notes a wide range of variation in the hybrid forms and the presence of a series of intermediates from the one species to the other. Variation in the form of the leaves has been reported—from narrower as in C. simplex, to relatively broad and deeply pinnately divided, which approaches C. obvallatum, but usually almost glabrous beneath. With reddish florets and the presence of dark brown bands on the involucral bracts, the hybrid forms are closer to C. simplex and with the broader leaf lobes look like C. obvallatum. 100. C. simplex C.A.M. Verzeichn. (1831) 70; in Mém. Acad. Sc. Pétersb. VI, sér., sc. nat. VI, 49; DC. Prodr. VI, 652; Ldb. FI. Ross. II, 742; Boiss. Fl. or. HI, 542, excl. var.; Petrak in Tr. Tifl. Bot. Sada, XII, 1, 29, 45; in Vestn. Tifl. Bot. Sada, 24, 12; in Izv. Kavk. Muz. VIII, 1-2, 62, excl. spp.; Grossh. Fl. Kavk. IV, 188, excl. var.; Charadze in Fl. Gruzii, VII, 511.—Exs.: Petrak, Cirsioth- Univ. No. 132. Perennial. Rhizome with string-like roots, densely covered with remnants of basal leaves. Stem ascending, 15-50 cm high, weakly flexuous, simple, lacking simple hairs, but with scattered crisped hairs, often purple, leafy to tip. Leaves glabrous on both sides, entire or sinuate-toothed, densely ciliate, with thin, more or less long spines; basal leaves numerous, broadly-lanceolate or oblong-obovate to lin- early-lanceolate, 9-25 cm long, 17—25 mm wide; cauline leaves like basal, gradually reduced, toward stem tip, surpassing internodes; apical leaves lanceolate, long-spinose, as long as capitula or somewhat sur- passing them. Capitula solitary at tips of stem, or 2-3, subsessile, obconical, with florets 2.5 cm long, and 1.5 cm in dia in middle. Involucral bracts usually dark purple, from outer ovate to middle ob- long-lanceolate, short-spinescent; inner bracts linearly-lanceolate, nar- rowed into scarious tip. Florets purple-red, to 20 mm long; limb to 12 mm long. Achenes light reddish-brown, 3.5 mm long; pappus silvery- white, slightly yellowish basally, with few, thin, apically scabrous hairs. Flowering July to August. Alpine and occasionally subalpine zone, in swampy meadows, near streams, on overgrown moraines, usually to 2,700 m.—Caucasus: Ciscaucasia, Eastern and Western Transcaucasia. Endemic. Described from Western Caucasus. Type in Leningrad. 101. C. armenum DC. Prodr. VI (1837) 647; Somm. and Lev. in Nuov. Giorn. Bot. Ital. N. S. II, 1, 17.—C. simplex B. armenum Boiss. 20 — 199 Fl. or. II] (1875) 543.—C. simplex ssp. armenum Petrak in Izv. Kavk. Muz. VIII, 1-2 (1914) 63.—C. simplex var. armenum Grossh. Fl. Kavk. IV (1934) 188.—Exs.: Sint. It. Orient. No. 7169. Perennial. Rhizome horizontal, short. Stem leafy, 30-45 cm high, sulcate, purple, simple, with scattered arachnoid hairs. Leaves linearly- lanceolate, subglabrous or with more or less scattered arachnoid hairs on both sides, pinnately lobed, with deltoid- or rotund-toothed lobes, densely long ciliate-spinous, usually with longer, hard, and short thin spines; basal leaves to 25 cm long and 2.8 mm wide, narrowed at base into winged petiole; cauline leaves sessile, with auricles decurrent on petioles, acuminate, surpassing internodes; apical leaves at base of inflorescence, somewhat surpassing it. Capitula at tips of stems, in clusters of 4—5, occasionally solitary, 2.3-2.8 mm in dia, subsessile, forming racemose-capitate general inflorescence. Involucral bracts elongately lanceolate, greenish, subglabrous or with weakly scabrous, narrowed into elongate, yellowish spine; inner bracts linearly lanceolate, spinulose. Florets yellowish; corolla about 18 mm long; limb about 12 mm long. Achenes 3.5 mm long; pappus yellowish-white, with few thinly setose hairs at apex. Flowering July to August. Swampy meadows, in subalpine zone.—Caucasus: Southern Transcaucasia. General distribution: Asia Minor, Armenia and Kurdistan. Described from Erzerum. Type in Paris; isotype in Leningrad. Note. The isolated range together with stable morphological char- acters, such as, for example, the yellow color of the corolla, dentition of the leaves, and the presence of sturdier spines, give this species a distinct appearance, readily distinguishing it from C. simplex C.A.M. The latter species extends to the region of the Western Caucasus and Transcaucasia. C. armenum DC., described from eastern Anatolia, geographically replaces C. simplex C.A.M. Subsection 9. Acaulia Petrak in Beih. Bot. Central-bl. XXXV, II, 2-3 (1917) 258; Kitam. in Acta Phytot. et Geobot. III, 1 (1934) 3.— Plants stem less or almost stem less, occasionally with distinct stem, and then stem short, leafy, simple. Capitula clustered at stem tip, occasionally solitary, usually more or less numerous, in center of basal rosette, usually with floral leaves at base, occasionally without them, globose or ovate-oblong, medium- or large-sized. Involucral bracts scatteredly hairy or subglabrous; outer and inner involucral bracts usually appressed, with straight or somewhat falcately recurved tip, short-spinescent, occasionally with long, hard spine, sometimes scabrous and ciliate on margin from middle to tip, gradually longer from outer to inner; inner bracts linear, long-acuminate, sometimes with unevenly incised-toothed, more or less expanded ligulate tip. Pappus almost as 202 200 long as florets, scarcely longer or shorter. Achenes not more than 4 mm long. Type of subsection: C. acaule (L.) Scop. The species of the subsection are distributed in North America and Eurasia. 102. C. esculentum (Sievers) C.A.M. in Mém. Acad. Sc. Pétersb. VI, sér., sc. nat. VI (1849) 42 (excl. var. B.) diagn.; in Beitr. Pfl. Russ. Reich. 5, 43; Nym. Consp. FI. Eur. I, 407; Korsh. Tent. Fl. Ross. or. 247; Schischk. in Fl. Zap. Sib. XI, 2894.—C. esculentum q. sibiricum C.A.M. op. cit. (1849) 43, a) Acaule C.A.M. op. cit. 43, and b) Caulescens C.A.M. op. cit. 44.—C. esculentum var. acaulis Trautv. in Bull. Soc. Nat. Mosc. XXXIX, 2 (1866) 377; Schischk. op. cit. 2895.— C. gmelini Tausch in Flora, XI, 31 (1828) 482; Turcz. Fl. baic.-dahur. II, 1, 127.—C. acaule Ldb. FI. alt. TV (1838) 11, excl. syn. p. max. p. non Scop.—C. acaule f. sibiricum Ldb. Fl. Ross. II (1846) 743; Schmalh. Fl. II, 106.—C. acaule y. gmelini DC. Prodr. VI (1837) 652.—Cnicus esculentus Sievers in Pall. Neust. Nord. Beitr. II (1796) 362; Georgi, Beschr. Russ. Reich. III, 4, 1225.—Cnicus gmelini Spreng. Hist. Rei Herb. II (1808) 270.—Ic.: Gmel. Fl. sibirica, II (1749) tab. XXV. Perennial. Rhizome reduced, turnip-shaped, with numerous fibrous roots. Stem to 35 cm high (var. caulescens Trautv.), sulcate, crisped- hairy or plants stemless (var. acaule Trautv.). Basal leaves numerous, narrowed into spinose-toothed elongate petiole, elongately oblong-lan- ceolate, 12-30 cm long, on both sides densely covered with flexuous hairs, arachnoid-hairy beneath along midrib, more or less deeply pin- nately incises into broad, ovate-deltoid crenate-toothed lobes, spinu- lose along margin, teeth, spinulescent; cauline leaves smaller than basal, somewhat decurrent, with spinose-toothed petiole; apical leaves at base of capitula 1-3, small, almost as long as capitula or somewhat longer. Capitula clustered at stem tip in capitate inflorescence, occasionally solitary, axillary, on short peduncles (var. caulescens Trautv.), often subsessile, clustered in center of basal rosette, usually 5—12 (var. acaule Trautv.), oblong, to 3 cm in dia, cm long. Involucre glabrous, with elongate bracts; outer bracts ovate-lanceolate to broadly-lanceolate, lower bracts to 10 mm long, gradually elongating inward, narrowed into long obtuse tip, spinulescent, yellowish or greenish, sometimes purple, denticulate-ciliate; inner bracts elongate, more or less linearly- lanceolate, narrowed into recurved scarious point. Florets red, to 27 mm long; corolla tube to 14 mm long; limb unequally 5-fid. Pappus sordid-white, to 28 mm long, almost as long as florets or somewhat longer, with numerous filiform hairs; achenes to 4.5 mm long, yellow- 203 201 ish-grayish, compressed, indistinctly finely ribbed throughout. Flower- ing July to September. Steppe region, on solonchaks, in wet saline meadows, along river banks, in low-herb vegetation of mountain meadows.—European Part: Volga-Kama, Middle Dnieper, Volga-Don, Trans-Volga, Black Sea Region, Lower Don, Lower Volga; Western Siberia: Upper Tobol, Irtysh, Altai; Eastern Siberia: Angara-Sayans, Dauria; Soviet Central Asia: Lake Balkash Region, Dzhungaria-Tarbagatai. General distribu- tion: Dzhungaria-Kashgaria, Mongolia (northern). Described from Soviet Central Asia (Lake Balkhash Region). Type in Leningrad. Note. It is distinguished from the closely related species distrib- uted in Eurasia by the numerous capitula clustered at the stem tip, elongate, more or less large involucral bracts, and pappus a little longer than the florets with fruits. The following hybrid is known: C. vulgare (Savi) Ten. x C. esculentum (Sievers) C.A.M.—C. ispolatovii Iljin (C. lanceolatum x C. esculentum) in Izv. Bot. Sada Akad. Nauk SSSR, XXX, 3-4 (1932) 351.—By its tall stem and decurrent leaves and also by the form of the involucral bracts it is closer to C. vulgare (Savi) Ten. It resembles C. esculentum (Sievers) C.A.M. in having the ca- pitula clustered in a dense inflorescence at stem tip.—European Part: Trans-Volga (Buguruslan District). 103. C. rhizocephalum C.A.M. Verzeichn. (1831) 70; in Mém. Acad. Sc. Pétersb. VI ser., sc. nat. VI (1849) 46; DC. Prodr. VI, 651; Ldb. FI. Ross. II, 744; Boiss. Fl. or. III, 542; Bornm. in Beih. Bot. Centralbl. XX, Abt. II, 2, 162; Grossh. Fl. Kavk. IV, 488; Charadze in Fl. Gruzii, VIII, 512; in Fl. Turkm. VII, 242.—C. rhizocephalum B. caulescens Boiss. 1. c. (1875) 542; Lipsky, Fl. Kavk. (1899) 359.—C. rhizocephalum y. sinuatum Boiss. loc. cit. p. p. (excl. syn.).—C. grumosum Fisch. and Mey. in C.A.M. op. cit. (1849) 48.—C. esculentum B. caucasicum C.A.M. op. cit. (1849) 44; C. Koch in Linnaea, XXIV, 405.—C. esculentum var. caucasicum Grossh. Fl. Kavk. IV (1934) 188.—C. esculentum var. caucasicum f. caulescens Grossh. Fl. Kavk. IV (1934) 188.—C. esculentum ssp. caucasicum (C.A.M.) Petrak in Tr. Tifl. Bot. Sada, XII, 1 (1912) 45.—C. acaule Ldb. FI. Ross. II (1846) 743 p. min. p. non Scop.—Exs.: Herb. Fl. cauc. No. 198; Petrak, Cirsioth. Univ. No. 131. Perennial. Rhizome short, thick, with cluster of string-like roots. Plants stemless, with capitula clustered in center of rosette in inflorescence, or stem short, to 25 cm high (var. caulescens Boiss.), simple or with short branches, weakly angular, more or less densely crisped-hairy with articulate hairs. Basal leaves numerous, rosulate, 20 os 202 elongately elliptical or oblong-lanceolate, 7-25 cm long, 2—3 cm wide, short-acuminate, narrowed into short narrow-winged petiole with thin yellowish spines, 3—5 (10) mm long, pinnately parted into approxi- mate, roundish or deltoid-ovate, indistinctly 2—3-fid lobes, toothed, abruptly narrowed into more or less long spine to 10 mm long, usually shallowly-pinnately parted, occasionally almost entire, subglabrous above or with scattered flexuous hairs along midrib, pubescent be- neath; cauline leaves distant, linearly-lanceolate, lobate, long-spinose, basally narrowed, with narrow wing decurrent on stem; apical leaves at base of capitula linearly-lanceolate or narrowly-linear, long pinnate- spinose, usually 2—3, as long as capitula or surpassing them. Capitula ovate, 1.5—3.5 cm in dia, in clusters of 3—5(7) in center of basal ro- sette, often forming racemose general inflorescence. Involucre gla- brous, imbricate, usually greenish or purple above; bracts lanceolate, outer ones appressed, narrowed into thin, yellowish, spines, 5 mm long, sparsely ciliolate from middle to tip along margin; inner bracts linearly-lanceolate, gradually longer from middle to inner, narrowed into short, purple tip. Florets 25—27 (28) mm long, pink, occasionally whitish; limb slightly shorter than corolla tube, inequally 5-fid to middle. Pappus at end of flowering and fruiting almost as long as florets, usually to 27 mm long, sordid-white, with numerous, apically plumose, filiform: hairs; achenes yellowish, with reddish-brown streaks, to 4 mm long. Flowering July to September. Mid-montane to alpine zone, in wet places, usually as weed in low-herb meadows and secondary carpet-like meadows of subalpine and alpine zones.—Caucasus: throughout Caucasus; Soviet Central Asia: mountainous Turkmenia. General distribution: Armenia and Kurdistan, Iran. Described from Eastern Caucasus (Tfandag). Type in Leningrad. Note. It varies greatly in the leaf lobing and the length of the spines of the outer involucral bracts and leaves; stemless forms pre- dominate. From C. esculentum (Sievers) C.A.M., to which this species was referred in part by authors of the Caucasian flora as var. caucasicum C.A.M., it is distinguished by the form of the narrower bracts, usually tapered to more or less elongate spines, fewer capitula, a pappus not longer than the florets, and other small differences. From the European species C. acaule (L.) Scop., it is well distinguished by the root sys- tem and the imbricate, narrower outer involucral bracts. C. frickii Fisch. and Mey., described from the Caucasus, differs from this species by having somewhat taller growth and larger terminal capitula. C. rhizocephalum C.A.M., reported for eastern Anatolia and Kurdistan, has been cited for Iran (Parsa) and also for Afghanistan (Petrak, Kitamura). Possibly, C. brevipapposum Tschern. described 205 203 from Soviet Central Asia and close to C. rhizocephalum C.A.M., grows in Afghanistan. Specimens resembling C. brevipapposum Tschern. are also known from the Caucasus, and, possibly, they were cited by Meyer as C. esculentum B. caucasicum a) rhodanthum C.A.M. (op. cit. 1849, 44). 104. C. frickii Fisch. and Mey. in Mém. Acad. Sc. Pétersb. VI sér., sc. nat. VI (1849) 47; Grossh. Fl. Kavk. IV 188; Charadze in FI. Gruzii, VIII, 512.—C. esculentum ssp. frickii Petrak in Tr. Tifl. Bot. Sada, XII, 1 (1912) 29. Resembles the preceding species; differs from it in the following characters: stem well developed, to 50 cm high; capitula solitary, ter- minal, to 5 cm in dia, and smaller axillary; strongly elongated leaves, pinnately divided in broadly-deltoid, toothed, long-spinose lobes; yel- lowish(?) florets. Flowering August. Forest zone.—Caucasus: Southern Transcaucasia. Described from southern Georgia (Tsalka). Endemic. Type in Leningrad. 105. C. brevipapposum Tschern. in Fl. Uzbek. VI (1962) 517, 373, Plate XXXIV, Fig. 2.—C. acaule auct. non Scop.: O. and B. Fedtsch. Perech. Rast. Turk. IV (1911) 248 p. p.—C. esculentum auct. fl. As. Med. et Cauc. p. p. non C.A.M. Perennial. Rhizome short, thick. Basal leaves numerous, deeply (up to midrib) sinuate-pinnately divided into 4—5-toothed ovate-del- toid lobes; teeth subacute, with thin, hard spines, usually to 5 mm long; cauline leaves thinly spinose-ciliate, glabrous on both sides, on midrib crisped-hairy, more or less long-acuminate, on short, long-spi- nose petioles, oblong-lanceolate, lower leaves 10-12 cm long, up to 3 cm wide. Capitula to 2 cm in dia, in clusters of up to 5 in center of basal rosette. Involucre more or less purple; outer involucral bracts glabrous, smooth along margin or scabrous from middle; middle bracts to 14 mm long, ovate-oblong, weakly narrowed above into elongate, obtuse, spinescent tip with scarcely noticeable spine; inner bracts to 20 mm long, linearly lanceolate, glabrous, acute. Corolla to 23 mm long; limb to 10 mm, inequally 5-fid almost to 1/2. Pappus shorter than florets, to 17 mm long, sordid-white, reddish-brown to brown below, with few more or less equal hairs, inner hairs with filiform tips. Flowering July to August. Mountain meadows.—Caucasus (rarely in high-mountain regions); Soviet Central Asia: Tien Shan, Pamiro-Alai. Described from pamiro- Alai. General distribution: ?Asia Minor. Type in Tashkent. Note. It differs from C. rhizocephalum C.A.M. by having involu- crat bracts that are smooth along the margin, with a scarcely noticeable 206 204 apical spine and reddish florets. From C. esculentum (Sievers) C.A.M., it differs by having a short pappus and smaller capitula. 106. C. acaule (L.) Scop. Fl. Carn. ed. 2, II (1772) 131 (acaulos); Lam. Fl. Fr. II (1778) 26 (acaulos); All. Fl. Pedem. I, 153; DC. Prodr. VI, 652 p. p. excl. var.; Koch, Syn. (1837) 398, ed. 3, II (1902) 1544; Fedtsch. and Fler. Fl. Evrop. Ross. 1012.—C. acaule var. genuina Briq. and Cav. in Burnat. Fl. Alp. Mar. VII (1931) 47.—C. acaule var. acaule Arénes in Bull. Jard. Bruxelles, XXIV, 4 (1954) 267.—C. allionii Sch. and spenn. FI. Frib. III (1829) 1079.—Carduus acaulis L. Sp. pl. ed. 2 (1763) 1156.—Cnicus acaulis Willd. Fl. Berol. Prodr. (1787) 260; Willd. Sp. pl. III, 3 (1803) 1681.—Onotrophe acaulis Cass. in Dict. Sc? nat. XX XVII (1825) 148.—Ic.: Rchb. Ic. Fl. Germ. XV (1852) tab. 840, I and II (var. caulescens); Coste, Fl. Fr. II (1903) 375; Fiori and Paol. Ic. Fl. Ital. (1904) 453; Fedtsch. and Fler. loc. cit. Fig. 1014; Javorka and Csapody, Iconogr. Fl. Hung. 540, fig. 3872; Hegi, Ill. Fl. VI, 2, fig. 577, 578.—Exs.: Hult. Pl. Suec. exs. (1946) No. 1584. Perennial. Rhizome oblique, somewhat thickened, ligneous, with long string-like roots. Plants almost stemless or with stems 7-9 cm high. Stems leafy, more or less crisped-hairy, solitary of 2—3, occa- sionally elongate shoots in center of basal rosette with solitary termi- nal capitula. Leaves rosulate, 12—16(20) cm long, oblong, to 3 cm wide, sinuate-pinnately lobed almost to base, with approximate, round- ish, 2—3-fid lobes, often with subacute strongly spinescent teeth with spines to 5 mm long, setose-spinose; leaves narrowed into strongly reduced long-spinose petiole, subacute; cauline leaves 6.5—10.0 cm long, up to 2.5 cm wide, sessile, subglabrous or scatteredly hairy above, more or less densely pubescent with crisped, articulate, brownish hairs beneath on veins; upper cauline leaves small, 2-4 cm long. Capitula subsessile, in rosula of basal leaves, solitary or 2—3, on short pe- duncles, oblong, 1.3—2.3 cm in dia, 3.0-3.3 cm long. Involucre imbri- cate; outer bracts ovate or ovate-lanceolate, subobtuse, spinulescent, ciliolate on upper margin; inner bracts linearly-lanceolate, 2 times as long as middle, slightly broadened at apex, scarious, subacute, with distinct spine. Florets red; corolla to 32 mm long; corolla tube about 20 mm long, limb inequally 5-parted to deeper than middle. Pappus 25 mm long, sordid-white, whitish above, with apically scabrous inner hairs; achenes 3—4 mm long. Flowering August to September. Dry low-herb meadows, pastures, along forest edges, heathers, roadsides, from lowlands to alpine zone.—European Part: Baltic Re- gion, Ladoga-Ilmen, Upper Dnieper. General distribution: Scandinavia, Central Europe, Atlantic Europe, Mediterranean, Balkans (northwest- ern). Described from western Europe. Type in London. S AT TNS 20 [e,<) 206 Note. It differs from the closely related species by having outer involucral bracts that are shorter than the inner ones and 1-3 capitula on each plant, and from C. esculentum (Sievers) C.A.M. it is distin- guished also by the pappus, which is not as long as the florets, and by an unthickened rhizome. Section 6. Pseudo-eriolepis (Nakai) Kitam. in Acta. Phytot. et Geobot. III, 1 (1934) 2; Comp. Japon. I, 37.—Pseudo-eriolepis Nakai in Tokyo Bot. Mag. XXVI (1912) 357, pro subgen.—Breea Sch. Bip. ex Maxim. in Bull. Acad. Sc. Pétersb. XIX (1874) 510 and in Mél. Biol. IX, 331 p. p. pro subgen. non Breea Less. (1832) pro gen.— Capitula nodding, lacking leaves. Involucre ovate, basally bulged; involucral bracts deflexed, linear, spinescent, outer 1/2 as long as inner. Florets with long filiform corolla tube, 4—5 times as long as limb almost to base 5-parted. Pappus at fruiting almost as long as florets. Basal leaves withering early; cauline leaves not decurrent on stem, pinnatifid. Type of section: C. pendulum Fisch. Note. Of the two eastern Asian species of this section, only one is widespread in Eastern Siberia and in the Far East. C. pendulum Fisch., in the structure of florets, exhibits affinity to the section Cephalonoplos (Neck.) DC. With its large nodding capitula and form of the involucral bracts and leaves, it approaches members of the sub- section Sinocirsium Kitam. 107. C. pendulum Fisch. in DC. Prodr. VI (1837) 650; Ldb. FI. Ross. II, 739; Maxim. Prim. fl. amur. 173, excl. syn. B. oligocephalum Trautv. and Mey.; Rgl. Tent. Fl. Ussur. 95; Turcz. Fl. baic.-daur. II, 1, 126; Korsh. in Tr. Peterb. Bot. Sada, XII, 2, 361; Kom. Fl. Man’chzh. III, 749; Nakai in Tokyo Bot. Mag. XXVI, 357; Kom. and Alis. Opred. Rast. Dal’nevost. Kr. II, 1085; Kitam. Comp. Japon. I, 37; Popov, FI. Sr. Sib. II, 752.—Cnicus pendulus Maxim. in Bull. Acad. Sc. Pétersb. XIX (1874) 510 and in Mél. Biol. IX, 332.—Ic.: Makino, Nature Col. Wild. Pl. II (1933) 82, var.; Terasaki, Jap. Bot. Hlustr. Album (1933) 487. Perennial. Rhizome with numerous ligneous roots. Stem hard, 1-2 m high, sulcate, weakly angular, more or less branched, sparsely hairy below, more or less densely arachnoid-hairy above, thinly gray- ish-tomentose below capitula. Leaves glabrous, larger below, numer- ous, deeply twice pinnately incised into unevenly sinuate-toothed lobes, glaucesent beneath, spiny-setose, teeth and lobes narrowed into spines to 4-5 mm long; basal leaves withering early, oblong-elliptical, 25—35 cm long, 7-8 cm wide, gradually narrowed into petiole; lower cauline 209 207 leaves to 15 cm long, short-petiolate or subsessile, upper leaves re- duced, to 5 cm long, sessile. Capitula nodding from base, numerous, in racemose-paniculate inflorescence in upper part of stem and on axillary branches. Involucre basally with roundish prominance, at be- ginning of flowering 1.5—2.5 cm in dia, at fruiting 3.5—5.0 cm in dia, scarcely arachnoid-hairy; involucral bracts narrowly-linear or linearly- subulate, dark green or brownish-purple, spinulescent; inner bracts narrowly-linear, membranous, apically scarious, slightly deflected. Florets red, numerous, usually somewhat exserted from capitula; co- rolla to 23 mm long or slightly shorter, with long, narrow tube, 3—4 times as long as broad limb more than 1/2 lobed. Pappus 20-21 mm long, light reddish-brown; achenes 3.5 mm long, brownish, above with narrow border, basally narrowed. Flowering July to September. Moist meadows, forest edges, near streams, in river valleys, along banks of peat bogs.—Eastern Siberia: Angara-Sayans (Irkutsk, Balagansk), Dauria; Far East: Zeya-Bureya (Amur-Bureya Lowland), Uda River area (Lower Amur), Ussuri (along Amur and Ussuri rivers), Sakhalin. General distribution: China (northeastern), Japan (Nippon, Yezo), Korean Peninsula. Described from Dauria. Type in Geneva; isotype in Leningrad. Note. C. pendulum Fisch. stands alone among the eastern Asian species of the genus, distinguished by narrow, elongate, outer involu- cral bracts, a very narrow corolla tube several times as long as its limb, and numerous capitula nodding from the base. Japanese bota- nists attach a somewhat exaggerated importance to these characters. For example, Nakai (1912) places this species in the separate subge- nus Pseudo-eriolepis Nakai, although Kitamura (1934) considers it possible to treat Pseudo-eriolepis as a section. Nodding capitula are not typical of either the European or the Siberian species of the genus, but are found in the eastern Asian species as well as in the group of Caucasian species usually placed in the Mediterranean-Western Asian section Epitrachys DC. and segregated by us as the section Caucasigena Charadze. C. pendulum Fisch. in the structure of its corolla approaches species of the C. arvense (L.) Scop. alliance. Regel and Tilling de- scribed a variety 8. oligocephalum Rgl. and Till. from Ayana; this variety, in fact, should be transferred to C. schantarense Trautv. and Mey., which was correctly pointed out already by Herder. Varieties a. genuinum Herd., B. microcephalum Herd. and y. recurvum Regl. described by Herder in C. pendulum apparently have no taxonomic importance. Section 7. Cephalonoplos (Neck.) DC. Prodr. VI (1837) 643; Endl. Gen. pl. (1836-1840) 478; Turcz. Fl. baic.-dahur. II, 1 (1856) 125; O. 210 208 Hoffm. in Pflanzenfam. IV, 5 (1893) 322.—Cephalonoplos Neck. Elem. Bot. I (1790) 68, pro gen.; Nevsky in Tr. Bot. Inst. Akad.. Nauk SSSR, Ser: 1, 4 (1937) 290; Kitam. Comp. Japon. I (1937) 134.—Cirsium subgen. Cephalonoplos Nakai in Tokyo Bot. Mag. XXVI (1912) 355.— Cirsium sect. Breea (Less.) Boiss. FI. or. III (1875) 525.—Breea Less. Syn. Comp. (1832) 9.—Cnicus Benth. in Benth. and Hook. f. Gen. pl. II (1873) 468 p. p.—Cnicus subgen. Breea Sch. Bip. ex Maxim. in Bull. Acad. Sc. Pétersb. XIX (1874) 510 and in Mél. Biol. IX (1874) 331 p. p.—Carduus Baill. Hist. des Plantes, VIII (1882) 74 p. p— Plants dioecious; capitula homogamous with numerous florets. Involu- cre oblong; involucral bracts many-rowed, imbricate, appressed, outer ones with short spinescent tip, inner ones elongate, apically scarious. Limb in staminate florets 5-parted to more than 1/2 corolla tube 2 times as long as limb; filaments papillose or glabrous. Pistillate florets filiform; corolla tube 4—5 times as long as limb, 5-parted almost to base. Pappus at flowering shorter than corolla, at end of flowering elongating and at fruiting longer than florets; achenes 4 mm long. Perennial plants with rooted underground stem; stem strongly branched above. Type of section: C. arvense (L.) Scop. Note. The majority of authors treats Cephalonoplos as part of Cirsium. The structure of florets characterized by a long and narrow corolla tube, which is several times as long as the limb, development of unisexual florets in the capitula, and dioecy, serves as the justifica- tion of some authors, such as Kitamura (1937, 1939) and Nevsku (1937), for placing species of the section Cephalonoplos in a separate genus. Among present-day Cirsium, we find transitions to species of Cephalonoplos. The closest to them in the length of the pappus and structure of the corolla is the eastern Asian C. pendulum Fisch., which otherwise has little in common with this group. A pappus almost as long as florets is also characteristic of the members of the subsection Acaulia Petrak. Evidently, C. arvense (L.) Scop. often forms hybrids with members of the section Cirsium, which also provides a basis for putting this group of species in a separate genus. A study of the pollen of the genus Cirsium (Shtepa, 1961) also showed that in the structure of the pollen the species of this section should be referred to Cirsium Mill. 108. C. setosum (Willd.) MB. FI. taur.-cauc. III (1819) 560 p. p. excl. pl. cauc.; DC. Prodr. VI, 643; Ldb. Fl. alt. IV, 10; Schischk. in Fl. Zap. Sib. XI, 2896 p. max. p.—C. setosum B. subulatum Ldb. FI. alt. IV (1833) 10.—C. setosum var. subulatum Schischk. op. cit. 2897.—C. arvense auct. p. p. non Scop.: Ldb. Fl. Ross. II (1846) 734; 209 Klok. in Vizn. Rosl. URSR, 569.—C. arvense y. setosum Ldb. FI. Ross. II (1846) 753; Kryl. Fl. Alt. III (1904) 697.—C. arvense C. setosum Schmalh. FI. II (1897) 106.—C. arvense var. setosum Kom. Fl. Man’chzh. III (1907) 749; Fedtsch. and Fler. Fl. Evrop. Ross. 1012; Nakai in Tokyo Bot. Mag. XXVI, 355; Miyabe and Miyake FI. Saghal. (1915) 272.—C. arvense ssp. setosum Iljin in Fl. Yugo-Vost. VI (1936) 408.—C. arvense y. setosum f. subulatum Ldb. F1. Ross. II (1846) 735.—C. arvense y. integrifolium Wimm. and Grab. FI. Siles. II, 2 (1829) 92; Koch, Syn. (1837) 400.—C. laevigatum Tausch in Flora, XI, 31 (1828) 483 (excl. syn. Gmel.).—C. argunense DC. Prodr. VI (1837) 644; Turcz. Fl. baic.-dahur. II, 1, 125.—Cnicus setosus Bess. Prim. Fl. Galic. II (1809) 172.—Cnicus arvensis y. setosus Maxim. in Bull. Acad. Sc. Pétersb. XIX (1874) 51 and in Mel. Biol. IX, 333.—Serratula setosa Willd. Sp. pl. III, 3 (1803) 1664.— Cephalonoplos setosum (MB.) Kitam. in Acta Phytot. et Geobot. III, 1 (1934) 8; Comp. Japon. I, 135.—Ie.: Gmel. Fl. sibirica, II (1749) tab. XXII; Rchb. Ic. Fl. Germ. XV (1852) tab. 842, II; Sorn. Rast. SSSR, IV (1934) 287, Fig. 477. Perennial. Rhizome vertical, robust, often with numerous rooted branches. Stem erect, (40)50-130 cm high, robust, weakly sulcate, usually ribbed, arachnoid-hairy above, glabrous below, with upward- spreading branches. Basal leaves withering after flowering; cauline leaves crowded in middle of stem, oblong-lanceolate or ovate-lan- ceolate, (5)10—20 cm long, 1.7—5.5 cm wide, obtuse, with tiny spines on margin, basally narrowed gradually, sessile, sometimes with winged lobes in lower part of stem, short-decurrent (var. subulatum Ldb.), undivided or coarsely and distantly sinuate-toothed tiny spiny-ciliate along margin, glabrous on both sides, usually green above, occasion- ally scatteredly somewhat floccose-arachnoid-hairy, pale green beneath or arachnoid-hairy; cauline leaves gradually reduced above, oblong- lanceolate or lanceolate, basally cuneately narrowed, spiny-ciliate, in inflorescence strongly reduced, usually undivided, 1.5 cm long, 0.5 cm wide. Capitula unisexual, numerous, in lax corymbose inflorescence, on slender peduncles, erect; staminate capitula ovate, to 13 mm long, 10 mm in dia, pistillate ones to 20 mm in dia, 16—20 mm long. Involu- cre weakly arachnoid-hairy, purple, basally obtuse; involucral bracts appressed, imbricate, lowermost strongly reduced, oblong-lanceolate, subacute, spinulescent, middle ones lanceolate, like outer brownish- purple above, narrowly scarious on margin; inner bracts linear, apically scarious, somewhat broadened, acuminate. Female florets 17 mm long, somewhat broadened, acuminate. Female florets 17 mm long, with corolla tube 3—5 times as long as limb, limb 5-parted almost to base. Pappus shorter than corolla, to 11 mm long, at fruiting to 22-24 mm; 212 210 pappus hairs brown, apically deflected, slightly thickened, plumose; _ achenes yellowish to reddish-brown, 2.5 mm long, compressed. Flow- ering July to August. Forest and steppe zones; forest edges, riverbanks, on post-forest meadows, weed in fields, crops, along roads, old fields —European Part: Dvina-Pechora, Ladoga-IImen, Volga-Kama, Trans-Volga, Lower Volga; Western Siberia: all regions; Eastern Siberia: all regions; Far East: Okhotsk, Zeya-Bureya, Uda River area, Ussuri, Sakhalin. Gen- eral distribution: Central Europe, Dzhungaria-Kashgaria, Mongolia, Japan, northern China. Described from Silesia. Type in Berlin. 109. C. incanum (S.G. Gmel.) Fisch. ex MB. FI. taur.-cauc. III (1819) 561; Fisch. Cat. Hort. Gorenk. 35, nom.; Ldb. FI. alt. IV, 9; DC. Prodr. VI, 643; Grossh. Fl. Kavk. IV, 178; Schischk. in Fl. Zap. Sib. XI, 2898; Klok. in Vizn. Rosl. URSR, 570; Charadze in Fl. Gruzil, VIII, 516; in Fl. Turkm. VII, 244; in Fl. Azerb. VIII, 413.—C. incanum var. pinnatilobum Sosn. ex Grossh. |. c. (1934).—C. arvense 6. incanum Ldb. Fl. Ross. II (1846) 753; Kryl. Fl. Alt. III, 697.—C. arvense var. incanum O. and B. Fedtsch. Perech. Rast. Turk. IV (1911) 287.—C. arvense 6. vestitum Wimm. and Grab.-Fl. Sil. II, 2 (1829) 92; Koch Syh. (1837) 400.—C. arvense ssp. vestitum Petrak in Schedis (1913); Arénes in Bull. Jard. Bruxelles, XXIV, 4, 272.—C. arvense var. com- mune f. incanum Beck. v. Monagg. Fl. Nied.-Oest. (1890) 1239.—C. arvense Vierh. in Oest. Bot. Zeitschr. LVI, 3 (1907) 108 p. p.—C. arvense ssp. incanum Iljin in Fl. Yugo-Vost. VI (1936) 408.—Serratula incana 8.G. Gmel. Reise, I (1770) 155.—S. ciliata MB. F1. taur.-cauc. II (1808) 267.—Ic.: S.G. Gmel. Reise, I, tab. 36; Rchb. Ic. Fl. Germ. XV (1852) tab. 842, III].—Exs.: Petrak, Cirsioth, Univ. Nos. 81, 82, 83. Perennial. Rhizome virgate, with horizontal branches, rooting and developing rosula of almost undivided broadly elliptical leaves. Stems ascending, sulcate, thinly white-tomentose above, cymosely branched, to 100 cm high. Lower cauline leaves ovate or broadly-elliptical, 7—8 cm long, 3-4 cm wide, undivided or shallowly sinuate pinnately parted, white-tomentose, beneath, green above, glabrous or thinly arachnoid- hairy, ciliolate-spiny, narrowed at both ends, sessile, with long- spinescent oblong or obtuse lobes, spinose at base of leaves; upper leaves strongly reduced. Capitula 1—4, at tips of stem and branches on long peduncles, in corymbose inflorescence, usually oblong, apically narrowed, with florets exserted from capitula, pappus hairs in pistillate capitula longer than florets. Involucral bracts greenish, with short, purple, usually deflected spine, weakly arachnoid-hairy below; outer and middle bracts ovate, inner ones linearly-lanceolate. Corolla lilac- ANS) 211 red, to 15 mm long; corolla tube long, narrow, in pistillate florets much longer than limb; limb up to base 5-parted. Pappus sordid-white; achenes 3 mm long. Flowering May to August. Forest-steppe and steppe zones, in mountains from lowlands to mid-montane zone, grows in thickets or singly along roads, on old fields, terraces in ravines of rivers and as weed in fields.—European Part: Crimea and southern regions of Ukraine; Caucasus: All regions; Soviet Central Asia: mountainous Turkmenia. General distribution: Central Europe (south), Balkans-Asia Minor, Armenia and Kurdistan, Iran. Described from eastern Ciscaucasia. Type in Leningrad. 110. C. ochrolepideum Juz. in Sorn. Rast. SSSR, IV (1935) 278; Charadze in Fl. Turkm. VII, 245.—C. arvense ssp. hookerianum Petrak in schedis.—Cephalonoplos ochrolepideum Nevsky in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 4 (1937) 290.—C. arvense var. horridum O. and B. Fedtsch. Perech. Rast. Turk. IV (1911) 287 p. p. Perennial. Rhizome virgate, with lateral rooting branches, to 1—1.5 m long; stem yellowish, to 120 cm high, branched above, more or less arachnoid-hairy below capitula. Leaves oblong-lanceolate or broadly lanceolate, coriaceous, 8 cm long, 1.5—2 cm wide, glabrous on both sides, glaucescent or more or less finely arachnoid-hairy beneath; unequally pinnately lobed, with crenate lobes, spinose, lobes and teeth tapered to spines up to 5 mm long. Capitula oblong, with florets 2.5 cm long, 13-15 mm in dia, with pale yellowish, glabrous, involucral bracts, sometimes (in staminate capitula) violet; outer bracts ovate or ovate- lanceolate, spinescent, inner narrow, linearly-lanceolate. Florets lilac; corolla to 15 cm long, corolla tube to 11 mm. Pappus hairs basally brown, to 20 mm long; achenes smooth or very weakly longitudinally sulcate, to 3.5 mm long. Flowering July to August. Grain fields, on irrigated lands, in cotton fields along irrigation channels.—Soviet Central Asia: mountainous Turkmenia, Pamiro-Alai, Syr-Darya, Tien Shan. General distribution: Dzhungaria-Kashgaria. Described from Kugitang. Type in Leningrad. 111. C. arvense (L.) Scop. Fl. Carn. ed. 2, II (1772) 126; Wimm. and Gr. Fl. Siles, II, 2, 92 p. p. (a. horridum and B. mite); DC. Prodr. VI, 643, excl. var. b. Koch, Syn. (1837) 400 (a. horridum and B. mite); Ldb. Fl. Ross. II, 734 p.-p.; Boiss. Fl. or. III, 552 p. p.; Vierh. in Oest. Bot. Zeitschr. LVII, 3, 107 p. p. (f. mite and f. horridum; Grossh. Fl. Kavk. IV, 177; Klok in Vizn. Rosl. URSR, 569 p. p.; Charadze in FI. Gruzii, VIII, 515; in Fl. Azerb. VIII, 412; Hegi, Ill. FI. VI, 2, 903; Majevski, Fl. ed. 8, 602 p. p.—C. arvense ssp. arvense Arénes in Bull. Jard. Bruxelles, XXIV, 4 (1954) 269.—C. arvense ssp. 214 Zi rubricaule Novopokr. in Engl. Bot. Jahrb. LXII (1929) 317.—C. rubricaule Novopokr. ibid. in adnot.—C. horridum Stank. in Stank. and Tal. Opred. Vyssh. Rast. (1949) 664, non MB.—Serratula arvensis L. Sp. pl. (1753) 820; Willd. Sp. pl. HI, 3, 1646.—S. spinosa Gilib. Fl. Lithuan. III (1781) 184.—Cnicus arvensis Hoffm. Deutschl. Fl. ed. 2, IL (1804) 130.—Carduus arvensis Robins. Brit. Fl. (1777) 163.— Breea arvensis Less. Syn. Comp. (1832) 9.—Iec.: Rehb. Ic. Fl. Gern. XV (1852) tab. 842, I; Vierh. in Oest. Bot. Zeitschr. LVII, 3 (1907) 109 (f. horridum and f. mite); Javorka and Csapody, Iconogr. Fl. Hung. (1933) 541, fig. 3873 (C. arvense f. horridum); Fiori, Fl. Ital. III (1933) No. 3712; Dostal. Kvét. CSR (1950) 1673, tab. 554; Hegi, Ill. Fl. VI, 2 (1929), Taf. 271, fig. 1, pag. 889, fig. 605 —Exs.: Pl. Ital. exs. No. 993 (var. kurridum Wimm. and Gr.); Petrak, Cirsioth. Univ. No. 59 (Var. horridum Wimm. and Gr.); Schultz. Herb. norm. n. s. No. 1853. Perennial. Rhizome virgate, with horizontal rooting suckers. Stems hard, ascending, 50-150 cm high, sulcate and weakly angular, often reddish, glabrous or sometimes with scattered hairs, with short, veg- etative, axillary branches below, branched above, below capitula often grayish-arachnoid-hairy. Leaves glabrous, glaucescent beneath or more or less finely arachnoid hairy, oblong or oblong-lanceolate; lower leaves 7-17 cm long, 1.5—4.5 cm wide, narrowed into more or less short petiole; upper leaves sessile, gradually reduced toward stem apex, nar- rowed at both ends, more or less deeply sinuately-pinnately parted with roundish obtuse lobes, with 2—S mm long spines; usually leaves soft, internodes lacking spines (f. mite) or more or less coriaceous and leaves at base of stem with long and thin spines, merging on intern- odes (f. horridum). Capitula ovate-oblong, 10-15 mm in dia, numer- ous on long, small-leaved peduncles, forming paniculate general inflo- rescence. Involucre imbricate, greenish or dark purple; outer involucral bracts thinly arachnoid-hairy on margin, with short deflected spine at apex; inner bracts linearly-lanceolate, acuminate, with scarious, purple tip. Florets lilac-pink; corolla to 15 mm long; corolla tube long, nar- row, limb strongly reduced in pistillate capitula. Pappus sordid-white, yellowish below, hairs 20-22 mm long; achenes 2.5—4.0 mm long, yellowish or reddish-brown. Flowering July to September. Forest and steppe regions, more rarely in mountainous areas; grows along forest edges, along roads, in old fields and crops as a weed.— European Part: in southern regions sporadically, in the north — Baltic Region; Caucasus: all region. General distribution: Scandinavia, Central Europe, Atlantic Europe, Mediterranean, Balkans-Asia Minor, North America (introduced). Described from western Europe. Type in London. Note. Great variability and the presence of intermediate forms often make a clear delimitation of the species of the C. arvense (L.) 215 218 Scop. alliance difficult. Out of the numerous forms and varieties re- ported for Western Europe, A.A. Grossheim cites the following vari- eties for the Caucasus: var. commune Beck., var. mite Koch, var. horridum Koch, var. setosum Koch, var. obtusilobum Beck. Evidently, following Vierhapper (op. cit.), it is necessary to con- sider the varieties mite and horridum closest to the typical form of C. arvense. Vierhapper recognizes four forms: f. mite, f. horridum, f. vestitum, and f. incanum. He considers the typical form of the species the most widespread in Europe as being intermediate between horridum and mite. He distinguishes f. vestitum Wimm. and Grab. from C. incanum. The synonymy given for the four forms, however, needs verification. We recognize C. incanum (S.G. Gmel.) Fisch. ex MB., and f. vestitum Wimm. and Grab. should be considered a synonym of it. Usually, all more or less pubescent forms were treated by authors of the Russian flora as a variety of C. incanum. C. incanum is the southern vicariant race of C. arvense (L.) Scop. It usually has leaves that are almost entire and grayish-arachnoid-hairy or white-tomentose beneath. The western European C. arvense, in turn, shows consider- able polymorphism, including the forms horridum, mite, and a thinly arachnoid-hairy one, but, unlike C. incanum, has more deeply pin- nately divided leaves. C. arvense is replaced in the east by C. setosum (Willd.) MB., which is widely distributed from Eastern Europe through Siberia and Eastern Asia to the Japanese Islands. This species with almost entire leaves is replaced in the east of its range by a pinnately- toothed form and in Dauria by a form with winged stems, described as C. argunense DC. and usually treated as var. subulatum Ldb. of C. setosum. The latter, cited by Bieberstein for the Caucasus, apparently should be excluded from its flora. C. incanum (S.G. Gmel.) Fisch. ex MB. usually grows in the Caucasus; forms of C. arvense are occasion- ally found: f. mite and a weakly arachnoid-hairy one called var. vestitum Wimm. and Grab. by European authors. C. horridum (Willd.) Stank., split off by S.S. Stankov from C. arvense, cannot be treated as sepa- rate, the more so because f. horridum is considered by some authors as the type form of C. arvense. The name C. horridum (Willd.) Stank. is a later homonym: cf. C. horridum (MB.) MB. and C. horridum (Adams) Petrak. Separation of C. rubricaule Novopokr., reported from the northern Caucasus, on the basis of the stem color is also unaccept- able. In the mountain regions of Soviet Central Asia, the pubescent forms of C. ochrolepideum Juz. are widespread; these usually were referred to C. incanum (S.G. Gmel.) Fisch. ex MB., but the glabrous forms (including the type of species) to var. horridum. True C. arvense f. horridum, more widespread in the Mediterranean region, is found only sporadically in the south of the European Part of the USSR and in the Baltic Regions. 214 GENUS 1593. Notobasis Cass.! * Cass, in Dict. Sc. Nat. XXV (1822) 225; XXXV (1825) 171. Capitula medium-sized, at stem tips and in axils of upper leaves, surrounded by apical leaves modified into pinnately incised spines, subsessile at stem tip, in clusters of few, below on short peduncles solitary, unisexual and many-flowered. Involucre broadly ovate or ovate- globose, 1.5—2(2.5) cm in dia, weakly arachnoid-hairy; involucral bracts many-rowed, imbricate, coriaceous, glabrous on outside, with numer- ous pale, punctate, sessile glands or linearly-subulate outer to lan- ceolate, longer inner, terminated at tip in straight or somewhat de- flected spinule, innermost bracts long-acuminate, on both sides pubescent in upper part. Receptacle with narrow, linear, long scales. All florets tubular, purple, somewhat surpassing involucre, with 5- parted glabrous corolla; peripheral florets outwardly resembling bi- sexual, fertile, middle florets but with undeveloped ovary and empty (lacking pollen) anthers, sterile, because of which capitula considered heterogamous. Anthers with short, fimbriate basal appendages, apical appendages acute; stamen filaments flattened, covered with papilli- 216form hairs. Style fused to tip, very shortly-lanate, entirely exserted from staminal tube, branched or unbrached, bifid with collar of hairs at base of branches. All achenes alike, obliquely obovate, 5-6 mm long, rounded above, lacking corona, with flat, oval, upper scar of attachment, smooth, entirely glabrous; hilum slightly oblique, slit-like, pappus 3 times as long as achenes, whitish, yellowish below, simple, fragile, falling entirely, plumose, soft bristles, more or less of same length, connate at base in ring. Annual herbs with erect stem and spiny, sinuate-lobate, amplexicaul leaves. A monotypic genus. 1. N. syriaca (L.) Cass. in Dict. Sc. nat. XXXV (1825) 171; DC. Prodr. VI, 660; Ldb. Fl. Ross. II, 2, 746; Boiss. FI. or. III, 553; Grossh. Fl. Kavk. IV, 177 and Opred. Rast. Kavk. 481; Gadjiev in Fl. Azerb. VIII, 389.—Carduus syriacus L. Sp. pl. (1753) 823.—Cirsium syriacum (L.) Gaertn. De fruct. II (1791) 383.—C. bracteatum Link in Linnaea, IX (1834) 380.—Cnicus syriacus (L.) Willd. Sp. pl. I, 3 (1803) 1683 p. p. excl. syn. Bauh.—Ic.: Gaertn. 1. c. tab. 163, fig. 2; Schur. Bot. Handb. ed. 2 (1808) tab. 229; Fiori and Paol. Iconogr. fl. Ital. 2 (1899- 1904) 452.—Exs.: Petrak, Cirsioth. Univ. fasc. IV, No. 34; Fl. Terr. Israel. exs. No. 581; Fl. Palaest. exs. No. 197. 'Treatment by S.K. Czerepanov. *From the Greek words notos—back, and basis—base; named for oblique achenes. 218 ZU5 Annual. Stem (20) 30-70 cm (probably even more) high, erect, sulcate, usually simple, rough, entirely sparsely arachnoid-hairy. Leaves oblong, sinuate-lobate, with deltoid, strongly spiny lobes, glabrous above, very weakly arachnoid-hairy beneath, lower leaves short-peti- olate, others sessile, amplexicaul. Capitula at tips of stem and in axils of upper leaves, surrounded by apical leaves modified into pinnately incised spines and surpassing capitula, at stem tip subsessile, clus- tered, on short peduncles and solitary below. Involucre broadly-ovate or ovate-globose, 1.5—2.2 cm long and 1.5—2(2.5) cm in dia, weakly arachnoid-hairy; involucral bracts smooth, with numerous, pale, punc- tate, sessile glands, from linearly-subulate outer to oblong-lanceolate, longer inner, terminating in straight or somewhat deflected short spine; innermost bracts long-acuminate, on both sides pubescent in upper part. Florets purple. Achenes pale reddish-brown, obliquely obovate, 5—6 mm long, 4—4.5 mm wide, glabrous; pappus 15-17 mm long. Flowering May to July. Dry slopes, weedy places, near roads, old fields, edges of crop fields, up to lower mountain zone.—Caucasus: Eastern Transcaucasia, Talysh. General distribution: Mediterranean, Balkans-Asia Minor, Armenia and Kurdistan (former Karsk Region). Described from Syria. Type in London. GENUS 1594. Picnomon Adans!: 2 Adans. Fam. II (1763) 116; Cass. in Dict. Sc. nat. XL (1826) 187; DC. Prodr. VI (1837) 634.—Cirsium sect. Acarna DC. in Duby, Bot. Gall. I (1828) 286 p. min. p. Capitula homogamous, homochromous, medium-sized, more or less oblong. Involucre ovate-conical; outermost involucral bracts foliaceous, considerably longer than florets, imbricate, appressed, with cuneately narrowed, plumose, echinate, recurved appendages, innermost squa- mose, with weakly toothed appendage, spinulose. Receptacle densely long-squamose. Florets all fertile, bisexual; corolla pink-purple; sta- men filaments glabrous or glandular. Style branches free above, diver- gent. Achenes obcuneate, weakly angular, lustrous, slightly flattened, with very narrow almost slit-like lower hilum above, with distinct 5- lobed, stalked tubercle with roundish lobes; pappus as long as corolla, consisting of sericeous, lustrous, finely plumose hairs, somewhat thick- ened above, many-rowed, connate at base in ring and falling off 'Treatment by S.G. Tamamschian. "From the Greek word picno—dense, densely flowered. 216 etc al fy iy Ning 217 Plate X. 1—Lamyra echinocephala (Willd.) Tamamsch.; 2—Ancathia igniaria (Spreng.) DC.; 3—Modestia mira (Iljin) Charadze and Tamamsch. 219 217 entirely. Annual-biennial plant with erect, weakly branched stem, and narrow spinous-toothed, alternate, decurrent leaves. A monotypic genus. 1. P. acarna (L.) Cass. in Dict. Sc. nat. XL (1826) 188; Grossh. Fl. Kavk. IV, 190; O. and B. Fedtsch. Perech. Rast. Turk. IV, 282.— Carduus acarna L. Sp. pl. (1753) 820.—Cnicus acarna L. Sp. pl. ed. 2 (1763) 1158.—Carthamus canescens Lam. FI. Fr. II (1778) 10.— Carlina acarna MB. FI. taur.-cauc. II (1808) 282.—Cirsium acarna Moench, Meth. Suppl. (1802) 226; Boiss. FI. or. III, 549; Schmalh. FI. II, 104. Annual or biennial. Plants 15—40 cm high, appressed-thinly arach- noid-hairy or grayish, sometimes whitish-pubescent. Stem white, sul- cate, almost entirely covered with narrow, subentire, oblong, spiny- ciliate wings, densely leafy, usually branched above. Lower leaves deciduous, middle and upper ones to 10-11 cm long, 13-15 mm wide, lanceolate, ciliolate, weakly pinnate-sinuate, with long, deflected, yellowish spines, 8—9 cm long, coriaceous, decurrent, with prominent reticulate veins beneath, and thin, white, glabrous midrib above, on both sides uniformly thinly whitish- or grayish-pubescent. Capitula clustered at stem tips in corymbose-paniculate inflorescence, narrowly- oblong, sessile, solitary or in clusters of 2—4; apical leaves with long spines. Involucral bracts appressed, densely pubescent. Corolla pink- ish or purple, raised above pappus hairs, 12-13 cm long; corolla tube considerably longer than limb. Achenes 4-5 mm long, about 2 mm wide in upper part; pappus white or grayish 3—4 times as long as achenes, with narrow squamiform bristles and sericeous, thin hairs. Flowering July to August. Dry, abandoned places, near roads.—European Part: Crimea; Caucasus: all regions; Soviet Central Asia: mountainous Turkmenia, Syr-Darya. General distribution: southern Europe, eastern Mediterra- nean, Armenia and Kurdistan, Iran and Afghanistan. Described from Europe. Type in London. GENUS 1595. Lamyra Cass." * Cass. in Bull. Soc. Philom. Paris (1812) 168; in Dict. Sc. nat. XXV (1822) 218, XLI (1826) 305.—Platyraphium Cass. in Dict. Sc. 'Treatment by S.G. Tamamschian. *From the Greek word /amyros—prickly-pointed. 220 218 nat. XXXV (1825) 173, XLI (1826) 305.—Chamaepeuce Dc. Prodr. VI (1837) 657 p. p. Capitula many-flowered, tubuliforms, homogamous and homochromous. Involucre ovate; bracts imbricate, more or less herba- ceous or stiff, with basal callose thickening, always cuspidate, ap- pressed, or outer deflected. Receptacle somewhat fleshy, bulged, with numerous, free, unequal bristles. Corolla almost symmetrically lobed above, open; filaments hairy, basal anther appendages long, subulate, membranous, lobed. Style branches very short. Achenes initially with thick pericarp, becoming hard and woody with progressive maturation, ovate-hemispherical or more or less pyriform, glabrous, lacking ribs, and with or without inconspicuous veins, reddish-brown or black, yellowish or gray, often with marbled pattern; hilum broad, roundish or somewhat trapezoid, sometimes slightly truncate, upper scar flat or with barely noticeable border, horizontal or slightly truncate; pappus many-rowed, of white, almost equal, thin, fine barbate bristles, thick- ened above, often plumose, basally connate, falling entirely. Biennial or perennial herbs with alternate, entire, pinnately parted or lobate, usually prickly leaves, more or less densely tomentose beneath, gla- brous above, coriaceous, sometimes lustrous and stiff; stem glabrous or pubescent, simple or branched. Type of genus: Lamyra casabonae (L.) Tamamsch. The genus includes about 10 highly ornamental species, distrib- uted in Southern Europe and Asia Minor. Of these, one species is found in the USSR. 1. L. echinocephala (Willd.) Tamamsch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVI (1954) 470.—L. angustifolia Cass. in Dict. Sc. nat. XXV (1822) 221.—Cnicus echinocephalus Willd. Sp. pl. II, 3 (1803) 1685; MB. Fl. taur.-cauc. II, 280.—Serratula echinocephala Poir. in Lam. Encycl. meth. VI (1804) 561.—Cirsium echinocephalum MB. F1. taur.-cauc. III (1819) 559; Spreng. Syst. veg. 3, 374; Boiss. Fl. or. HI, 551.—C. wachtangii Woron. in Byull. Kavk. Muz. XI (1917) 20.—Chamaepeuce echinocephala DC. Prodr. VI (1837) 660; Ldb. Fl. Ross. II, 746; Grossh. Fl. Kavk. IV, 190.—C. wachtangii Grossh. Fl. Kavk. IV (1934) 190.—Exs.: Petrak, Cirsioth. Univ. fasc. X, No. 97; GRF No. 1369 (sub Chamaepeuce echinocephala DC.); Herb. Fl. cauc. No. 599. Perennial. Stem erect, 15-50 cm high, simple or cymosely branched above, cylindrical, weakly sulcate, tomentose, densely leafy, occasion- ally with single capitulum. Leaves glabrous, coriaceous, weakly rug- ose, and dark- or light-green above, snow-white-tomentose beneath, lanceolate, pinnately incised to narrowly-linear or deltoid lobes with 219 long tip and revolute margins, in upper part of stem more or less semiamplexicaul, midrib above, in lower part, thickened and densely tomentose, throughout glabrous beneath; terminal leaf lobe usually broaden than laterals and 2—3 or 5—6 times as long, ensiform or lan- ceolate, occasionally subulate. Capitula terminal, 4—5 cm long, broadly- cylindrical at flowering, on short peduncle. Involucral bracts glabrous, stiff in mature capitula, appressed, narrowed above and terminating in thin spine almost as long as bract; outermost bracts deflected, middle ones straight, appressed, inner ones purple, scarious. Florets dark pink or purple, to 3—3.5 cm long; stamen and style strongly exserted from corolla. Achenes large, to 5 mm long, thickened above, 3 mm wide, roundish in cross section, narrowed below, truncate, obliquely pyri- form, often with black marbled pattern on dark gray surface, upper scar of attachment slightly oblique, so that corolla and pappus some- what lateral; pappus much shorter than corolla, but longer than tube. Flowering July to August (September). (Plate X, Fig. 1.) Stony screes and steep slopes, on rocks.—European Part: Crimea; Caucasus: Ciscaucasia, Western Transcaucasia. Described from Crimea. Type unknown; isotype in Leningrad. Note. A highly ornamental plant, strongly varying in the color of the achenes and the form and size of the leaf lobes, especially the terminal lobe. The broader and considerably longer terminal lobe com- pared to the lateral lobes served as the basis for Yu.N. Woronov to treat the specimens from Georgia as a separate species. The species of Chamaepeuce reported for Soviet Central Asia, in fact, belong to the genus Cirsium (cf. Tamamschian in Zam. po Sist. i Geogr. Rast. Tbil., 22 (1961)). GENUS 1596. Lamyropappus Knorr. and Tamamsch.!? Knorr. and Tamamsch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVI (1954) 463. Capitula homogamous, homochromous, large, solitary at tips of stems. Involucre globose-ovate; involucral bracts numerous, imbricate, appressed, stiff, with stiff, straight or recurved tip. Receptacle shal- low-alveolate, with numerous long bristles on edges of alveoli. Florets numerous, slightly zygomorphic, bisexual. Lower appendages of an- thers long, caudate, subentire, weakly parted only at tip to 2-3 bristles; 'Treatment by S.G. Tamamschian. From the similarity of the pappus to that of the genus Lamyra. 222 220 filaments glabrous, almost smooth. Style exserted from corolla, style branches almost inconspicuous. Achenes 9-11 mm long, 3 mm wide, weakly deltoid, more or less flattened, weakly longitudinally sulcate, slightly narrowed at ends, slightly truncate; hilum straight or slightly oblique; upper scar patellate, horizontal, with scarcely noticeable rem- nants of corolla and style in middle (tubercle); pappus rowed, with light green, yellowish, plumose bristles racemosely thickened above, later pointed, scarcely exserted from involucral bracts. Perennial herbs with entire, coriaceous, large leaves. A monotypic genus as yet known only from Soviet Central Asia. 1. L. schakaptaricus (B. Fedtsch.) Knorr. and Tamamsch. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVI (1954) 466.—Cirsium schakaptaricum B. Fedtsch. in O. and B. Fedtsch. Perech. Rast. Turk. IV (1911) 287.—Ic.: Knorr. and Tamamsch. op. cit. 465. Perennial. Rhizome thick, ligneous, branched, Stems basally white- lanate, covered with tomentose remnants of previous years leaves, numerous, to 75 cm high, weakly branched, cylindrical, sulcate, floc- cose-tomentose, with solitary capitula. Leaves numerous, dense below, sessile, in upper part distant, highly variable in size and form, lower ones large, broadly-obovate, 14-15 cm long, 9.0—9.5 cm wide sessile; petiole flat, ampliate below, 5-6 cm long, 1.0—1.5 cm wide, in lower part weakly grooved, basally crenate, with occasional thin spines; up- per leaves oblong-obovate, ovate, elliptical, or lanceolate, 1.5—8.0 cm long, 0.6—3.5 cm wide; apical leaves reduced to spinules; all cauline leaves sessile, sometimes weakly decurrent, coriaceous, glabrous above, light green veins with plumose hairs, lateral veins terminating at mar- gin to hard spines, densely white-tomentose-lanate beneath, with promi- nent midrib. Capitula very large, 5.0-5.5 cm long, 2.5—3.0 cm in dia. In volucre of numerous, dense, appressed, imbricate bracts; outer bracts oblong-obovate or spatulate above, with recurved tips, middle and inner ones linear with straight tip. Florets large, 3.0—3.2 cm long; corolla light violet. Pappus considerably shorter than corolla, 2.5 cm long; achenes light gray, light reddish-brown, or with dark reddish- brown spots or bands on lighter background or all straight or slightly truncate, usually 1/2 as long as pappus. Flowering July to September. River valleys on chalk deposits, on gypseous strata and red clays.— Soviet Central Asia: Lake Balkhash Region (southwestern part), Tien Shan (southern part), Syr-Darya (eastern part). Endemic. Described from Shakaptar. Type in Leningrad. 223 224 GENUS 1597. Ancathia DC.! 2 DC. in Guill. Archiv. bot. II (1833) 331; Prodr. VI, (1837) 557. Capitula homogamous, homochromous. Involucre ovate globose; involucral bracts imbricate, outer and middle bracts recurved, inner ones appressed, almost entirely scarious, dry, purple or flame-red, very much like inner bracts of genus Carlina, lanceolate, acuminate. All florets similar, bi-sexual, pink or purple; corolla obliquely 5-lobed, corolla tube almost as long as limb. Filaments glabrous; basal append- ages of anthers on both sides with bushy-tailed. Achenes glabrous, oblong, more or less narrowed below, ribbed- angular, longitudinally sulcate on sides, dark reddish-brown or almost black; hilum oblique, upper scar angular with thin toothed edge; pap- pus usually two-rowed, falling off entirely, hairs connate below, al- most equal, thinly plumose. Perennial herbs with short, usually simple, sometimes branched, solitary or many stems with peduncles bearing single capitula; leaves linear or linearly-lanceolate, alternate, almost entire but with sharp spines on revolute margin, glabrous above, like stem densely grayish- or whitish-tomentose beneath. A monotypic genus. 1. A. igniaria (Spreng.) DC. in Guill. Archiv. bot. IT (1833) 331; Prodr. VI, 557; Ldb. Fl. Ross. II, 681; O. and B. Fedtsch. Perech. Rast. Turk. IV, 265; Grossh. Fl. Kavk. IV, 190.—Cirsium igniarium Spreng. Syst. Veg. 3 (1826) 375; Ldb. Fl. alt. IV, 10; Kryl. Fl. Zap. Sib. XI, 2900.—Cnicus igniarius Benth. in Benth. and Hook. f. Gen. pl. II (1873) 469.—Ie.: Ldb. Ic. pl. Fl. Ross. V, tab. 477; DC. in Deless. Ic. Sel. IV, tab. 73. Perennial. Rhizome vertical, mostly branched, to 1.5 cm thick. Stems straight, solitary or several, sometimes spreading, simple or with few branches, whitish-tomentose, un-winged. Leaves linear or narrowly- lanceolate, 5-20 cm long, 3—15 cm wide, lower leaves withering be- fore flowering, middle and upper ones more or less decurrent, weakly amplexicaul, entire, with strongly revolute lamina, with 2, 3 or large number of hard, acute, yellowish spines on margin above, lustrous, sometimes weakly arachnoid-hairy, mostly glabrous, white-tomentose beneath; uppermost cauline leaves reduced, usually subtending capitula. Capitula solitary terminal on peduncles, large, 2.5—4.0 cm in dia. In- volucre arachnoid-floccose; bracts imbricate, many-rowed; all bracts stiff, lanceolate, narrow, with long, hard spine, outermost bracts re- curved, middle ones straight, inner ones thin, membranous, orange or 'Treatment by S.G. Tamamschian. *From the Greek word ankos, meaning spiny, sticky [or clinging]. 200 reddish, whitish above. Corolla flame-red or lilac; limb lobed to 1/3 or slightly more. Achenes quite large, black, to 6 mm long. Flowering July to September. (Plate X, Fig. 2.) Stony steppes, small knolls, atop clay domes (hillocks), on solo- netzes, on rubble slopes, screes of limestone rocks, rarely in alpine meadows or in fescue and shrubby steppes.—Western Siberia: Irtysh (south), Altai; Eastern Siberia: Angara-Sayans (south); Caucasus: Dagestan; Soviet Central Asia: Lake Balkhash Region, Dzhungaria- Tarbagatai, Kzyl-Kum (northwestern). General distribution: Dzhungaria-Kashgaria (western), Mongolia (northwestern and western). Described from Altai. Type in Leningrad. GENUS 1598. Modestia Charadze and Tamamsch.!:? Charadze and Tamamsch. in Zam. po Sist. i Geogr. Rast. Tbil. 19 (1956) 40. —Cirsium sect. Anacantha Iljin in Bot. Mat. Gerb. Glavn. Bot. Sada, III (1922) 57. Capitula homogamous, homocromous, erect, in clusters of 3—4, in corymbs, occasionally solitary. Involucre more or less ovate, at fruit- ing obconical; involucral bracts many-rowed, imbricate, attenuate into cusp with more or less long spine, outer bracts shorter, ovate-lan- ceolate or lanceolate, middle ones elongately elliptical, innermost lin- early-lanceolate. Receptacle alveolate, covered with unequal sparse, subulate, scarious bristles. Florets bisexual, slightly zygomorphic, 3.0— 3.5 cm long; filaments glabrous or with thin papillae only in upper part, basal anther appendages caudate, bifid, glabrous. Achenes straight or more or less curved below, to 9 mm long, tetraquetrous, on sides weakly sulcate-rugose, with prominent ribs above and thin rectangular, toothed ridge, often with remnants, of corolla and style in center, below also with more or less quadrate hilum, end of achene visible in its centre; pappus many-rowed, of white, sericeous, short-plumose, nu- merous (outer shorter) bristles, connate at base in ring, hence falling together. Perennial, densely tomentose, with ligneous, basally winged stems and alternate, un-divided, decurrent leaves. Type of genus: M. darwasica (Winkl.) Charadze and Tamamsch. An oligotypic genus of 2-3 species. 1. Capitula usually clustered at stem tip, on peduncles, 2—6, very rarely solitary. Involucre more or less hemispherical, glabrescent, at fruiting sub-glabrous; involucral bracts, both middle and outer, with 'Treatment by S.G. Tamamschian. 7In honor of Soviet Botanist Modest Mikhailovich Iljin. 225 223 hard, spiny tip, longer than bracts, spiny tip sometimes weakly developed .......... 1. M. darwasica (Winkl.) Chardze and Tamamsch. + Capitula solitary, at stem tip, on long peduncles. Involucre nar- rower, at fruiting obconical, whitish-pubescent from beginning to maturity; involucral bracts with inconspicuous tip; only inner bracts of unopened capitula terminating in more or less. hard tip. .................. Mons «elle 2. M. mira (Ijin) Charadze and Tamamsch. 1. M. darwasica (Winkl.) Charadze and Tamamsch. in Zam. po Sist. i Geogr. Rast. Tbil. 19 (1956) 41.—M. jucunda Charadze and Tamamsch. 1. c.—Cnicus darwasicus Winkl. in Tr. Peterb. Bot. Sada, IX, 2 (1886) 427.—C. jucundus Winkl. 1. c.—Saussurea darwasica Lipsky in Tr. Peterb. Bot. Sada, XXVI, 2 (1910) 437 p. p.; O. and B. Fedtsch. Perech. Rast. Turk. IV, 234.—S. jucunda O. and B. Fedtsch. 1. c.—Cirsium darwasicum Ijin in Bot. Mat. Gerb. Glavn. Bot. Sada, III (1922) 59.—C. jucundum Ijin, ibid. 58. Perennial. Plants snow-white or greenish, densely tomentose. Stems numerous, branched, occasionally simple, straight, with 2—6 capitula. Leaves undivided, oblong-elliptical, occasionally lanceolate, lower ones usually larger, 10—12 cm long (excluding decurrent part), 2.0—2.5 cm wide, strongly decurrent such that stem broad-winged over internode, cuspidate greenish and weakly pubescent or grayish above, more or less densely grayish-pubescent beneath, like stem densely white- or grayish-tomentose. Capitula large, at fruiting almost 4 cm long, about 2 cm in dia. Involucre at fruiting more or less campanulate; outer involucral bracts small, 1-2 mm long, middle ones longer, both, long _ with deflected, long spine, inner bracts much longer, 2.0—2.8 cm long, whitish below, colored above, with thick midrib and 2 thin lateral veins, continued into tip but not spiny, ciliolate in elongated part, often darker at tip, remaining part stramineous. Corolla pink, scarcely exserted from pappus. Pappus as long as inner involucral bracts, be- fore falling off far exserted from involucre, with achenes 3.5—3.8 cm long; achenes almost white, pale or reddish-brown, 7—9 mm long. Fruiting August to September. Mountain and high-mountain slopes along river valleys, on gypsif- erous limestone slopes, sometimes on sands and gravelly sediments.— Soviet Central Asia: Pamiro-Alai (southern and southeastern. parts). Endemic. Described from Tadzhikistan. Type in Leningrad. 2. M. mira (Iljin) Charadze and Tamamsch. in Zam. po Sist. i Geogr. Rast. Tbil. 19 (1956) 41.—Saussurea darwasica Lipsky in Tr. Peterb. Bot. Sada, XXVI, 2 (1910) 437 p. p.—Cirsium mirum [jin in Bot. Mat. Gerb. Glavn. Bot. Sada, III (1922) 61. 226 224 Perennial. Rhizome vertical, woody. Stem to 50-55 cm high, stocky, branched almost from base, sulcate, densely white-tomentose. Leaves lanceolate, 2-7 cm long (excluding decurrent part), 0.5—1.0 cm wide, entire, light green and weakly pubescent above, except white midrib, from pubescence, densely tomentose beneath with whitish or light gray midrib, strongly decurrent, such that internodes throughout narrowly- winged, wings considerably narrower than in preceding species, only in beginning 2—3 mm wide, becoming narrower downward, at end less than 1 mm wide. Capitula solitary, on peduncles, terminal distinctly smaller than in preceding species, 1.0—1.5 cm long, 0.5—0.8 cm in dia. Involucre in open capitula obconical, slightly rounded below, sur- rounded by small apical leaves; all bracts with distinct, deltoid, dark spot above; outer and middle bracts firmly appressed, whitish-tomen- tose; inner ones weakly pubescent, lighter in color, as long as pappus. Mature achenes unknown, immature reddish-brown. Flowering July. (Plate). X; Fig. 3.) Mountain slopes along river banks.—Soviet Central Asia: Pamiro- Alai (southern part). Described from Karategin. Type in Leningrad. GENUS 1599. Cynara L.'? Le Sp. YpliGl753) (827: Capitula large, on peduncles, homogamous, homochromous, with many setae. Involucre globose; involucral bracts imbricate, many-rowed, appressed at base, almost straight, coriaceous, narrowed into hard, conical spine, occasionally without spine, revolute. Receptacle flat, but fleshy, with long bristles. Corolla regular, 5-lobed; limb with un- equal lobes, 1/2 as long as broad tube. Filaments glandular; basal appendages of anther short, more or less barbate, apical ones obtuse. Style branches convergent. Achenes obovate, thickish, somewhat bulged and weakly 4-angled, hard, glabrous and smooth, with almost straight or somewhat truncate hilum, upper scar lacking annulus; pappus of many.-rows of numerous, plumose, almost equal bristles, somewhat scariously broadened below, connate at base in ring and falling with it. Large perennial, prickly herbs, with basal rosette of leaves. Type of genus C. scolymus L. Of the 10-11 species in the Mediterranean Region and Canary Islands one is found in cultivation in the USSR. 1. C. scolymus L. Sp. pl. (1753) 827. 'Treatment by S.G. Tamamschian. "From the Greek word meaning dog. 227 225 Perennial. Plant 0.5—2 m high. Stem thick, straight, densely gray- ish- or whitish-arachnoid-hairy, weakly sulcate. Leaves larger, forming broad rosette at stem base, arachnoid-hairy beneath; lower leaves oblong or broadly lanceolate, about 1 m long and 50 cm wide, decurrent, twice pinnately incised, primary segments decurrent on winged axis and divided into irregularly lobate or toothed lobes, lobes and teeth unarmed, glabrous and green above, densely tomentose beneath; middle and upper cauline leaves reduced, sessile, weakly decurrent, upward reduced to linear and oblong apical leaves, 5—6 cm long. Capitula large, 7-10 cm long, solitary, terminal on short branches. Involucre ovate or subglobose; involucral bracts subglabrous, smooth, stiff, co- riaceous, with more or less fleshy base, outer bracts slightly de- flected, erect, or recurved, middle and innermost bracts straight, im- bricate, appressed; middle bracts more or less narrowed above, acute, lacking spine, inner ones with short, roundish or obtuse appendage. Florets numerous; corolla bluish-purple. Achenes 6—7 mm long, ob- long, reddish-brown with inconspicuous white and darker, sometimes black spots; pappus considerably longer than achenes, 2—3 cm long, more or less yellowish. Flowering June to October. Cultivated in the south on fertile soils. Note. It has been suggested that C. scolymus L. is a cultivar of the wild Mediterranean species C. cardunculus L., which is not found in the USSR in the wild. Economic Importance. It is a culinary delicacy. Its fleshy recep- tacle and the fleshy base of the involucral bracts are eaten boiled. It also is grown as an ornamental plant for its attractive leaves and large red capitula. GENUS 1600. Silybum Adans’” Adans. Fam. II (1763) 116. Capitula terminal, solitary, large, nodding, homogamous. Involu- cre broadly globose; involucral bracts imbricate, outer and middle bracts with foliaceous, prickly-toothed appendage, terminating in long, erect, lanceolate-subulate straight spine; inner bracts undivided, with or with- out small appendage. Receptacle flat, fleshy, scarious-ciliate. All flo- rets alike, fertile, bisexual; corolla violet, purple, pink, occasionally white; corolla tube narrowly, cylindrical, long, limb campanulate, short, broadened; corolla lobes narrow, linear, undivided, with lateral veins 'Treatment by S.G. Tamamschian. *From a Greek name applied to a prickly plant used as food. 228 226 converging above. Filaments free above, connate above base because of dense short mucilaginous hairs covering them; basal anther append- ages linear-cilia-like, as long as connective; apical appendages sagittately deltoid. Style thickened above, somewhat truncate, covered with long, branched hairs; style branches approximate, slightly diver- gent above, like style covered with short, branched hairs on outside. Achenes obovate, slightly truncate on sides, glabrous, with indistinct ribs, hilum straight, tubercle (nectary) on upper scar raised; pappus of many scaly, toothed bristles, connate at base in ring with crown of tiny, lustrous, approximate hairs. Annual or PE herbs, with hard, leafy stem and spiny alternate leaves. A monotypic genus, widely distributed throughout the Mediterra- nean Region, in central and southern Europe, and in Central Asia. 1. S. marianum (L.) Gaertn. De fruct. II (1791) 378; Grossh. FI. Kavk. IV, 191.—S. maculatum (Scop.) Moench, Meth. (1794) 555.— S. martiae (Cranz) S.F. Gray, Nat. Arr. Brit. Pl. TT (1821) 436— Carduus marianus L. Sp. pl. (1753) 823.—C. mariae Crantz. Inst. I (1766) 248.—Mariana mariana (L.) Hill. Hort. Kew. (1768) 61.—M. lactea Hill. Herb. Brit. I (1769) 76.—Cirsium maculatum Scop. FI. Carn. ed. 2, II (1772) 130.—Carthamus maculatus (Scop.) Lam. Encycl. meth. I (1783) 638.—Ic.: Kozlovskaya in Fl. BSSR, V (1959) Plate LXIX. Annual or biennial. Stem usually to 1.5 m high, sometimes quite low (30 cm), very rarely 10 cm, or very tall, taller specimens to 2—3 m, glabrous or weakly arachnoid-hairy, sulcate, straight, branched, occasionally simple, also farinose, cylindrical. Leaves green, lustrous, with large white spots, sometimes to 80 cm long and 30 cm wide; lower leaves rosulate, together with middle and upper ones oblan- ceolate or elliptical, spiny-toothed pinnately lobed or pinnately in- cised, lobes prickly or prickly-toothed, petiolate; upper leaves reduced, sessile, with broad, amplexicaul and short-decurrent base, deeply or shallowly pinnatifid, prickly-toothed, uppermost still reduced, with large spiny amplexicaul lobes and acute tip. Capitula medium- or large- sized, 3-6 cm long, oblong, often globose, solitary on slender pe- duncles. Involucral bracts weakly pubescent or glabrous, imbricate; outer and middle bracts appressed at base, with erect foliaceous, hard, spinescent appendage and 4—6 spines on margin, appendages as long as or longer than florets; innermost bracts straight, coriaceous, with small appendage or narrowed into nonspiny tip. Florets numerous, pink, violet, or white. Achenes elliptical or obovate, to 7.3 mm long, weakly transversely rugose, dark reddish-brown or black with longitudinal stria- tions or more or less white-spotted, tubercle on somewhat angular 22 \o 2277 stalk, cylindrical, 3-lobed pappus hairs unequal, white or yellowish, outer hairs thin, setaceous, inner ones somewhat thickened, scarious, 1-2 cm long, 2-3 times as long as achenes, ring with thin, very short, delicate hairs, 2 mm long. Flowering July to August. Uncultivated lands, roadsides, abandoned fields.—European Part: Ladoga-Ilmen, Baltic Region, Volga-Kama, Upper Dnieper, Middle Dnieper, Volga-Don, Trans-Volga, Upper Dniester, Black Sea Region, Bessarabia, Crimea; Caucasus: Ciscaucasia, Dagestan, Eastern and Southern Transcaucasia, Talysh; Western Siberia: Upper Tobol (south- western); Soviet Central Asia: Kara-Kum, Syr-Darya, Pamiro-Alai. General distribution: central and Atlantic Europe, Mediterranean, northern Africa. Described from western Europe. Type in London. Note. It is highly variable in habit, color of corolla, and length of spines on the involucral bracts. In view of this, the described varieties and the species S. leucanthum Jord. and Fourr. can be considered only as forms of this species. GENUS 1601. Onopordum L.'” L. Sp. pl. (1753) 827.—Acanos Adans. Fam. II (1763) 116. Capitula homogamous, homochromous, many-flowered, and uni- sexual, solitary or numerous, terminal, occasionally axillary. Involucre ovate, oblong-globose or globose; involucral bracts imbricate, lanceolate or cuneate, apically without or with long or short spines, appressed, divergent or strongly deflected, sometimes more or less uncinate, nar- row or broad, coriaceous or stiff, more or less thick, hollow or flat, subglabrous, pubescent or glandular-hairy. Receptacle fleshy, deeply alveolate, alveoli membranous along margin and sinuately irregularly toothed. Corolla 5-fid, slightly open, saccate, purple, violet, pink, yel- lowish, or white. Filaments glabrous or weakly glandular, lacking simple hairs; apical appendages of anthers linearly-cuneate or subulate, basal appendages shortly caudate, undivided, cuneate, truncate or straight and narrower, sometimes weakly lobed. Style long; style branches 1/5—1/3 as long as style, usually folded and entirely or partly exserted from corolla. Achenes oblong or obovate, sometimes more or less compressed, indistinctly or distinctly 3—4-quetrous, with distinct or indistinct ribs, on sides with oblique or parallel transverse-wrinkles or tubercles, with many or few, longitudinal, thin veins and furrows, 'Treatment by S.G. Tamamschian. "From the Greek words onos—ass, and pordon—flatulence. 230 228 4—7 mm long, dark brown or light- and dark-gray with darker, some- times black spots, marbled; upper scar of attachment with thin edge, usually angular, with distinct or inconspicuous tubercle, hilum straight or slightly oblique, entirely or partly hidden under thick callose ap- pendage; pappus fragile, falling off entirely, consisting of 2—3, occa- sionally many rows of equal bristles, of which, in many species, one row with longer and thicker bristles than others. All florets ciliolate, smooth, barbate or plumose above, connate in broader, angular, thick, waxy ring. Biennial, occasionally perennial, herbs, stemless or with erect, tall or short, solitary stem, simple or branched from middle or weakly branched only above, spiny-winged, with large, spinose, green or pubescent leaves. Type of genus O. acanthium L. A widely distributed Mediterranean genus, comprising about 50 species; in the USSR, it is represented by 10 quite polymorphic species. Economic Importance. Good honey-producing plants. Their seeds contain oil which is similar in chemical composition to sunflower oil (Lazurevsky, 1934). In horticulture, the West European species O. bracteatum and O. acanthium are grown as ornamental plants. 1. Plants not very spiny; capitula 3 cm long, 1.0—1.5 cm in dia, un- opened ovoid, opened oblong, axillary and terminal; involucral bracts small allappresseds. 21228 2a 6. O. frickii Tamamsch. + Plants highly spiny; capitula 5-7 cm in dia, globose, somewhat depressed or oblong, often solitary at apices of branches, occasion- ally in clusters of 2—3; involucral bracts erect, deflected or ap- PRESSE a. PAR Bir ed WR ah OA Oe OO eae 2. 2. Involucral bracts coriaceous, broad, green, glabrous but with nu- merous small glands on midrib, inward and on margin........ EER SUMS MERE, VARI OBIS Star a0, 4. O. tauricum Wild. + Involucral bracts narrower, more or less strongly pubescent, eglandular or with few intermixed glands only on midnib................ Bt 3. Pappus bristles thin, scabrous-ciliate, often smooth at tips; involu- Cral Gractsiehe Gta 208 Oe 1 Re A BO A Be hk Ieee 4. + Pappus bristles barbate, weakly or distinctly plumose; involucral bracts straight, appressed, or only middle ones recurved............... 6. 4. Capitula depressed-globose, solitary or terminal on branches; in- volucral bracts strongly divergent, erect, pubescent below, glabrous above, fleshy, thick, oval in cross-section, hollow; corolla almost white or light pink... ee eeeeeeeeeeeeee 3. O. prjachinii Tamamsch. + Capitula globose; involucral bracts glabrous or weakly or strongly pubescent, thinner than in preceding species, roundish in cross- 231 section, not hollow; inner bracts straight, outer ones erect............ 5. 5. | Whole plant tomentose-arachnoid-hairy, sometimes scarcely 1 m high; involucral bracts always covered with intertwined gray, long, arachnoidihains eter: 2 22. Ee 2. O. cinereum Grossh. + Pubescence of plant variable, sometimes completely glabrous; in- volucral bracts pubescent at beginning of flowering, later glabrous, if pubescent then never densely tomentose-arachnoid-hairy.............. buddies Wey, egos AU LAU AAC tis sel hd, Mean tae 1. O. acanthium L. 6. All involucral bracts appressed, more or less erect, similar; capitula oblongonelobOses-veeee Haid, keke, tes: eee, eA OS ile 4. + Outer and innermost involucral bracts appressed, middle ones at beginning of flowering erect, later recurved; capitula very large, globoser cede nutes: binaries 10. O. heteroacanthum C.A.M. Stem usually simple or with few branches from base.................... 8. + Stem solitary with short racemose branches and oblong-globose capitula; involucral bracts unequal; pappus of barbate bristles....... ta brectets mei cite, coma niet) concen ba 9. O. seravschanicum Tamamsch. 8. Plants snow-white and densely tomentose; capitula solitary, 2—2.5 cm in dia, on short solitary branches; involucral bracts appressed, spinose and densely pubescent, snow-white outside, ciliate-glandu- lar on margin, abruptly acuminate, with yellowish erect spine. Corolla pinkish-violetene)e: nuh Somebelestepes sree 8. O. candidum Nabelek + Plants not snow-white, but more or less densely grayish- or sordid- white pubescent; stem simple, sometimes weakly branched from middle or above. Capitula 3—6 cm in dia... eee eee eeseeeeeeeneeees 9. 9. Involucral bracts lacking spines or with inconspicuous spines, sub- glabrous. Pappus bristles ciliolate. Plants 30-65 cm high..................... barotits pants dues, Merits toe Boveub, as. casted | 5. O. armenum Grossh. + Involucral bracts with long, sharp spines, densely pubescent below. Pappus bristles plumose. Plants larger.............. 7. O. leptolepis DC. a Section 1. Onopordum.—Sect. Reflexa Rouy in Bull. Soc. Bot. Fr. III (1896) 581; Arénes in Not. syst. Paris, IX (1942) 215.—Outer involucral bracts curved or upward-spreading, numerous, very narrow or broad, coriaceous, green or pubescent, roundish or oblong in cross section, hollow, finely glandular on both sides or only outside. Corolla broadened only at throat or including limb, violet or purple, occasion- ally light colored. Pappus bristles ciliolate, smooth above, not plu- mose. Type of section: type of genus. Series 1. Acanthia Tamamsch.—Involucral bracts narrow, more or less equal, glandulare only on midrib and on margin outside, glabrous or arachnoid-hairy. 232 230 1. O. acanthium L. Sp. pl. (1753) 827; DC. Prodr. VI, 716; Ldb. Fl. Ross. II, 716; MB. FI. taur.-cauc. II, 281, III, 561; Boiss. Fl. or III, 559; Grossh. Fl. Kavk. IV, 192; Takht. and Fed. Fl. Erevana, 323; Dumbadze in FI. Gruzii, VIII, 522; Kozlovskaya in Fl. BSSR, V, 146; Gadjiev in Fl. Azerb. VIII, 420 p. p.; Kryl. Fl. Zap. Sib. XI, 2902; Nikitin in Fl. Turkm. VII, 247, p. p—0O. acanthium ssp. araneoso- tomentosum Rech. f. in Anz. Ost. Acad. Wiss. 88 (1951) 263.—O. acanthifolium Gilib. Fl. Lithuan. IIT (1781) 190, non C. Koch—Acanos spina Scop. Fl. Carn. ed. 2, II (1772) 132.—Ie.: Hegi, Ill. Fl. VI, 2, Wat272, fig ol: Biennial. Tall, mostly branched plant. Stem 0.3—2.0 m high, gla- brous or arachnoid-hairy, winged to tip, wings 6-15 mm wide, sinuate, narrower on branches, with broadly deltoid spiny lobs, spines to 5 mm long, stiff, yellow. Leaves unequal, lower ones 10—30 (35) cm long, 4-15 cm wide, oblong- or broadly ovate, lowermost narrowed into petiole; middle and upper cauline leaves ovate or ovate-lanceolate; all pinnately lobed, toothed, terminating into sharp spine, subglabrous or softly arachnoid hairy on both sides, with distinct, especially beneath, reticulate venation of uniformly thick veins. Capitula usually in clus- ters of 2-3 on upper branches, occasionally solitary, globose, 3-5 cm in dia. Involucre of immature capitula usually arachnoid-hairy, later glabrous or subglabrous; involucral bracts almost equal, inner ones erect, middle and outer ones recurved or horizontal, in lower appressed part ovate-lanceolate, subulate above, strongly spinescent, with occa- sional lustrous glands on midrib, along margin short-ciliate, and some- times with occasional glands. Corolla considerably longer than involu- cral bracts, purple or light violet, sometimes pink, very rarely white 2.0—2.3 cm long; corolla tube as long as limb including throat. Style branches almost entirely exserted from corolla. Achenes oblong or oblong-obovate, usually indistinctly triquetrous, slightly narrowed, 4— 6 mm long, transversely rugose, with inconspicuous veins along mar- gin, reddish-brown or brown with darker or grayish spots; pappus rubiginous, 1.5—2 times as long as achene; pappus bristles weakly ciliate. Flowering June to August. (Plate XI, Fig. 2). Weedy places, old fields and in crops as weed.—European Part: Almost all regions but Far North; Caucasus: all regions except high mountains; Western Siberia: Upper Tobol, Irtysh, Altai; Eastern Sibe- ria: Angara-Sayans (rarely); Soviet Central Asia: Kara Kum, Kyzyl- Kum, Tien Shbn, Pamiro-Alai, Aralo-Caspian Region, Lake Balkhash Region. General distribution: Central Europe, Atlantic Europe, Medi- terranean, Balkans-Asia Minor, Armenia and Kurdistan, Iran-Afghani- stan. Introduced in North America. Described from western Europe. Type in London. 231 Note. The hybrid O. acanthium L. x O. tauricum Willd., known from the vicinity of Yalta, is characterized by involucral bracts that are intermediate in form. 2. O. cinereum Grossh. in Tr. Tifl. Bot. Sada, II, 1 (1920) 38; Grossh. Fl. Kavk. IV, 192. Biennial. Whole plant densely grayish-arachnoid-tomentose. Stem 20—90(100) cm high, white, sulcate, laterally winged; wings 3—4 rowed, 5—6 mm wide, weakly sinuate, with short deltoid lobes and hard, yel- low or apically reddish-brown spine, equal to or 2 times as long as wing width; branches in upper part more or less spreading. Leaves, like stem, densely grayish-tomentose, 5-25 cm long, broadly oval, weakly pinnate-sinuate with deltoid lobes terminating into long, yel- low spine, soft and thick from dense grayish pubescence. Capitula large, to 7 cm in dia when open. Involucre globose, sordid-white- tomentose; involucral bracts densely tomentose below, densely arach- noid-hairy in middle, and subulate above, glabrous and with or without occasional glands on midrib, ciliolate, tiny-ciliate on margin. Achenes narrowed, but distinctly 4-angled, obovate, sometimes strongly nar- rowed below, brown or reddish-brown, marbled with black spots, trans- versely rugose, upper scar of attachment quadrate, hilum narrowly, oval, almost entirely concealed under fleshy appendage; pappus 2-3 times as long as achne; pappus bristles barbate at base, weakly ciliate above. Flowering July to August. Weedy places, roadsides, old fields in mountain zone.—Caucasus: Eastern Transcaucasia (south), Talysh; Soviet Central Asia: Kara-Kum, Amu-Darya (in extreme south). General distribution: Iran-Afghani- stan. Described from Talysh. Type in Tbilisi. 3. O. prjachinii Tamamsch. in Addenda, XXVII, 60S. Biennial. Plants 60—100 cm high, light colored, greenish-gray from pubescence. Stems branched above, to tip spiny-winged; wings in lower part 1.5—2.0 cm wide, gradually narrowed above, with network of distinct thin veins, undulate-sinuate, with hard thickish spine, decur- rent on midrib of deltoid lobes. Middle leaves 13—25 cm long, 2.5—4.0 cm wide, decurrent, spinescent, inequally pinnately lobed, lobes del- toid with long white or yellowish spine; upper leaves narrowly lan- ceolate, considerably reduced, sometimes entire, but always prickly at tip, like others on both sides uniformly grayish-arachnoid-hairy, be- neath with more distinct reticulate venation. Capitula large, subglobose, to 5 cm across, solitary at tips of erect branches. Involucre imbricate, oblong or globose in immature capitula; middle and inner involucral bracts erect, terminating in long, straight, yellow spine; bracts more or 234 232 less tomentose below, all bracts erect in mature capitula, outer ones sometimes strongly curved; all bracts almost equal, flat, oblong-ovate below; subulate above, glabrous, hollow in cross section, narrowed into yellow spine, lustrous inside, glabrous, often with constriction at point of transition of flat lower part to subulate upper part, ciliate, scabrous. Corolla light pink or almost white, 2—5—2.9 cm long; corolla tube 1.5—1.6 cm long, style branches and style considerably longer than corolla. Immature achenes 5—6 mm long, tetraquetrous, trans- versely rugose on sides; pappus yellowish or light golden, its longest bristles more than 2 times as long as achenes, one bristle usually thicker and longer than rest. Flowering May to July. In thickets of shrubs.—Soviet Central Asia: Pamiro-Alai. Endemic. Described from banks of Vakhsh River from M.I. Pryakhin’s collec- tions. Type in Leningrad. Series 2. Taurica Tamamsch.—Involucral bracts glabrous, densely fine glandular on both sides. Corolla limbs and throat uniformly ex- panded; filaments very finely glandular. 4. O. tauricum Willd. Sp. pl. III, 3 (1803) 1687; Boiss. FI. or. III, 559; MB. FI. taur.-cauc. II, 281, Ldb. Fl. Ross. II, 717.—O. virens DC. Fl. Fr. V (1815) 456.—O. virens B. tauricum DC. Prodr. VI (1837) 618.—O. viscosum Horn. apud Spreng. Syst. veg. 3 (1826) 386.—O. elatum Sibth. and Sm. Fl. Graeca, IX (1813) 23. Biennial. Whole plant subglabrous, green, slightly viscid from tiny glands. Stem stiff, thinly sulcate, to 50-60 cm high, with narrow wings; wings 2—3-rowed, unevenly sinuate, with deltoid lobes and long, very thin spines. Leaves large, oblong-lanceolate, irregularly coarsely toothed, glabrescent, green, weakly coriaceous; leaf lobes spinose, more densely glandular beneath; lower leaves to 15—20 cm long, 5-7 cm wide, middle and upper ones gradually reduced, with deltoid prickly lobes, with distinct light-colored midrib like lower, basally consider- ably broader. Capitula solitary, large, at tips of branches winged al- most to tip. Involucre 5—6 cm in dia, semi-globose; involucral bracts green, yellowish or reddish, large, elongate, to 5 cm long, almost equal, stiff, outer ones somewhat erect or deflected, like others with midrib slightly raised on outside, terminating in straight, subulate spine, with 3 distinct veins inside; corolla 3.0—3.2 cm long, its tube as long as limb including throat. Achenes gray, black-spotted, slightly compressed, 4-angled, with fine longitudinal veins on sides; pappus yellowish, some- times rubiginous, 2 times as long as achene but 1/2 as long as corolla tube. Flowering May to July. 236 236 233 Stony river banks, around estuaries, roadsides.—European Part: Crimea. General distribution: Balkans-Asia Minor; introduced in Cen- tral Europe. Described from Crimea. Type was in Berlin. Section 2. Erecta Rouy in Bull. Soc. Bot. Fr. III (1896) 581.— Involucral bracts appressed, flat, glabrous or pubescent, sometimes only middle bracts deflected. Pappus bristles ciliate, barbate, or plu- mose. Corolla light-colored, pinkish, creamish or whitish. Type of section: O. leptolepis DC. Series 1. Armena Tamamsch.—Pappus bristles ciliate; involucral bracts appressed, lacking spine or with scarcely noticeable spine. 5. O. armenum Grossh. in Tr. Tifl. Bot. Sada, II, 1 (1920) 40; Grossh. Fl. Kavk. IV, 192; Takht. and Fed. Fl. Erevana, 323. Biennial. Whole plant, except capitula, light grayish pubescent. Stems 30-65 cm high, simple or dichotomously branched from middle, deeply sulcate, 1 cm in dia, winged and spinose almost to capitula; wings 4—5-rowed, weakly sinuate, to 1 cm wide in lower part of stem, above gradually strongly reduced, weakly sinuate, with yellowish spines 4-8 mm long. Leaves large, on both sides tomentose, lower leaves with more or less winged petiole, middle cauline leaves decurrent, 6— 15 cm long, 2-6 cm wide, ovate-lanceolate, shallowly pinnately lobed, with broadly deltoid, unequal lobes bearing yellow spines, midrib broad and tomentose at base on both sides; upper leaves like middle, gradu- ally reduced, absent at tip. Capitula solitary, 3—S cm in dia, oblong- or broadly-globose, slightly flattened. Involucral bracts imbricate, outer- most tomentose, others glabrous, sometimes lustrous, strongly ap- pressed, lacking spine or with short, scarcely noticeable apical spine; outer bracts about 1 cm long, middle and inner ones longer, ovate-lan- ceolate, often violet at tips, ciliolate scabrous, smooth inside, stramineous, lustrous, greenish inside along margin. Corolla light pinkish, sometimes almost white, 3.0-3.5 cm long; corolla tube 1.8—2.0 cm long. Style branches strongly exserted from corolla. Apical appendages of anthers violet. Achenes weakly tetraquetrous, 5—6 mm long, 1/2 as long as pappus; pap- pus bristles ciliolate, unequal, as long as or shorter than corolla tube, basally connate in 4—S-angled ring. Flowering June to August. Weedy places, stony slopes, roadsides.—Caucasus: Eastern and Southern Transcaucasia. Endemic. Described from Dzervesh in vicin- ity of Erevan. Type in Tbilisi. 6. O. frickii Tamamsch. in Addenda, XXVII, 605. Annual or biennial. Plants more or less grayish-pubescent. Stem 234 Plate XI. 235 1—Onopordum heteracanthum C.A.M., 2—O. acanthium L., achene, pappus, pappus bristles. 237 235 80-100 cm high, branched above or from middle; branches many, long, upward spreading, winged like stem; wings grayish-arachnoid- hairy, weakly sinuate, 0.5—0.8(1.0) cm wide, with tiny, yellow, short spines. Middle leaves broadly-lanceolate, 12 cm long, about 2 cm wide, sessile, short-decurrent, remotely coarsely toothed; teeth some- times bidentate and with yellow spines of different length, gradually reduced in upper part, sessile, long-decurrent, narrowly-lanceolate with spine 1—2 mm long; upper leaves like lower soft from dense arachnoid pubescence on both sides, and with prominent midrib beneath. Capitula solitary, terminal and axillary, 3 cm long, 1 cm in die, ob- long-cylindrical. Involucral bracts imbricate, flat, most diverse in size, outer ones 0.6—-1.2 cm long, ovate-lanceolate and lanceolate, subse- quent rows of bracts gradually longer, somewhat curved, linearly-lan- ceolate; innermost bracts from oblong, 0.6 cm long, 0.3 cm wide, glabrous base narrowly subulate, darker on midrib and along margin glandular ciliate; all inner bracts glabrous, light stramineous, lustrous. Corolla almost as long as inner involucrat bracts, light colored. Achenes 5—6 mm long, 2-3 mm wide, ovate, slightly narrowed, 3—4-angled, with 3—4 thin longitudinal veins along sides, transversely rugose, gray, sometimes with black longitudinal spots along ribs, upper scar raised and hilum almost completely concealed under thick appendage; pap- pus setose, indistinctly two-rowed or one-rowed; pappus bristles ciliolate, almost smooth, unequal, of which longest (one of them thick- est) 1/2 as long as inner involucral bracts and 2 times as long as achene. Fields, rocky places.—Caucasus: Southern Transcaucasia. Endemic. Described from collections of Frick. Type in Leningrad. Note. A very distinctive plant, distinguished by weak spines and relatively small capitula—both characters that in general are not char- acteristic of this genus. Series 2. Leptolepidia Tamamsch.—Involucral bracts narrow or more or less broad at base, usually erect, with long spine, or middle involucral bracts deflexed; pappus bristles barbate or plumose. 7. O. leptolepis DC. Prodr. VI (1837) 619; Boiss. Fl. or. III, 564; O. and B. Fedtsch. Perech. Rast. Turk. IV, 276; Grossh. Fl. Kavk. IV, 191.—O. lanigerum Boiss. 1. c. (1875) pro syn.—O. heteracanthum var. ortholepis Bornm. in Beih. Bot. Zentralbl. LX (1939) 302 p. p. — O. olgae Rgl. and Schmalh. in Izv. Obshch. Lyubit. Estestv., Antrop. i. Etn. 34, 2 (1882) 51; O. and B. Fedtsch. op. cit. Biennial. Tall, densely white-pubescent plants. Stem straight, soli- tary, more or less branched or simple, fistular, broadly-winged; wings 238 236 5—7-rowed, weakly sinuate, lobes toothed, with yellowish spines. Leaves oblong-lanceolate or elliptical, densely lanate on both sides; lower leaves pinnately parted with ovate or sinuate lobes, spiny-toothed, narrowed into straight, yellow, glabrous spine. Capitula 4—6 cm in dia, oblong-broadly ovate or globose, solitary on apical branches, some- times in axils of upper leaves, sessile and then usually undeveloped. Involucral bracts appressed, straight, deflected even at blooming, gradu- ally narrowed, lanceolate, dorsally floccose; spines erect, hairy, at- tenuate, triangularly accuminate; bracts more or less glabrous below, ciliolate only along margin, narrowed above, along midrib on outside and often inside, ciliate on margin; innermost bracts 3.0—3.5 cm long, to 2 mm wide in broadest part in large capitula, gradually narrowed into tip 1.2-1.5 cm long, scabrous outside, glabrous inside, lustrous, almost as long as light-colored corolla. Achenes narrow, 6 mm long, 1.8—2 mm wide, distinctly tetraquetrous, brown or light brown, ob- liquely rugose on sides, longitudinally finely sulcate, with 2—3 distinct veins and sometimes with darker spots; ribs well developed, broader above; upper scar of achenes with scaly, more or less quadrate border, with distinct tubercle, hilum straight, to 1/2 or more concealed by callous appendage; pappus 2.0—2.5 cm long, 4 times as long as achene; bristles plumose, one of them thicker, to 3 cm long. Flowering May to June. Mid-montane zone.—Soviet Central Asia: In all regions, except northern part of Aralo-Caspian. General distribution: Armenia and Kurdistan, Iran, Dzhungaria-Kashgaria. Described from Iran. Type in Paris. Note. The hybrid O. leptolepis DC. x O. acanthium L. is known from Samarkand. 8. O. candidum Nabelek. It. turc.-pers. II (1925) 35. Biennial. Plants arachnoid-hairy, snow-white. Root simple, dark brown at neck. Stem simple, occasionally weakly branched, solitary, with solitary capitulum, erect, 18—40 cm high, indistinctly angular below pubescence, with decurrent leaves, narrowly-spiny-winged, leafless above. Basal leaves large, falling before flowering; cauline leaves recurved, oblong, ovate, or narrowly-elliptical, 4-5 cm long, coarsely toothed, pinnate-undulate or pinnately parted, lobate with spine to 4 mm long, narrowed into petiole 5—7 cm long; upper and middle leaves sessile, 4—9 cm long, 2—3 cm wide, gradually reduced to simple spine, 1 cm long, 0.5 cm wide; all leaves soft, densely white-arachnoid-hairy above, more densely snow-white-tomentose beneath. Capitula solitary, ovoid-cylindrical, basally narrowed, lacking constriction, pedunculate, 3.5—4.5 cm long, 2—2.5 cm in dia. Involucral bracts appressed, spinose 239 237 or densely arachnoid-hairy snow-white, at tip sometimes colored, cili- ate-glandular along midrib and margin, with yellowish upward-spread- ing spine, triangular or lanceolate below, 8-10 mm long, 2-3 mm wide. Corolla pinkish-violet, about 3 cm long; corolla tube 1.5 cm long; limb linear, uniformly broad; style branches to 3.4 cm long, exserted from corolla. Achenes tetraquetrous, dark brown, transversely rugose; pappus to 14 mm long; bristles barbate, thin, one of them erect, longer and thicker than rest. Flowering July to August. Mountain slopes, dry stony places, uncultivated fields —Caucasus: Southern Transcaucasia. General distribution: Asia Minor. Described from vicinity of Van. Type in Bratislava (Czechoslovakia). Note. Nabelek cites fragrant florets as a distinctive feature of this species; however, fragrance is charactersitic of most species of this genus in general. 9. O. seravschanicum Tamamsch. in Addenda, XXVII, 605. Biennial. Short light gray plant. Stem thick, almost uniformly thick from bottom to top, to 1 cm in dia, angular, weakly sulcate, densely white-tomentose, divaricately branched, with solitary terminal capitula at tips of branches, sometimes with 1—2 undeveloped capitula in axils of upper leaves; wings all over stem and branches 1.0—1.5 cm wide, undulate-sinuate, lobes of wings deltoid or plain, with thin spine, 0.5— 0.7 mm long, yellow, densely grayish-pubescent like leaves. Basal leaves about 12 cm long and 2.5 cm wide with deltoid lobes; often bidentate, terminating into thin, yellow spine almost as long as width of lobes, lamina soft from dense indumentum above, denser beneath, rugose- crisped, especially at base with light colored, thick veins, more promi- nent beneath, spinescent; upper leaves gradually reduced, below capitula 2 cm long, 0.5 cm wide, lanceolate, entire or denticulate, but always spinose and with thick midrib. Capitula oblong, 3.5—-4.0 cm long, 1.5—2 cm wide. Involucre oblong-globose, basally somewhat constricted, more or less glabrous; outer involucral bracts very small, with cusp 0.5—0.6 cm long, ovate-lanceolate, middle-linearly-lanceolate, somewhat divergent, to 2—2.5 cm long, spinescent, like outer bracts glabrous at base, in middle weakly arachnoid-hairy, glandular above and on midrib, ciliate-glandular along margin; innermost bracts 2.5— 2.8 cm long, 1.0—1.5 mm wide, at base narrowly-linear, subulate, gla- brous below, glandular and ciliate-glandular above and along midrib and margin, pale violet. Corolla whitish, with occasional small glands, 2.8 cm long; corolla tube shorter than limb. Style branches exserted from corolla. Immature achenes 4—5 mm long, reddish-brown, unequally 4—5 angled; pappus white, barbate; pappus bristles unequal, one of them longest reaching throat. Flowering July to August. 240 238 Alpine zone.—Soviet Central Asia: Pamiro-Alai. Endemic. Described from vicinity of Rarez from Zeravshan. Type in Leningrad. Note. The species, possibly of hybrid origin, resembles O. polycephalum Boiss. in habit, which is distributed in the subalpine zone of the Sicilian Taurus. A highly ornamental plant. 10. O. heteracanthum C.A.M. Verzeichn. (1831) 68; DC. Prodr. VI, 618; Ldb. Fl. Ross. II, 717; Boiss. Fl. or. III, 563; Grossh. FI. Kavk. IV, 192; Dumbadze in Fl. Gruzii, VIII, 522; Gadjiev in FI. Azerb. VIII, 420.—O. heteracanthum £. anisacanthum Boiss. 1. c. (1875).—O. anisacanthum Boiss. Diagn. ser. I, 10 (1849) 93.— O. carmanicum Bornm. in Beih. Bot. Zentralbl. LX (1939) 302 p. p. Biennial. Light grayish and densely pubescent plant. Stem up to 50-60 cm high, occasionally shorter, branched, branches, like stem, to 1 cm in dia, ribbed-angular, densely white-pubescent, to top with many rows of spiny wings; wings in lower part of stem 1.5—1.8 cm wide, on both sides white-tomentose-arachnoid-hairy, weakly sinuate but with very long, light colored spine. Leaves greatly variable; lower leaves petiolate, 15-26 cm long, 4-8 cm wide, shallowly or deeply lobed almost to midrib into large deltoid spinescent lobes with long thick spine sometimes longer than lobe; middle and upper leaves gradually reduced, lanceolate, almost undivided, but with long spine, like lower leaves tomentose-arachnoid-hairy on both sides, with thick midrib and distinct lateral veins above. Capitula solitary, terminal on branches, large, unopened about 7 cm in dia, opened 10—11 cm, usually flat- tened-globose. Involucre surrounded by lanceolate apical leaves, some- what appressed above, densely white-pubescent, glabrescent; involu- cral bracts dissimilar, 3—4-rowed, small, lanceolate, with short tip, imbricate, appressed; middle bracts cuneate, sub-glabrous above, often violet or purple, with thick bristles along margin, spinescent, deflected in upper half, inner bracts straight, shorter than florets, often colored, with whitish spine. Corolla large, bright pinkish, almost white, 3.5—3.8 cm long; corolla tube very narrow, considerably longer than limb. Achenes 5—6 mm long, oblong, somewhat narrowed below, distinctly tetraquetrous, ribbed, on sides grayish, with many longitudinal furrows and transverse dark brown striations, rugose, not maculate, hilum straight, somewhat concealed by thick appendage, upper scar almost quadrate; pappus long, longest bristles barbate, bristles longer than corolla tube and 3 times as long as achene, basally connate in angular, waxy ring, to 1 mm in dia. Flowering May to July. (Plate XI, Fig. 1.) Up to mid-montane zone, roadsides, in weedy places, abandoned fields —Caucasus: Eastern and Southern Transcaucasia. General dis- tribution: Iran-Afghanistan. Described from Zuvant. Type in Leningrad. 24 — 289 Subtribe 3. Centaureinae O. Hoffm. in Pflanzenfam. IV, 5, (1893) 324.—Trib. Carduaceae Cass. in Dict. Sc. nat. VII (1817) 94 p. p.— Trib. Centaurieae Cass. In Dict. Sc. nat. VII (1817) 376, X (1818) 156, XX (1821) 358, L. (1827) 246, LX (1830) 370; Opusc. phyt. I (1826) 294, II (1826) 202.—Trib. Carduineae Cass. in Dict. Sc. nat. X (1818) 156 p. p., XX (1821) 359 p. p., XLI (1826) 308 p. p., L (1827) 463; p. p.; LX (1830) 571 p. p.; Opusc. phyt. I (1826) 296 p. p., II (1826) 202 p. p.—Subtrib. Centaurieae Less. in Linnaea, VI (1831) 85 and Synops. Comp. (1832) 4 p. p.; DC. Prodr. VI (1837) 557; Benth. in Benth. and Hook. f. Gen. pl. II (1873) 214; Boiss. FI. or. III (1875) 585.—Subtrib. Serratuleae DC. op. cit. 662 p. p—Subtrib. Characopappeae Sch. Bip. Div. Centaurineae Sch. Bip. in Linnaea, XIX (1846) 325.—Subtrib. Characopappeae Sch. Bip. Div. Serratuleae Sch. Bip. ibid. 326 p. p.—Capitula heterogamous, with one row of sterile, often campanulate peripheral florets, mostly longer than bisexual central florets, or homogamous. Achenes with lateral, ob- lique, very rarely straight, areola or attachment (scar [or hilum]), gla- brous, less often hairy, usually with pappus; pappus simple, compris- ing 2 or more rows of bristles or setaceous scales, or double, inner row comprising mostly considerably shorter scales, scale-like bristles or [simple] bristles. KEY TO GENERA OF SUBTRIBE CENTAUREINAE 1. Capitula enclosed by spiny-toothed, toothed-ciliate, stiffly, setose, rarely/almost entirelsapicalsleaviesmsadin..:2krrnle See 2 ae 2. + Capitula not enclosed by apical leaves................eeceeeseeeeeeeereeeenes 5, Capitula small, (4-7 mm in dia, excluding recurved spines and cusp); achenes compressed, not angular, lacking ribs, smooth...... 3. + Capitula larger (10-25 mm in dia, excluding squarrose outer involucral bracts); achenes angular or ribbed, large (4—8.5 cm NS 3. Involucral bracts herbaceous, white-membranous along margin sa at tip, terminating in stiff-membranous, nonprickly, subulate cusp; florets yellow; achenes rather large (3.5—4.5 mm long). Plants stemless or almost stemless; leaves on margin with white, thin, stiff, mostly palmately branched bristles................eeeeeeceeeeeeeeeeeeeeeees PS ANAS ie Retr rae ere em ae er 1618. Schischkinia [ljin + Involucral bracts coriaceous, outer ones at tip with oblong append- age, on margin with toothed-ciliate, green, leaf-like appendage, not decurrent on bracts; middle bracts with long (10-22 mm), simple, strongly recurved, white spine; florets lilac-pink; achenes small 242 240 (2—2.3 mm long). Plants with stem, stem dichotomously spreading- branched, mostly from base; leaves with stiff, simple, subulate cilia dlone 'marein'(ef: also couplet 27)203:.aly ee) ARI seem A oy 1624. Centaurea L. (subgenus Tetramorphaea DC. Czer.) Outer and middle involucral bracts more or less gradually narrowed into lanceolate, ovate-lanceolate, or ovate, pinnately, spiny-toothed, rarely almost entire, acuminate, or spiny-cuspidate, green, leaf-like, coriaceous, appendage. Achenes tetraquetrous, obovate, lacking, pa- ppus or with pappus of numerous flat bristle-like scales................... ANAL ARE BAL dep ate heed 1626. Carthamus L. Outer involucral bracts with long, filiform, stiff, appendage at tip, middle and inner bracts with narrowly lanceolate, coriaceous appendage; spiny-pinnate along margin; achenes many-ribbed, oblong-cylindrical, with pappus of few, narrow bristles ............. sal, wine aee.. Chea eet nel peepee 1627. Cnicus L. Capitula homogamous; all florets bisexual, rarely unisexual (and then plants dioecious), fertile; sometimes capitula heterogamous, with peripheral pistillate florets, and large, central bisexual flo- nets. .Aduc senuetions atekoaiag iain sete heel on mina Sa 6. Capitula heterogamous; peripheral florets sterile, sometimes with staminodes, small, tubular, or large and then tubular-infundibuli- formycentrall florets bisexualy fertile xt. 20). ee ee. ee 16. Receptacle with few, very readily deciduous, narrow-lamellate, long bristles; floret corolla with tube long pubescent in upper part; achenes appressedseniceous 22)... 2h WO... 1603. Russowia Winkl. Receptacle densely covered with persistent, narrow bristles; floret corolla glabrous; achenes entirely glabrous or very rarely with scattered, readily deciduous, long hairs, and almost glabrous at Thatunibyastone ter leh en J ee nh women Bb Ue ee celta as 2 Ws Outer and middle involucral bracts acuminate, cuspidate, awned or spinescent, rarely obtuse; achenes glabrous...............cccscccceseceeeeees 8. Outer and middle involucral bracts with scaly or membranous api- cal appendage, sometimes as very narrow border, weakly delimited from coriaceous part of bracts, but then involucral bracts roundish and achenes (especially immature) scatteredly pilose................. 1s Florets yellow. Cauline leaves sessile, amplexicaul, short-decurrent (Kopetdae)minaccevme eek eet. Leila eae 1610. Schumeria I]jin Florets pink, violet, lilac-purple, purple, reddish, rarely white (al- bins) che). 8). Ales. 22. Laur, 6.00) ee bene lly, cumin on meee > Achenes small (2—2.5 mm long), with straight, punctate hilum; pappus double, outer one consisting of few, scabrous, hairlike bristles, inner of 1 (rarely 2) scabrous, stiff bristles, appreciably broadened below, longer than outer pappus bristles, 243 10. Ids. 13. 14. and 3—5 small, smooth, unequal scales. Annual plants................... Ji A racer neenteacsa sees va duncemdeaialae. ines anterad 1617. Oligochaeta C. Koch Achenes large (4-8 mm long); pappus simple, consisting of mostly numerous weakly scabrous, serrate, less often short-plumose bristles, gradually becoming longer from outer to inner. Perennial polaintSe-crahiedset nected tibiae cemeed eeih un Rear Races 10. Stamen filaments densely, more or less faneoducatans involu- cral bracts very narrow (about 1 mm wide), outer bracts subulate, middle apically more or less abruptly attenuate into fine hairlike, 8—12-mm-long awn. Semishrub with thick woody root and many- headed caudex; stems numerous (20-30) with virgate branches. Leaves rather fleshy, narrow, sessile; middle and upper cauline leaves linear and narrow-linear, entire or with 1—2 small teeth at base, erect (southern Darvaz Region)........... 1606. Tricholepis DC. Stamen filaments glabrous or with very small, inconspicuous papil- lae; involucral bracts wider, lacking thin and hairlike, more or less long-awned tips. Plants of different habit...............cceeeeeeeeeeeees bl. Achenes with straight hilum. Basal appendages of anthers cleft into two hairlike bristles. Leaves entire or subentire; cauline leaves sessile easereeaee vereeglist aeretlao, wlaehests sneer. 4h 1609. Syreitschikovia Pav]. Achenes with oblique hilum. Basal appendages of anthers undi- Anthers free, connate only by basal appendages, which are fused on inner side with filaments. Capitula drooping, large. Leaves broadly triangular-ovate, green above, glabrous, white tomentose beneath; basal and lower cauline leaves sinuate-lobate with cordate base...... scott eines ceaes eentasess tr vauts dee getter bee Bho 1608. Synurus Iljin Anthers connate in anther tube. Capitula not drooping, mostly medium-sized or rather small. Leaves of different shape, green, sometimes glaucous, glabrous or scabrous from scattered, usually somewhat stiff hairs, pinnately incised, pinnately parted, lyrate, lobate, less often undivided and entire............... 1607. Serratula L. Achenes tetraquetrous, more or less ribbed or rugose-tuberculate, truncate at tip, with well-developed crown around margin; append- ages of outer and middle involucral bracts large, round, broadly ovate, or ovate, often divided into several lobes, convex, membra- nous,notdecurrpentionsbracte ters. 22s) |we. deowied di aes es: 14. Achenes not angular, smooth, apically rounded, without crown around margin; appendages of involucral bracts of different struc- CUS pelted oh HAS ede esl uses absent boctilneut tydeah. calneieh eames een ule iS. Annual plants, with slender roots, scabrous due to sharp spine-like hairs; leaves sessile, pinnately incised with narrowly linear, entire, decurved segments terminating in small bristle. Pappus bristles 244 242 15. 16. by: 18. Serrate eresilientel!s...2 2 Aen ev nen tess 1615. Callicephalus C.A.M. Perennial plants, with long or shortened rhizomes, weakly arach- noid-hairy tomentose, rarely glabrous. Leaves pinnately incised, pinnately parted or lyrate, with almost round, ovate, elliptical or lanceolate, toothed or lobed segments, less often undivided basal and lower cauline leaves petiolate. Pappus bristles short-plumose on barbate; SOftens..Nern. 22.0.0. etree 1611. Rhaponticum Adans Plants entirely glabrous, with virgate, vertical root, not producing suckers. Outer and middle involucral bracts with very narrow, semicoriaceous or stiff-membranous, glabrous upper margin, inner with small, wide, membranous appendage. Filaments papillose. Achenes sparsely pilose, mature achenes almost glabrous, with lat- eral hilum; pappus as long as achene, pappus bristles serrate (cf. alsorcOuplet 22)... ceassh Aen Saeed 1605. Plagiobasis Schrenk. Plants arachnoid-pubescent, with very long root and with lateral, more or less horizontal or obliquely ascending suckers. Append- ages of involucral bracts scarious, in outer and middle bracts wide, semi-transparent, with short-hairy border, innermost bracts very narrow, linear-lanceolate, densely pilose. Filaments glabrous. Achenes glabrous, with very slightly oblique hilum; pappus usually 3 times as long as achene, pappus bristles serrate-ciliolate, inner- most bristles short-plumose above.................. 1616. Acroptilon Cass. Achenes sericeous or rather profusely appressed-hairy. Exclusively annualsor.‘byennial! plants) . sili ieed. ede eased nee rete es cates We Achenes glabrous or scatteredly hairy, subglabrous when MMMATUIES FIs, GL Ye. TLGST OAS OLAS. Sook SOE PE ed TON SS DRE. 18. Involucral bracts herbaceous, from acute outer to long-acuminate inner ones, lacking appendages. Florets purple, few (only 1-6 achenes develop); peripheral florets not enlarged, tubular, with 5- parted corolla. Pappus double, outer one of serrate-scabrous, stiff bristles; inner of S—10 smooth, small scales. Leaves sessile, pin- nately divided, with narrowly-linear segments..............:ceeseeeeseeeeeeee ToT SLIT TRIN, Beco. Bh SANE ge aan 1604. Crupina Cass. Involucral bracts coriaceous, outer and middle ones obtuse, lacking appendages, inner ones with small membranous appendages; florets pink or yellow, numerous, usually strongly enlarged, with infundibu- liform, 4—20-lobed corolla; pappus simple, of serrate-toothed seta- ceous scales. Leaves toothed, pinnately lobed or pinnately parted; basal and lower cauline leaves with quite long petioles................ wceeacenvatsneamuivkse meatie aaneaaree ata sctid el moat rata tae 1611. Amberboa (Pers.) Less. Pappus bristles plumose. Perennial plants..................eeseesseeeeeeee 19. Pappus bristles scabrous, serrate or serrate-ciliolate or achenes lack- INS *pPappus. AAI. ABLAS. IN. aL Se, ee eee ose Ze 245 20. ale 22 23). 243 Florets bright pink, peripheral ones quite enlarged, tubular-infundibu- liform; pappus simple; outer involucral bracts membranous, more or less densely short-pubescent, fimbriate-ciliate; appendages of middle and inner involucral bracts like outer ones, large, ovate, entirely covering bracts. Rhizomatous plants; leaves undivided; lyrate-pinnately divided or incised, arachnoid-hairy or densely arach- noid-tomentose (mostly beneath), less often glabrous.................... le, PERM Ss CR OE DR 1619. Aetheopappus Cass. Florets yellow, peripheral ones not enlarged, tubular. Combination Of other/characters: different. 13, (RRs. Pere eine etl. catheltved Sonco0e 20. Appendages of involucral bracts more or less large, membranous, almost round to ovate, mostly lacerated, erose-toothed. Achenes scatteredly hairy; pappus double. Leaves entire, scabrous from spine-like hairs; cauline leaves sessile, long-decurrent on stem as Wak Ge, LEER UCD, AESOP, HE og. 28 1614. Chartolepis Cass. Appendages of involucral bracts small, coriaceous, with palmately divaricate, almost equal, small spines. Achenes glabrous; pappus simple. Leaves entire or sinuate-pinnately lobed, scabrous from stiff hairs; cauline leaves sessile, not decurrent ................ prec uan pond sereeowcunt oa AE RORLE. ROMANE cease 1620. Cheirolepis Boiss. Achenes apically rounded; florets pink. Perennial plants with en- tire, toothed, or more or less pinnately lobed leaves.................... D2. Achenes apically truncate, with small-toothed, rarely almost obtuse CLOW Ma sceicascacsaszecasscassectse che sane Mate ee aah bttds at Reon cette NRO ta on 23. Plants entirely glabrous; root unbranched, vertical. Outer and middle involucral bracts with very narrow, inconspicuous, semicoriaceous or stiff-membranous, entire border at tip. Peripheral florets with staminodes, not enlarged, tubular. Achenes with simple pappus of few very readily deciduous, serrate bristles (cf. also couplet LES BYES. Beare ected. MOLE. ae zoe ee 1605. Plagiobasis Schrenk Plants pubescent, with long, more or less branched rhizome. Appendages of outer and middle involucral bracts like wide mem- branous margin, more or less decurrent on bracts acuminate or spinescent, pectinate-ciliate. Peripheral florets without staminodes, enlarged, tubular-infundibuliform; achenes lacking pappus ........... I crea eM ot panei aN IN Da ah te 1623. Phalacrachena I]jin Stamen filaments entirely glabrous. Outer and middle involucral bracts with thin-coriaceous or membranous, broadly triangular or lanceolate, pectinate-ciliate apical appendage, cuspidate or with long- awn; inner bracts at tip irregularly short-toothed, setose-fimbriate or setaceous-awned. Annual plants.................:cesssesseeeseeeeesneseneees 24. Stamen filaments covered with papilliform hairs or papillae. Involucral bracts at tip with diverse appendages, less often lacking 244 appendagesieiierciyes cuaw urns laeterl adit) Mek celal meted a 25. 24. Florets saffron- or golden-brownish-yellow, flesh-colored or white; peripheral florets lacking staminodes, enlarged; all achenes with pappus, with 2-4 transverse furrows above, below crown; pappus 2 times:as longias: achenesscwd Siete ates: 1602. Zoegea L. + Florets yellow; peripheral florets with staminodes far exserted from corolla tube, not enlarged. Achenes lacking transverse furrows, smooth, outer ones often lacking pappus; pappus almost as long as achene or slightly shortet................:ccsccsesseeees 1622. Stizolophus Cass. 25. | Achenes tetraquetrous, more or less transversely rugose, glabrous; pappus simple, bristly, long (12-16 mm), 2 times as long as achenessau.sead. det eee terme adbess 1621. Tomanthea DC. + Achenes not angular, smooth, scatteredly short-hairy (mature achenes more or less glabrous), very rarely glabrous (subgenus Centaurea of genus Centaurea L.); pappus double, sometimes simple, but then shorter (to 6 mm long), much shorter than or equal to achenes, less ofteniachenes lacking pappusifnemen: kudos erivele he een ate 26. 26. Scales of inner pappus much shorter than scale-like bristles of outer pappus, 8, broadly linear, more or less hairy on outer side, with tuft of rather long hairs at apex. Annual plants.....00....0.....eeeeeeneeeeeees Oia ies eres oc Mon ive ogee 1623. Hyalea (DC.) Jaub. and Spach. + Scales of inner pappus (often strongly reduced or not differing in size and shape from bristles of outer pappus), always glabrous on outer side, lacking tuft of hairs at apeX............ceeeeseeeeeeeeeeeeeeneee 2. 27. Capitula very large, (20) 30-50 mm in dia; peduncles strongly thickened below capitula; appendages of involucral bracts membra- nous, irregularly fimbricately lacerate or deeply palmately divided into stiff cilia; florets yellow (Transcaucasia)............::ccsscceseseseeeeees achat, aa ataeentabcoa beeline 5, Sas 1613. Grossheimia Sosn. and Takht. + Capitula usually smaller; peduncles not thickened below capitula; appendages of involucral bracts of diverse shape and size (cf. also Couplet3 iectecsel: Newt never elle cece, herevenai, te 1624. Centaurea L. GENUS 1602. Zoegea L.':? L. Mant. pl. (1767) 15; DC. Prodr. VI (1837) 562; Boiss. Fl. or. III (1875) 697; Czer. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX (1959) 443; in Fl. Turkm. VII (1960) 248.—Cyanus sect. Zoegea (L.) DC. in Ann. Mus. Paris, XVI (1810) 158. 'Treatment by S.K. Czerepanov. *Named after J. Zoega, a student of Linnaeus and investigator of the flora of Iceland. 247 245 Capitula heterogamous, medium- or quite small-sized, with many florets; at tips of stem and lateral branches, solitary, in lax paniculate or almost corymbose inflorescence. Involucres ovate, 6-12 mm in dia (excluding spreading cilia), weakly tomentose to glabrous; involucral bracts many-rowed, imbricate, almost herbaceous, with veins greenish (somewhat yellowing during drying), lanceolate, acute; appendages of outer and middle bracts not decurrent on bract, membranous, lanceolately triangular, pectinate-ciliate, setaceous cuspidate or long- awned at tip; cilia of appendages thin, hair-like, serrate-scabrous; in- ner involucral bracts white in upper part, irregularly short-toothed to long-awned at tip. Receptacle setose. Florets saffron- or golden-brown- ish-yellow, flesh-colored or white, heterogeneous; corolla glabrous, central florets (disk florets) bisexual, tubular, several-rowed, very slightly shorter than involucre, with deeply 5-parted corolla, periph- eral florets (ray florets) sterile, lacking staminodes and rudiment of style, tubular infundibuliform or tubular, 1-rowed, broadened, longer than involucre, with more or less shallow 4-parted, sometimes 4-toothed corolla. Anthers with small basal appendages, their apical appendage obtuse; stamen filaments flattened, glabrous. Style woolly above, with collar of hairs, bifid, distinctly exserted from anther tube. All achenes similar, ellipsoid-obovate to oblong, 2-5 mm long, truncate above, with crown of short teeth, compressed, smooth, only above with 2—4 transverse furrows, weakly hairy or glabrous scar lateral; pappus longer than achene, persistent, double, outer one of several rows of serrately scabrous, white or very slightly reddish bristles, from very short outer to inner longer ones, inner 1/3 or 1/4 as long as outer, of 10 oblong- spatulate, fimbriate at tip, brownish-green, rarely brownish-purple, broad scales, or subulate, smooth, very narrow bristles. Annual herbs with erect, mostly well-built, branched stem, moderately leafy almost to tip and scabrous lower leaves mostly lyrate, pinnately incised or lyrate, short-petiolate; middle and upper cauline leaves entire, subsessile. Type of genus: Z. /eptaurea L. Seven species belong to the genus Zoegea L. They are distributed from the Sinai Peninsula to the southwestern Pamiro Alai. 1. Florets golden-brownish-yellow; achenes 3.5—4.0 mm long; inner pappus of achenes Scaly..............:scseeees 1. Z. baldshuanica Winkl. + Florets white or flesh-colored; achenes 2.0—2.2 mm long; inner pappus of achenes bristly.............cccssesseseeees 2. Z. purpurea Fresen. Section 1. Zoegea—Czer. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX (1959) 446.—Capitula medium-sized; appendages of outer and middle involucral bracts bristly at tip, cuspidate; cusp shorter 246 than lateral cilia; inner involucral bracts irregularly short-toothed above, bristly-toothed or bristly-fimbriate with distinct bristle-like cusp, often lacking cusp; florets saffron- or golden-brownish-yellow, large (corol- las of central florets 10-13 mm long, of peripheral florets 15—28 mm long, tubular-infundibuliform); achenes weakly undulate; inner pappus scaly, of 1 row of oblong spatulate, fimbriate at apex, brownish-green, rarely brownish-purple broad scales. Type of section: type of genus. Of the six species belonging to this section, only one is found in the USSR. 1. Z. baldshuanica Winkl. in Tr. Peterb. Bot. Sada, IX, 2 (1886) 426: O. and B. Fedtsch. Perech. Rast. Turk. IV, 273; Czer. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX, 449; in Fl. Turkm. VII, 249.—Z. crinita auct. fl. As. Med. non Boiss.—Ic.: Czer. Fl. Turkm. VII, Plate XXXIX, fig. 1.—Exs.: Sintenis, It. transcasp.-pers. 1900- 1901, No. 180. Annual. Whole plant weakly scabrous from short spiniform hairs. Stem 20-—80(100) cm high, erect, finely ribbed, branched from middle or above, lateral branches spreadingly upcurved, well-shaped, more or less long, with fewer branches, sometimes simple, glabrous above. Leaves somewhat thick, acuminate or acuminate-cuspidate, with punc- tate sessile, glands; lower, often middle, cauline leaves also lyrately pinnately incised or lyrate, short-petiolate, their lateral segments mostly lanceolately linear, entire, 1-2 on each side, terminal segment 2-3 times as long as lateral, narrowly lanceolate or lanceolately linear, entire, sometimes remotely crenate; middle and upper cauline leaves lanceolately linear or linear, entire, subsessile, middle ones (3) 4—6 cm long, 0.3—0.6(0.8) cm wide. Capitula crowded in lax paniculate or almost corymbose inflorescence. Involucre elongately ovate, (15)18-20 mm long, (8)10—12 mm in dia (excluding spreading cilia), weakly tomentose; outer and middle bracts with membranous, pecti- nate-ciliate appendages; cilia of appendage slightly reddish, somewhat spreading, hairlike, 3—6 times as long as thickness of slightly tubercu- late, lanceolate-triangular central part, (7)9-10 on each side; inner bracts 2 times as long as outer and middle ones, white in upper part, at tip, irregularly bristly toothed, lacking cusp. Florets golden-brown- ish-yellow; corollas of central florets 10-12 mm long, peripheral ones 15-17 mm long. Achenes ellipsoid-obovate or ellipsoid, 3.5—4 mm long, 2—2.5 mm wide, sparsely hairy in upper part, sparingly through- out (var. dasycarpa Nevski); pappus (6)7-9 mm long, inner ones 1/4 as long as outer. Flowering June. (Plate XII, Fig. 2). 249 247 Rubbly-stony, clayey slopes of foothills, loess mounds, rarely old fields; in low mountains up to 1,400 m.—Soviet Central Asia: moun- tainous Turkmenia, Pamiro-Alai (southwestern part). Endemic but possibly in Afghanistan and northeastern Iran. Described from Baldzhuan. Type and isotypes in Leningrad. Note. A species very closely related to Z. glabricaulis Czer. (south- eastern and probably central Iran) and Z. crinita Boiss. s. str. (south- western and western Iran). The three species constitute the natural phylogenetic series Crinitae Czer. (Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, X1X (1959) 446. Section 2. Subulatae Czer. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX (1959) 454.—Capitula quite small; appendages of outer and middle involucral bracts long-awned, terminal awn much longer than lateral cilia; inner involucral bracts long-awned in upper part; florets white or flesh-colored, small (corollas of central florets 5 mm long, peripheral ones 7 mm, tubular); achenes glabrous; inner pappus setose, of single row of subulate, smooth, white, very narrow bristles. A monotypic section. 2. Z. purpurea Fresen. in Muz. Senckenberg I (1834) 86; Boiss. Fl. or. III, 698; Hook. f. Fl. Brit. Ind. III, 384; Post, Fl. Syr. Palest. a Sin. 461, ed. 2, II (1938) 118; Parsa, Fl. Iran. III, 699; Czer. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX, 455; in Fl. Turkm. VI, 250.—Z. aristata DC. Prodr. VI (1837) 562.—Iec.: Fresen. loc. cit. Taf. V; Czer. in Fl. Turkm. VII, Plate XX XIX, fig. 2.—Exs.: Bornm. It. Pers.-turc. 1892-1893, No. 4052. Annual. Stem 10-40 cm high, erect, finely ribbed, with few branches from middle or above, rather densely hairy below with more or less long, somewhat intertwined, whitish, articulate hairs, scabrous above from short, stiff, mostly numerous hairs, with few upward spread- ing, rather well-formed branches, usually simple. Leaves acuminate or acute, terminating into very short cartilaginous cusp, punctate, with sessile glands, scabrous from tiny, stiff hairs, sometimes mixed with arachnoid-hairy pubescence; lower ones rarely and middle cauline leaves lyrate or lyrately pinnately incised, short-petiolate, their lateral seg- ments linear, less often lanceolate or oblong-obovate, entire, 2—3(5) on each side, terminal segment 2-3 times as large as laterals; middle and upper cauline leaves (often also lower ones) narrowly lanceolate, entire, sub-sessile, middle ones 1.5--4 cm long, 0.15—0.3(0.4) cm wide. Capitula clustered in lax paniculate or almost corymbose inflorescence. Involucre ovate, almost 10 mm long (excluding slightly purple awns of 250 248 inner involucral bracts), 6—7(8) mm in dia, glabrous or farinose; outer and middle involucral bracts with membranous, pectinate appendages; cilia of appendages whitish, occasionally slightly purple, usually not spreading, stiff, hair-like, 3-4 times as long as diameter [width] of brownish or purple, lanceolately triangular central field, (5)7—10 on each side; inner involucral bracts longer, mostly purple in upper part, long-awned at tip. Florets white or flesh-colored; corolla of central florets 5 mm long, of peripheral ones 7 mm long. Achenes ellipsoid, 2.0-2.2 mm long, 0.9-1.0 mm wide, glabrous; pappus 5 mm long, inner 1/3—1/4 as long as outer. Flowering second half of March to April. Rocks, in lower mountain areas.—Soviet Central Asia: Pamiro- Alai (Kugitang, vicinity of Khodzha-Karaul), mountainous Turkmenia (Firyuza). General distribution: Southwestern Asia (Sinai Peninsula, southern Palestine, Iran, and Afghanistan). Described from southwest- ern Asia (probably from Rocky Arabia [Arabia Petraea]. Type un- known. GENUS 1603. Russowia Winkl." ? Winkl. in Tr. Peterb. Bot. Sada, XI, 2 (1892) 281; Czer. in Fl. Turkm. VII (1960) 252.—Plagiobasis auct. non Schrenk: Boiss. FI. or. III (1875) 614. Capitula homogamous, quite small, with few florets, clustered in lax paniculate or almost corymbose inflorescence. Involucres cylindri- cal, 3.5—5 mm wide, glabrous; involucral bracts 4—5-rowed, loosely imbricate, semi-coriaceous, with 3—4 dark reddish-reddish brown veins, greenish (somewhat yellowing during drying), on margin thinner, white, from broadly ovate and roundish-elliptical obtuse outer ones to nar- rowly lanceolate, acute, innermost lacking appendage. Receptacle cov- ered with scales, with few, very readily deciduous long bristles. All florets tubular, pale lilac-pink, slightly surpassing involucre, similar, bisexual; corolla tube pilose above. Anthers with small basal append- ages, apical appendages obtuse, filament flattened, glabrous. Style branches very short-woolly, with collar of hairs at base, free up to collar, oblong, included in anther tube. All achenes similar, oblong, 3.5—4 mm long, truncate above, with finely toothed crown, slightly compressed, smooth, sericeous, hilum lateral; pappus longer than achene, white, persistent, simple, comprising more or less numerous serrate-scabrous bristles, gradually becoming longer from very short 'Treatment by S.K. Czerepanov. "Named for E. Russov, famous biologist and professor at the Yuriev University. 249 251 Plate XII. 1—Russowia sogdiana (Bge.) B. Fedtsch.; 2—Zoegea baldshuanica Wink. 252 250 outer to inner ones. Annual herbs with erect, well-formed, branched stem and glabrous, pinnately incised, lyrately pinnate leaves, some- times lower ones undivided. A monotypic genus. Winkler erroneously attributed to this genus characters not found in it, such as a glabrous receptacle and a double pappus on the achenes. 1. R. sogdiana (Bge.) B. Fedtsch. in O. and B. Fedtsch. Perech. Rast. Turk. IV (1911) 267; B. Fedtsch. Rast. Turk. 756; Czer. in FI. Turkm. VII, 252.—Plagiobasis sogdiana Bge. in Mém. Acad. Sc. Pétersb. div. etr. VII (1854) 361; Boiss. Fl. or. III, 614.—Russowia crupinoides Winkl. in Tr. Peterb. Bot. Sada, XI, 2 (1892) 282.—Ie.: Winkl. loc. cit. tab. I, II. Annual. Whole plant glabrous. Stem 15—70 cm high, erect, finely ribbed, branched from middle or above, rarely almost from base, with slender, upward-spreading branches, leafless below capitula. Leaves pinnately incised or lyrately pinnate, sessile, not decurrent; lower leaves often with remote small teeth, or even entire, narrowly lanceolate to oblong-oblanceolate, subobtuse or roundish, petiolate, lateral segments of leaves narrowly linear, lanceolately linear, sometimes oblong-linear or oblong (var. latifolia Winkl.), roundish at tip, entire, terminal seg- ment usually larger than laterals, mostly remotely small-toothed; seg- ments of strongly reduced upper cauline leaves thickened at tips. Capitula clustered in lax paniculate or almost corymbose inflorescence. Involucre cylindrical, 9-13 mm long, 3.5—5 mm in dia; involucral bracts lacking appendages, outer and middle bracts obtuse, inner ones acute. Florets pale lilac-pink. Achenes 3.5—4 mm long, about 1 mm wide; pappus 6—7 mm long. Flowering April to May. (Plate XII, Fig. 1). Stony Screes and limestones, pistachio plantations, loessic mounds.—Soviet Central Asia: Mountainous Turkmenia (Firyuza, Kushka, Kazachye Ravine), Amu-Darya (Charshanga), Pamiro-Alai (western part), Syr-Darya (south), Kyzvyl-Kum (Zhana-Darya River), Tien Shan (former Dzharkent District, Kairylgan Road). Endemic. Described from Zhana-Darya River (Yana-Darya). Type in Leningrad. GENUS 1604. Crupina Cass.'? Cass. in Dict. Sc. nat. XII (1818) 67, XLIV (1826) 35; L. (1827) S29 Xe (SSO) a7 te 'Treatment by S.K. Czerepanov. "Name of unknown origin, presumably from the Low German word krupen (kreichen)—to crawl; named, presumably, for the ability of the achenes to disperse with the help of their pappus. 25 Ww 251 Capitula heterogamous, medium-sized, with few florets clustered in corymbose or paniculate-corymbose, inflorescence. Involucres cylindrical, (4)5-8 mm in dia; involucral bracts few-rowed, loosely imbricate, herbaceous, membranous on margin and almost transparent, with distinct longitudinal striations and numerous, punctate, sessile glands on outer side, greenish, slightly violet at tip, from lanceolate- ovate and acute outer to oblong-lanceolate, long-acuminate innermost lacking appendages. Receptacle covered with long, relatively readily deciduous bristles. All florets tubular, purple, somewhat surpassing involucre, heterogeneous; central florets (disk florets) bisexual, with hairy corolla tube; peripheral florets (ray florets) sterile, lacking staminodes and rudiments of style, with subglabrous (corolla tube with occasional hairs) 5-parted corolla, not broadened. Anthers with short basal appendages, their apical appendages more or less acute; fila- ments flattened, papillose. Style very short-woolly above with collar of hairs, short, bifid, strongly exserted from anther tube. All achenes similar, 1-5(6) in each capitulum 4—5 mm long, truncate above with obtuse or indistinctly toothed crown, smooth, sericeous, hilum basal or lateral; pappus 1.5—2 times as long as achene, persistent, double, outer pappus comprising several rows of serrate-scabrous, stiff bristles, be- coming longer from very short outer to inner ones; inner pappus much shorter than outer, comprising 5—10 triangularly lanceolate or oblong, entire, small scales in one row. Annual herbs, with erect stem and pinnately incised, sessile leaves, nondecurrent. Type of genus: C. vulgaris Cass. The genus includes three or four species, of which two are with us. 1. Achenes obovate, compressed at base, with lateral linear hilum; pappus whitish-rusty-brown or pale rusty-browN.............ccceeseeeeeees BF tO eee ee 1. C. crupinastrum (Moris) Vis. + Achenes short cylindrical, truncate, not compressed, with circular hilum at base; pappus blackish-brown................. 2. C. vulgaris Cass. 1. C. crupinastrum (Moris) Vis. Fl. Dalm. II (1847) 42; Boiss. Fl. or. III, 699; Grossh. Fl. Kavk. IV, 193 and Opred. Rast. Kavk. 488; Ter-Khachaturova in Fl. Gruzii, VIII, 524; Agadj. in Fl. Azerb. VIII, 421.—C. vulgaris auct. p. min. p. non Cass.: DC. Prodr. VI (1837) 565; Ldb. Fl. Ross. II, 2, 684.—C. morisii Bor. Fl. Cent. Fr. ed. 2 (1849) 292.—C. zuccarinii Bge. ex Nym. Consp. FI. Eur. II (1879) 433, pro syn.—Centaurea crupinastrum Moris, Enum. sem. hort. Taur. (1842) 12.—Iec.: Vis. loc. cit. tab. 51, fig. 3—Exs.: Fl. cauc. exs. No. 75; Billot, Fl. Gall. et Germ. exs. No. 3424, Todaro, Fl. Sic. exs. No. 826; Pl. Herceg. exs. No. 93; Fl. Graeca exs. No. 468; Herb. Europ. sine numero. 254 252 Annual. Stem 20-80 cm high, erect, finely sulcate, branched, well- formed, whitish-pubescent below, with short-plumose, multicellular, intertwined, thin hairs, glabrous above, often with more or less sparse, dull, sessile glands. Leaves sessile, pinnately incised; leaf segments narrow, narrowly linear, toothed, with stiff, spines indistinctly stellate (visible under high magnification of stereoscopic microscope) at tip, lower leaves pubescent like stem, usually on both sides on midrib, lowermost leaves 2, very small, oblong-obovate, obtuse, entire, with- ering early. Capitula solitary at tips of stem and its branches, in lax corymbose or paniculate-corymbose inflorescence. Involucre cylindri- cal, 16—20 (22) mm long, 5—8 mm in dia; involucral bracts on outer side with distinct greenish longitudinal lines and numerous, punctate sessile glands, from lanceolate-ovate, acute outer ones to oblong-lan- ceolate, long-acuminate innermost lacking appendages. Florets purple. Achenes obovate, 4—4.5 mm long, 2.3—2.5 mm wide, plump, sericeous, at base compressed, with lateral linear hilum; pappus whitish-rusty- brown or pale rusty-brown, 8—9 mm long. Flowering second half of May to July (August). Dry clayey and rubbly slopes, among shrubs, on rocks from low- lands to mid-montane zone.—Caucasus: Dagestan, Eastern and South- ern Transcaucasia, Talysh. General distribution: Mediterranean Region, Balkans-Asia Minor, Armenia and Kurdistan, Iran-Afghani- stan. Described from Italy. 2. C. vulgaris Cass. in Dict. Sc. nat. XII (1818) 68; DC. Prodr. VI, 565, p. p.; Ldb. Fl. Ross. II, 2, 684 p. p.; Boiss. Fl. or. III, 699; Schmalh. FI. II, 114; Fedtsch. and Fler. Fl. Evrop. Ross. 1014; O. and B. Fedtsch. Perech. Rast. Turk. IV, 267; Hegi, Ill. Fl. VI, 2, 930; Grossh. Fl. Kavk. IV, 192 and Opred. Rast. Kavk. 488; Dobrocz. in Vizn. Rosl. URSR, 571; Ter.-Khachaturova in Fl. Gruzii, VIII, 523; Geideman, Opred. Rast. Mold. SSR, 291; Dostal, Klic, 730; Nikitin in Fl. Turkm. VII, 253; Agadj. in Fl. Azerb. VIII, 422.—C. pauciflora Kar. and Kir. in Bull. Soc. Nat. Mosc. XV (1842) 392, non Hoffmgg. and Link (1820).—Centaurea crupina L. Sp. pl. (1753) 909.—Serratula crupina (L.) Vill. Hist. Pl. Dauph. II (1789) 38.—Ie.: Hegi, op. cit. fig. 629; Javorka and Csapody, Iconogr. Fl. Hung. (1933) 544, fig. 3888; Dostal, op. cit. tab. 279, fig. 2493.—Exs.: Fl. exs. austro-hung. No. 3063; Fl. Gall. et Germ. exs. No. 685; Rchb. Fl. Germ. exs. No. 2524; Fl. exs. reip. Boh.-Slov. No. 881; Novopokr. Herb. Fl. Tanait. No. 67. Annual. Stem 20-80 (120) cm high, erect, finely sulcate, branched, well-formed, whitish-pubescent below with distinctly plumose, short, thin hairs, usually glabrous above. Leaves sessile, pinnately incised, their segments narrowly linear, entire or small-toothed, on margin with 25 a) 253 stiff spines indistinctly stellate (visible under high magnification of stereoscopic microscope!) at tip, lowermost leaves 2—4, very small, obovate to oblong-oblanceolate, obtuse, entire or toothed, withering early. Capitula solitary at tips of stem and branches, usually in clusters of 2—6 in corymbose or paniculate-corymbose inflorescence. Involucre cylindrical, (12)14—18(20) mm long and 4—6 (8) mm in dia; involucral bracts with distinctly dark green or green, longitudinal striations and numerous punctuate, sessile glands on outer side, from ovate or lanceolately ovate, acute outer ones to oblong-lanceolate, long-acumi- nate innermost lacking appendages. Florets purple. Achenes short-cy- lindrical, truncate, 3.5-6 mm long, 2-3 mm wide, plump, dark-brown below, yellowish-sericeous above, with round basal hilum; pappus blackish-brown, 5—8 (9) mm long. Flowering second half of April to July (August). ; Dry stony and rubbly slopes, on rocks and screes, among shrubs, on gravelly banks of rivers and streams, sometimes in gardens, vine- yards, old fields and crop fields —European Part: Bessarabia, Black Sea Region (south), Lower Don, Crimea (south); Caucasus: Almost all regions; Soviet Central Asia: mountainous Turkmenia, Pamiro-Alai, Tien Shan; Dzhungaria-Tarbagatai (Dzhungarian Alatau Range). Gen- eral distribution: southern and Central Europe, Balkans-Asia Minor, Armenia and Kurdistan, Iran-Afghanistan. Described from southern Europe. Type in London. Note. In Soviet Central Asia, there is a predominance of plants with more capitula on the average, in a more crowded inflorescence (in clusters of 2—6) at the tips of the stem and its branches, and on the average, larger involucres and achenes (1—5 per capitulum). However, in the Caucasus, in the south of the European part of the USSR and in Europe, specimens are found that do not differ from the Soviet Central Asian ones. The Soviet Central Asian plants were described as a sepa- rate species, C. pauciflora Kar. and Kir., which was found to be a later homonym (cf. synonymy). O.V. Tscherneva (F/. Uzbek., VI (1962) 387) proposed the new name—C. oligantha Tschern. GENUS 1605. Plagiobasis Schrenk’:? Schrenk in Bull. Acad. Sc. Petersb. III (1845) 108. Capitula heterogamous, rarely homogamous, medium-sized, with many-florets in paniculate, less often almost corymbose inflorescence. Involucres globose-ovate or ovate, 10—20 mm in dia, glabrous; involu- 'Treatment by S.K. Czerepanov. 2From the Greek words plagios—oblique, lateral, and basis—base; named prob- ably for the lateral hilum of achenes. 256 254 cral bracts many-rowed, compactly imbricate, coriaceous, smooth, greenish (yellowing during drying), with small dark reddish-brown, triangular or almost crescent-shaped macula at apices, from roundish- elliptical outer ones to oblong innermost, outer and middle ones with very narrow, subcoriaceous or stiff, membranous border at tip, weakly delimited from coriaceous part; inner involucral bracts with membra- nous, entire, small appendages. Receptacle bristly. All florets tubular, pinkish-violet, much surpassing involucre, mostly heterogeneous; co- rolla glabrous; central florets (disk florets) bisexual, in several rows; peripheral florets (ray florets) sterile, mostly with staminodes far ex- serted from corolla tube, and long rudiments of style, in smgle row, less numerous not broadened, sometimes bisexual (and then all florets similar). Anthers with short basal appendages, their apical appendages subacute; filaments flattened, sparsely short-papillose laterally, gla- brous below. Style branches short-woolly, at base with collar of hairs, free to 1/2—2/3 entirely or partly exserted from anther tube, oblong, divergent. All achenes similar, oblong-cylindrical, 5-6 mm long, rounded at tip, often very slightly curved, smooth, with occasional, long, readily deciduous hairs, mature achenes subglabrous or glabrous, hilum lateral; pappus almost as long as achene, whitish, simple, of few, very readily deciduous (at base not connate into ring), serrate bristles, gradually becoming longer from outer to inner ones. Perennial herbs with erect, branched stem and entire, thick, glabrous, leaves. A monotypic genus. 1. P. centauroides Schrenk in Bull. Acad. Sc. Pétersb. III (1845) 109; B. Fedtsch. Rost. Turk. 756; Popov Fl. Almat. Zapovedn. 48.— P. dshungaricus Ijin in Izv. Bot. Sada, Akad. Nauk SSSR, XXX, 3-4 (1932) 357.—Exs.: GRF No. 4299. Perennial. Whole plant glabrous, with cylindrical, vertical, simple or less branched root, thickened at collar. Stem solitary, occasionally several (2-3), 25—60 (80) cm high, erect, finely ribbed, branched from middle or above, with upward spreading branches, mostly leafy up to tip. Leaves thick, obovate to oblong, obtuse, subobtuse, less often subacute, usually irregularly toothed; basal and lower cauline leaves strongly approximate, petiolate, petioles 1.5—6.0 cm long; others sessile, sometimes very slightly amplexicaul; apical leaves greatly shortened. Capitula in paniculate, less often almost corymbose inflorescence. Involucre globose-ovate or ovate, 15S—20 mm long, 10-20 mm in dia; outer and middle involucral bracts with very narrow, semicoriaceous or stiff, membranous border at tip, weakly delimited from coriaceous part; inner bracts with small membranous appendages. Florets pinkish- purple. Achenes 5—6 mm long, 1.2—1.5 mm wide; pappus 5 mm long. Flowering July to September. (Plate XIII, Fig. 1). Poy AN I Slip fey | ay et Plate XIII. 1—Plagiobasis centauroides Schrenk; 2—Oligochaeta minima (Boiss.) Briq. 258 256 Outcrops of gypsic clays, conglomerates, and red sand-stones, rubbly slopes; in desert, foothills and low-mountains—Soviet Central ‘Asia: Dzhungaria-Tarbagatai (southern spurs of Dzhungarian Alatau; Dzhungarian Pass), Tien Shan (Ketmen Range, eastern spurs of Kungei Alatau and Trans-Ili Alatau ranges, Issyk-Kul Depression), Pamiro- Alai (northeastern foothills of Alai Range). General distribution: Dzhungaria-Kashgarai (Kuldzha, Kash River. Turfan). Described from Chingilda in Dzhungarian Alatau. Type and isotypes in Leningrad. Note. M.M. Iljin (op. cit.) described P. dshungaricus Ijin from the vicinity of Dzharkent (former Panfilov). I could not find characters reliably distinguishing it from P. centauroides Schrenk. It is repre- sented only by the type specimen, collected in the range of the latter species. GENUS 1606. Tricholepis DC.' 2 DC. in Guill. Arciv. bot. II (1833) 331, 515; DC. Prodr. VI (1837) 563; Benth. and Hook. f. Gen. pl. II (1873) 475. Capitula with similar, regular, bisexual, fertile florets. Involucre, many-rowed, of imbricate, narrow, more or less long-awned bracts, gradually reduced from inner to outer ones. Receptacle flat, alveolate, densely bristly with long white bristles. Corolla regular, with narrow tube and narrow limb 5-fid. Filaments densely ciliate-hairy; anthers sagittate at base with rather long, lacerate, incised appendages. Style with thin long branches. Achenes attached laterally, along edge of slightly cartilaginous hilum, roundish-obpymramidal, somewhat angu- lar, glabrous, weakly striped, truncate above, with broad (almost as wide as achene) areola at apex with projecting, small-toothed, upper margin; pappus many-rowed, much longer than achenes, consisting of numerous, sordid-white, very short-plumose unequal hairs, at base con- nate in ring and detaching together, innermost longer and at base some- what thickened. Perennials (in Pamiro-Alai, Hindukush and Himalayas) or annuals (mostly in central and southern parts of the Indian Penin- sula), with virgate, somewhat woody, more or less long branches, and pinnately incised or remotely-toothed lower leaves, and entire or subentire, middle and upper leaves greatly reduced in upper part; capitula lacking bracteal leaves, always solitary at apices of stems. The genus comprises about 15 species; one species’ is known in the USSR. 'Treatment by I.A. Linczevsky. "From the Greek words trix—hairs, and /epis—scales. 259 257 1. T. trichocephala Lincz. in Bot. Mat. Gerb. Bot. Inst. Akad. ‘Nauk SSSR, XVI (1954) 479.—Ic.: Lincezevsky op. cit. Plate 1, Fig. a-c. Perennial. Semishrubs with thick woody root and numerous cau- dices. Stems 40-60 cm high, numerous (20-30), weakly floccose-white tomentose below, entirely glabrous above, more or less strongly branched only below with slender, virgate, somewhat woody, long (to 30-50 cm), quite densely leafy branches. Leaves sessile, lower ones linear-lobate, uniformly remotely toothed, rather fleshy, sparsely pu- bescent with short crisped hairs, 25-40 mm long, 3—4 mm wide; middle and upper leaves linear and narrowly-linear, erect, entire or basally with 1—2 small teeth, very fleshy, glabrous, with prominent white- cartilaginous cusp, (2)5—15(25) mm long, (0.5)1.0—1.5(2.0)mm wide, abruptly reduced upward. Capitula solitary at tips of branches, oblong- ovate or conical, or subconical, with fruits broadened above, and then cylindrical or obconical, 20-25 mm long (with florets), at base round or almost truncate, 7-10 mm in dia. Involucre stramineous, weakly floccose-white-tomentose; involucral bracts quite numerous; middle bracts stiff below, strongly appressed and firmly imbricate, linear or oblong-linear, obtusely carinate, 4-12 mm long, about 1 mm wide, almost abruptly attenuated above into thin, hair-like, strongly (some- times horizontal) spreading, almost straight, light pink awn 8-12 mm long; outermost bracts much smaller, almost subulate, innermost lin- ear, 18-20 mm long and about 1 mm wide, gradually narrowed above, hairy and ciliate at tip. Receptacular bristles almost 2 times as long as mature achenes. Florets slightly surpassing capitulum; corolla pink (?), 18-20 mm long, limb about 10 mm long, its lobes narrow, linear, about 5-6 mm long; anther appendages about 1.0-1.5 mm long, irregularly lacerate-incised with unequal lobes. Mature achenes about 5 mm long, 2 mm in wide; pappus almost 3 times as long as mature achenes. Flowering ?; fruiting September to October. Stony slopes in lower mountain zone at about 1,200—1,400(1,800) m.—Soviet Central Asia: Pamiro-Alai (southern Darvaz Region). Gen- eral distribution: Very possibly found in northeastern Afghanistan. Described from basin of Zarbuz River—a tributary of Pyandzh River. Type in Leningrad. Note. The description of the lower and middle leaves (which are absent in the type specimens collected later) is given here on the basis of the material originating also from the southern Darvaz Region (south- western spurs of the Darvaz Range, southeast of the village of Sarygor, 1.VI. 1961, No. 286, Yu. Soskov and S. Yunusov), collected not far from the classic locality. 260 258 GENUS 1607. Serratula L.! 2 L. Sp. pl. (1753) 816. Capitula homogamous, rarely heterogamous, with many-florets, medium-sized or small, in lax corymbs or solitary, large. Involucres semiglobose, globose, ovate or almost cylindrical, comprising several (4-9) rows of firmly imbricate, numerous, more or less coriaceous bracts; outer bracts mostly ovate, short, lacking appendages, subobtuse, acute, or cuspidate, more or less scarious on margin; inner bracts lan- ceolate and linear with membranous, obtuse or subacute apical ap- pendage. Receptacle flat, bristly, densely pilose. Florets all tubular, pink, purple, lilac-purple or white, in upper part cup-shaped, almost regularly 5-parted, bisexual, less often mono- and dioecieous or pe- ripheral florets pistillate, surpassing bisexual central florets. Stamen filaments glabrous, smooth or with very fine papillae; anthers at base sagittate, with undivided, mostly linear appendages. Style usually bi- fid, less often undivided; lobes usually longer, divergent at maturity, arcuate. Achenes 4—6 mm long, oblong, terete, somewhat compressed laterally, glabrous, smooth or finely sulcate, with oblique hilum; pap- pus sordid-white or rusty-brown, many-rowed, simple, comprising ser- rate, less often plumose, persistent or gradually detaching bristles, becoming longer from outer to inner. Perennial, unarmed, glabrous or scatteredly hispid herbs, with erect, sometimes ascending stems or almost stemless with alternate, mostly stiff, sometimes coriaceous, pinnately incised, less often undivided, toothed or entire leaves; leaves scabrous on both sides or only beneath, or glabrous, less often with weakly crisped hairs beneath. Type of genus : S. tinctoria L. The genus comprises about 50 species distributed in Asia, Europe and North America. Economic Importance. Some species are used as dyeing agents to obtain yellow and green dyes (S. tinctoria and others); others are medicinal, vitamin-containing, meliferous and fodder plants (in the early stages of development). The stems of S. coronata contain traces of alkaloids. 1. Leaves compound, with 2-4 oblong or elliptical lateral leaflets and larger terminal leaflet (Caucasus) .............. 28. S. quinquefolia MB. + Leaves simple, undivided, pinnately lobed or incised .................. 2. 2. Stems divaricately branched from base, densely pubescent, basal 'Treatment by A.G. Borissova. "From the Latin word serra [saw]—projection of the saw-toothed leaf margin. 261 ea 259 leaves pinnately divided or incised. Capitula 0.4—1.5 cm in dia, mostly oblong, numerous, clustered in dense or lax corymbose or corymbose-paniculate inflorescence; pappus bristles few, decidu- Stems simple or branched in upper and middle part, sometimes plants stemless or almost stemless. Capitula large- or medium-sized, globose to ovate, few, solitary or in corymbose inflorescence Plant pubescent with articulate hairs; leaves pinnately divided ih lobes lacking cusp. Involucres glabrous ...............:eeceeseesceseeeceeeeeeeeeseeees teers h salte la clesnnacyides cu agitate diets erates tater a 3. S. erucifolia (L.) Boriss. Plants pubescent with simple stiff hairs; leaves pinnately incised with toothed lobes, terminating into cartilaginous cusp. Involucres pubenulentiansube labnou sisi aeee tes et dct eta a en wtesisanenes 4. Involucral bracts puberulent, with curved, subulate cusp 2-3 mm long; involucres cylindrical. Stony places and alkaline steppes lee leeoes ny janbiu tees. opel genesmed cancels leat by 4.S. dissata Ldb. Involucral bracts subglabrous, with straight, hard, 3—5-mm-long apical spine; involucres cuneately narrowed toward base, narrowly cylindrical. On sands ...............:00 5. S. angulata Kar. and Kir. Stems reduced, 2—5 mm high, or plants stemless or almost stemless, leaves lyrate or runcinate. Capitula sessile or subsessile; involucral bracts 4=-6-rowedwasdee.in venialaden eyeel achededasete ns Ls! 6. Stems well developed, 20-190 cm high; if shorter, then leaves coriaceous, undivided or only at base pinnately lobed. Capitula pedunculate, sometimes subsessile; involucral bracts 5—8-rowed Leaves densely spinose-serrate, lyrate, with 2—3 pairs of Yiteral lobes; terminal lobe much larger than laterals. Involucres 5S—6-rowed cata teas saa i ag i ell 29. S. lyratifolia Schrenk Leaves with occasional inconspicuous teeth, runcinate, pinnately incised into 3-5 pairs of irregular lobes, terminal lobe differing little from laterals. Involucres 4-roWed .00..........ccccceecececeeeneeeeeeeteees want ett PS, sch hat teeter 30. S. tianschanica Sap. and Nik. Stems sometimes leafless, often woody at base; leaves coriaceous, crowded mainly in lower part of plant (Soviet Central Asia) ..... By Stem leafy throughout or almost throughout ................cccceceeeeeees 8. Peripheral florets pistillate with large corolla, others bisexual; ca- pitula rather large, usually several in one cluster; involucre with rusty-brown tomentum, reddish-brown to reddish. Leaves large, deeply pinnately incised, coarsely serrately toothed; plants 80-150 rere ay) ofc Weer eee een a, Sere Ler eee 2. S. coronata L. Peripheral florets bisexual or unisexual (dioecious plants). Leaves 262 260 tel. 12% 13% 15. entire, toothed, serrulate, or pinnately divided, sometimes pinnately TMCISe chia. 340, ges, 04. TT, RS RIS, SETERUOISY oo: 2. Capitula mostly several in corymbose inflorescence, often constricted at base, 1.0-1.5 cm in dia. Plants dioecious, sometimes monoe- cious; leaves usually undivided or pinnately incised, serrulate (European partoff USSR) 252.00. as. ee eee 1. S. inermis Gilib. Capitula solitary, 1.5—4.0 cm in dia. Plants monoecious. Leaves undivided or pinnately lobed, sinuate-toothed or entire .............. 10. Capitula 2.5—4 cm in dia; involucral bracts with long, straight, re- curved, hard spines. Plants low, stems simple, with single terminal CAP er luMAK CAUCASUS )o avsee-2sbenceanmensstandiesteaongcaseceees-eacadonet eee 11. Capitula smaller. Stems simple, with single terminal capitulum, but then capitula 1—1.5 cm in dia, or branched and with several capitula vinatadanebaewntliraunnndacesrontnuacaenceraness ste: CLOT eae EY Sooo eR eERREE SS re 2k Capitula semiglobose. Leaves lobate at base, crenate-toothed ........... HUET: a8, RR eS, 32. S. serratuloides (Fisch. and Mey.) Takht. Capitula subglobose. Leaves large, undivided, irregularly spinose- GELLALS AY IEE. «FSI ILS. ONY. ue EE a, Ne 33. S. caucasica Boiss. Leaves large, deeply pinnately incised or divided, sometimes only lower ones or only reduced upper cauline leaves undivided; plants scabrous or glabrous, (60) 75—150 cm high ...........eeeeeeeeeeeeeeees ie Leaves undivided, sometimes lower ones pinnately lobed or sinu- ate-toothed; plants subglabrous or pubescent but not scabrous, 10— SOW Senmhigh ace Wit, CO Ae | Seseel ey Dice Yreiis..s 23. Stems more than 25 cm high, more or less branched, angular, erect, uniformly leafy mainly in lower part; leaves coarsely spinose-toothed, SOMME be Sale Ged ASE cs rate sre ce apeeeet aaceenauasuares secrertysoesenceeaeses tae 14. Stems mostly low, 5—20 cm high, simple, erect or ascending; leaves sharply-serrate or bluntly-toothed to crenate mostly in basal part of Je CIO i cdi a a RC EO epee ULE IR ae Mle, 1S) Capitula oblong, 0.7—1.2 cm in dia, at tips of lateral branches; involucral bracts greenish-stramineous, monochromatic. Stems branched above (Lake Balkhash Region, Dzhungarian Alatau, Tien Shain) Sete i ome ea I 21. S. suffruticosa Schrenk Capitula about 1.5 cm in dia, solitary on 15-cm-long peduncles; involucral bracts with dark brown spot at tip. Stems with few branches in lower part (Pamiro-Alai) ........... 19. S. chartacea Winkl. Peduncles much longer than leaves; leaves obovate, crenately toothed bluntly toothed or bluntly-lobed. Capitula about 1 cm in dia SCHOLL. SEES, UL, OSE ERE es, CUE 22. S. aphyllopoda Ijin Peduncles more or less leafy, as long as or slightly longer than Reaivie S 2c fis Perea ican iv vecacamanneatenesususnodulmtencnes SRO Pete anes 16. Capitula (2.5) 3.0-4.0 cm long and to 3 cm in dia, flat at base. 263 17. 18. We) 20. 21. 261 Leaves large, ovate or oblong, undivided or with several lobes at base, long-petiolate (Dzhungarian Alatau) ....... 31. S. modesti Boriss. Capitula 1.0-2.5 cm long, 1.0-1.5 cm in dia, roundish at base. Stems ascending, short, more or less leafy; leaves narrowly lan- ceolate, sharply serrate, short-petiolate ...... 20. S. procumbens Rgl. Basal and lower cauline leaves ovate or oval, undivided or incised at base itself, sinuate-coarsely toothed; cauline leaves pinnately in- cised or lobed. Capitula on long, straight peduncles, leafless above mmnicldlllcspetyebey cyt tetbep ey, myers eau: wh, 12. S. heterophylla (L.) Desf. All leaves similar, pinnately incised with entire or pectinate seg- IMIGHS ebecclern atueke, suaesnclteet el se. ween cut thet tet Ahem shee Do 18. Stems leafy to or almost to capitula; peduncles short; plants sca- brous-pubescent or subglabrous. ..........0..0cccccccecssesseseseseseeeeeeeee 19. Stems tall, leafless in upper part; leaves large, basal leaves long- petiolate. Capitula large, on peduncles 20-50 cm long .............. 2). Plants scabrous-pubescent, grayish, densely leafy; all leaves simi- lar, pectinate, pinnately incised into similar or almost similar, lan- ceolate or linearly-lanceolate, coarsely toothed lobes. Capitula to 2 cm in dia and to 3 cm long; outer involucral bracts with very short spines, inner with spiny-toothed appendages (Trans-Baikal) ...... Mirna: etna Tener censors Mine, chal 8 oe we septa cos ke: 6. S. centauroides L. Plants almost glabrous or weakly pubescent; leaves lyrate or pin- nately incised, [all] similar or lower undivided to pinnately lobed, uppermost leaves sometimes linear or linear-lanceolate; leaf lobes entire, less often with occasional teeth, sometimes terminal lobe longer than laterals. Involucral 0.7—1.5 cm in dia, 1.0—-1.5 cm long; involucral bracts with conspicuous, very thin, filiform spines ....... Involucral bracts with filiform cusp nearly 0.5 mm long. Upper leaves sometimes undivided, linear (south of European part of USSR) ae eae Ch lite Ud dh alte 8. S. radiata (Waldst. and Kit.) MB. Involucral bracts with recurved and erect cusp 1.5-3.0 mm long. Capitula constricted in upper part. All leaves lyrately pinnatifid, sometimes terminal lobe longer than laterals (Armenia) ................ i PRG tN 2A libres, | 9. S. biebersteiniana (Iljin ex Grossh.) Takht. Plant entirely glabrous, with slender stems; all leaves similar in shape; leaf lobes entire, linear to filiform in upper leaves, erect almost at right angle to rachis. Florets pale purple or whitish. On CHa RG A tro detec, PY ccc sesenss 11. S. tanaitica P. Smimn. Plants scabrous-pubescent, mainly in lower part, or subglabrous, 60-100 cm high, with thick, sulcate stems; leaves large, pinnately incised or divided; upper cauline leaves pinnately lobed or undi- vided, lanceolate, with broad lobes .0...0...0......c:cccecccessceccenceessees 22. 264 265 262 23. 24. WS). 26. 2a 28. 29. Involucral bracts yellowish-green, glabrous, 2.0—2.5 cm wide. Plants subglabrous, glaucous, 60-80 (100) cm high (Volga Region, South- em Urals, Western Siberia) .:.......0.0...000. 10. S. gmelinii Tausch. Involucral bracts with dark tips, sometimes with arachnoid-hairy pubescence. Plants scabrous-pubescent, 80-150 cm high, with sturdy, often violet-colored stem (Trans-Baikal) ....... 7.8. komarovii I]jin Leaves coriaceous or almost coriaceous, glabrous, short-petiolate, entire, erose-toothed to lyrate with remote lobes in lower part of leaf, crowded mostly in lower part of plant; upper leaves some- times pinnately incised with few linear lobes, sessile ................. 24. Leaves soft, not coriaceous, glabrous or short-ciliate, basal leaves long-petiolate, mostly undivided, entire, or upper leaves with large teethiorlobes- ie REO RO eons 28. Capitula oblong or ovate-oblong with cuneate base; involucral bracts yellowish, inner ones violet, lustrous, lacking cusp or with very short cusp often deciduous (Caucasus) 2.0.00... eeeeeeecesseeeeeeseeeees LGR. A QUE. 2 UA AR 27. S. coriacea Fisch. and Mey. Capitula ovate or oblong-ovate; involucral bracts with distinct cusp (Soviet:Central Asia)Gxi7. 2RU8e ARN I. 2 AE eens 2 Leaves undivided or entire, basal ones elliptical, gradually nar- rowed into petiole, obtuse or with short cusp; cauline leaves sessile, ovate-oblong to ovate, upper ones semiamplexicaul, auriculate. Ca- pitula numerous; involucres cupuliform, 1.2—1.5 cm in dia, yellow- ish-green (Talass Alatau) ............... 26. S. hastifolia Kult. and Korov. Plantstotiotherthabit:) 22) 2028, TE ROM, 26. Basal and middle cauline leaves undivided, less often sinuate-toothed in lower part, lanceolate and linear-lanceolate, upper ones entire; involucres about 0.8 cm in dia, oblong-ovate ..............scseeseeesseeeeees Basal and middle cauline leaves undivided, large-toothed or lyrate- lobate: withiremote:lobes. Ortecth nccciwsenen nnn cwunennnne’s Dar Involucral bracts with straight spines 0.5—1.5 mm long. Plants 25— 55 cm high (mostly in Pamiro-Alai) ............ 23. S. sogdiana Bge. Involucral bracts with divergent spines 3-5 mm long. Plants 30-80 cm high (mostly in Tien Shan) ................... 24. S. alatavica C.A.M. Stems branched above; leaves long-petiolate, mostly entire, some- times with several small teeth; capitula usually in clusters of a few (1-5), 0.8—1.0 (1.5) cm in dia; involucral bracts green, with small brownish spot at tip ..............00 13. S. cardunculus (Pall.) Schischk. Stems simple; capitula usually solitary, 1.5—2.5 cm in dia; involu- cral bracts green or blackish, with blackish brownish border ..... 29. Involucral bracts blackish or brownish; capitula 1.5—2.5 cm in dia weet Dcausessvenvereevaveree RRR, EM LE ERO RO Let 30. 266 263 30. _Involucral bracts usually black or with brownish-black border; leaves oblong, long-petiolate, usually entire or with remote, rare broad feethioy Mamas Pot, Binley 3 15. S. algida Iljin + Involucral bracts brown on margin and at tip. Leaves glaucous, cauline leaves usually toothed or pinnately lobed .................... arte tree eres. veal auecses,. Rr laye: 14. S. marginata Tausch. 31. Stems and basal leaves on surface and on margin glabrous; basal leaves undivided or subentire, rarely large-toothed, with very short cartilaginous cusp; involucral bracts with hard cusp ............:0000 eh See meer evel, Torre Leave eomaieis wh ooey . ! 16. S. kirghisorum Iljin + Stems pubescent; leaves ciliate on margin; basal leaves large-toothed or lobed; involucral bracts with thin soft cusp .............c:ccceeeees B22. 32. Involucral bracts densely tomentose, narrowly dark-bordered and with dark veins at apex; capitula 2—2.5 cm in dia, roundish at base; achenes oblong; pappus 10-15 mm long. All leaves large-toothed; cauline leaves narrowly lanceolate, appressed to stem ..................- kts ened} otal ine sates reel od ey. i, 17. S. dshungarica I]jin + Involucral bracts glabrous or young bracts sparsely pubescent, green; capitula 1-2 cm in dia, concave at base; achenes oblong-ovate; pappus 9 mm long. Cauline leaves oblong-lanceolate, spreading Ee IM LEST, TELE 18. S. transcaucasica (Bornm.) Sosn. ex Grossh. Section 1. Serratula.—Sect. Sarreta DC. Prodr. VI (1837) 667.— Plants dioecious or monoecinous; capitula homogamous, oblong or oblong-ovate, small, several, usually clustered in corymbose or corym- bose-paniculate inflorescence; florets unisexual or bisexual. Type of section: type of genus. 1. S. inermis Gilib. Fl. Lithuan. III (1781) 183; Exerc. phytol. 1, 152; Iljin in Fl. Yugo-Vost. VI, 412; Dobrocz. in Vizn. Rosl. URSR, 570; Majevski, Fl. ed. 8, 607.—S. tinctoria auct. fl. ross. non. L.— S. tinctoria ssp. inermis var. angustiloba M. Pop. Ocherk. Rast. i. FI. Karpat. (1949) 252, nomen.—Ie.: Fedtsch. and Fler. Fl. Evrop. Ross. Fig. 1018; Syreistsch. Ill. Fl. Mosk. Gub. III, 308, Kozlovskaja in FI. BSSR, V, Plate L.—Exs.: GRF No. 925. Perennial. Roots numerous, string-like; rhizome more or less ob- lique, short. Plant 30-120 cm high; stems erect, glabrous, hard, simple or branched above, with fibrous remnants of old leaves at base. Basal leaves long-petiolate, ovate or oblong-ovate, to 15 cm long and 8 cm wide, acute, at base more or less roundish, sharply serrulate, often undivided or with several lateral lobes, green, sometimes reddish, light 268 264 green beneath, stiff and scabrous from short papilliform hairs; middle cauline leaves short-petiolate to subsessile, elliptical to oblong-lan- ceolate, undivided or lyrate with large terminal lobe, acuminate; upper leaves reduced, often undivided. Capitula mostly several in clusters, at stem apices in corymbose-paniculate or corymbose inflorescence, small, 1.0-1.5 cm long, oblong or ovate, narrowed toward base and slightly constricted. Involucres oblong-cylindrical; involucral bracts dark purple entirely or only in upper part; outer and middle bracts broadly ovate to oblong-lanceolate, on margin and arachnoid-hairy or ciliate-arachnoid-hairy, in upper part, curved and acuminate; inner bracts longer than rest, lanceolately linear, acuminate. Florets lilac- purple, sometimes white, mostly unisexual and dioecious or bisexual. Corollas of pistillate florets about 13 mm long, narrow part of corolla [corolla tube] about 7 mm long, broad part [limb] about 5 mm long, with linear, acute lobes about 3 mm long and 0.5 mm wide; style as long as corolla or slightly exserted from it, about 13 mm long, brown- ish, with divergent, linear lobes about 1.5 mm long; staminate florets with well developed anther tube, as long as corolla or somewhat longer, ovary abortive, style short, with long linear branches [lobes]; bisexual florets with style sometimes piercing anther tube and surpassing it. Achenes oblong, 4—6 mm long, grayish or brownish; pappus about 7 mm long, comprising dense, weakly scabrous, short-toothed hairs, yellowish-brownish. Flowering June to September; fruiting August to September. Open broad-leaved and mixed forests, mostly oak, pine, and birch, forest edges, among shrubs, in cut-over forests, moist, sometimes al- kaline meadows, on sand ridges, limestones and chalk outcrops.—Eu- ropean Part: Baltic Region, Upper Volga, Volga-Kama, Upper Dnieper Middle Dnieper, Volga-Don, Trans- Volga, Upper Dniester, Bessarabia, Black Sea Region, Lower Don. General distribution: Central Europe (Poland). Described from vicinity of Grodno. Type in Kiev. Note. This species is similar to the European S. tinctoria L. and possibly even identical to it; the character of absence of spines on the involucral bracts is variable. Additional observations in nature are necessary. Within the USSR, we often come across plants with undi- vided leaves—f. integrifolia Ijin (op. cit. 413). In Western Europe, the form S. tinctoria L. with incised leaves is predominant; however there are individuals with undivided leaves that are described as a separate species, for example, S. germanica Wiesb. Section 2. Mastrucium (Cass.) DC. Prodr. VI (1837) 667.— Mastrucium Cass. in Dict. Sc. nat. XXXV (1825) 173, XLI (1826) 521.—Pereuphora Hoffmgg. Verz. Pfl. Nachr. II (1826) 173.— 265 Plate XIV. 1—Serratula quinquefolia MB.; 2—S. procumbens Rgl.; 3—S. trancaucasica (Bornm.) Sosn. ex Grossh.; 4—S. biebersteiniana (Iljin ex Grossh.) Takht. 269 266 Rhaponticum subgen. Mastrucium (Cass.) Less. Synops. Comp. (1832) 6.—Plants monoecious. Capitula heterogamous, quite large, ovate, several in clusters, forming corymbose inflorescence, sometimes soli- tary; peripheral florets pistillate, with larger corolla; disk florets bi- sexual, fertile, numerous. Type of section: S. coronata L. Note. Besides S. coronata L., this section includes S. insularis Iljin and S. koreana Iljin s. str. 2. S. coronata L. Sp. pl. ed. 2 (1763) 1144; DC. Prodr. VI, 667; Ldb. FI. alt. IV, 38; Fl. Ross. II, 2, 756; O. Hoffm. in Pflanzenfam. IV, 5, 326; Schamlh. FI. II, 112; Fedtsch. and Fler. Fl. Evrop. Ross. 1015; O. and B. Fedtsch. Perech. Rast. Turk. IV, 291; Grossh. Fl. Kavk. IV, 193; Iljin in Fl. Yugo.-Vost. VI, 413; Pavlov, Fl. Tsentr. Kazakhst. III, 315; Kitagawa, Lineam. Fl. Mansh. 470; Kryl. Fl. Zap. Sib. XI, 2935; Dobrocz. in Vizn. Rosl. URSR, 570; Majevski, Fl. ed. 8, 607; Popov, Fl. Sredn. Sib. II, 767.—S. coronaria Pall. Reise. II (1773) 280, 528.— S. altis-sima Steph. ex Ldb. Fl. Ross. II, 2 (1846) 761, in nota.—S. manshurica Kitagawa in Tokyo Bot. Mag. XLIX (1935) 229.— Mastrucium pinnatifidum Cass. in Dict. Sc. nat. XXXV (1825) 173, XLI (1826) 523.—Ic.: Gmel. FI. sibirica, II (1749) tab. XX; Syreistsch. Ill. Fl. Mosk. Gub. III, 309; Kom. and Alis. Opred. Rast. Dalnevost. Kr. II, Plate 326; Lapin, Korm. Rast. III, Fig. 473 —Exs.: GRF No. 296. Perennial. Rhizome woody, large, horizontal, with numerous string- like roots. Stems 35—150 cm high, sulcate, angular, in upper part more or less branched. Leaves green or reddish, sometimes glaucescent beneath, glabrous, less often sparsely hairy beneath on midrib, un- paired pinnate and pinnately divided, mostly with 3-8 pairs of lobes coalesed at base, oblong or elliptical, 10-40 cm long, to 12 cm wide, large-toothed, lobes serrate-toothed, spinulose-ciliate, sometimes with cartilaginous spines at tips of teeth, terminal lobe largest; lower leaves petiolate, cauline leaves sessile and smaller, with ovate-oblong or lan- ceolate lobes. Capitula quite large, ovate, at tips of stems and lateral branches, on conspicuous peduncles, sometimes solitary, often several in clusters, forming corymbose inflorescence. Involucres 1.0—1.8 cm in dia, 1.5-2.5 cm long, ovate to oblong-ovate, pubescent with short- appressed hairs, often brown or rusty-brown, tomentose, sometimes subglabrous; involucral bracts imbricate, usually reddish-brown, 5—6 rowed, all acuminate, outer ones ovate with short subulate cusp, inner ones oblong-lanceolate, narrow and long-acuminate. Florets lilac-purple, sometimes peripheral larger, undeveloped, with abortive stamens, ster- ile anthers and narrower, sometimes 3—4-parted corolla; disk florets 270 267 bisexual; corollas 20—27 mm long, corolla tube narrow, 15 mm long, limb about 10 mm long, lobes linear, about 5 mm long; anther tube about 10 mm long; anthers with filiform basal appendages about 3 mm long; style exserted from corolla, with 2 linear, grooved lobes about 3 mm long. Achenes 4-5 mm long and about 1.5 mm wide, oblong, smooth, finely sulcate, brownish; pappus 8—10 mm long, shorter than corolla. Flowering June to September. Forest and Steppe zones; forest and steppe meadows, on forest edges, shrubby thickets, insular forests, mountain slopes, in flooded and alkaline meadows, and sedge marshes.—European Part: Upper Volga, Volga-Kama, Upper Dnieper, Middle Dnieper, Volga-Don, Trans-Volga, Upper Dniester (?), Bessarabia, Black Sea Region, Crimea (?), Lower Don, Lower Volga; Caucasus: Cis-caucasia; Western Siberia: Upper Tobol, Irtysh, Altai; Eastern Siberia: Yenisei, Lena- Kolyma, Angara-Sayans, Dauria; Far East: Zeya-Bureya, Uda River area, Ussuri; Soviet Central Asia: Aralo-Caspian, Lake Balkhash Re- gion, Dzhungaria-Tarbagatai, Tien Shan. General distribution: Central Europe, Mongolia (?), Japan, China. Described from Siberia and Italy. Type in London. Note. A polymorphic, widely distributed plant. The size and num- ber of capitula vary on the plant. In the Far East, the new var. orientalis Boriss. has been discovered. Its capitula are much smaller than in European and West Asian plants; the involucres are copiously brown- ish-villous; all the plants are smaller in size; and the leaves often have fewer pairs of sparsely toothed lobes. Economic Importance. A melliferous, medicinal, and vitamin- containing plant; traces of alkaloids have been found in its stem. Section 3. Piptochaete Boiss. Fl. or. III (1875) 590.—Capitula 0.4-1.5 cm in dia, oblong or oblong-ovate, numerous, clustered in corymbose-paniculate inflorescence; all florets similar, bisexual; pap- pus bristles few, deciduous. Plants scabrously pubescent, branched from base; leaves pinnately incised or divided. Type of section: S. erucifolia (L.) Boriss. Series 1. Xeranthemoides Boriss.—Leaf lobes not cuspidate; outer involucral bracts with recurved cusp. 3. S. erucifolia (L.) Boriss. comb. nova.—S. xeranthemoides MB. Fl. taur.-Cauc. II (1808) 265, III, 545; DC. Prodr. VI, 669; Ldb. FI. Ross. II, 760; Boiss. Fl. or. III, 590; O. Hoffm. in Pflanzenfam. IV, 5, 326; Schmalh. FI. II, 112; Fedtsch. and Fler. Fl. Evrop. Ross. 1016; O. and B. Fedtsch. Perech. Rast. Turk. IV, 292; Grossh. Fl. Kavk. IV, 27 — 268 193; Iljin in Fl. Yugo-Vost. VI, 415; Pavlov, Fl. Tsentr. Kazakhst. III, 317; Ter-Khachaturova in Fl. Gruzii, VII, 525; Majevski, Fl. ed. 8, 609; Agadj. in Fl. Azerb. VIII, 426—Xeranthemum erucifolium L. Sp. pl. (1753) 858.—Ie.: Gmel. Fl. sibirica, I] (1749) tab. XLVII; Larin. Korm. Rast. III, Fig. 475. Perennial. Whole plant bristly from long, papilliform, flexuous moniliform-articulate hairs. Stems 20-60 cm high, with fibrous rem- nants of old leaves at base, divaricate, strongly dichotomously branched throughout, with woody, spreading branches, forming globose bush. Basal and lower cauline leaves petiolate, oblong-lanceolate, oblong- oval or oval, pinnately divided or incised only at base, less frequently pinnately incised or lobed with lanceolate, acute, toothed lobes; cauline leaves sessile, middle leaves pinnately incised into narrow, entire or sparsely-toothed lobes, gradually transitional to undivided and entire, linear, smaller upper leaves. Capitula oblong-ovate to cylindrical, more or less small 0.4-1.5 cm in dia, numerous, at tips of branches, clus- tered in broad, compound corymbose-paniculate inflorescence. Involu- cres dull greenish-yellow, glabrous or sparsely arachnoid-hairy; in- volucral bracts coriaceous, imbricate, outer and middle bracts ovate, broad, scarious on margin, with short or more or less long cusp at tip, usually recurved. Receptacle densely covered with long, smooth bristles, as long as involucre. Corollas light purple, almost pink, sometimes white (albinos), about 10 mm long; corolla tubes narrow, 4-5 mm long, limb about 6 mm long with linear teeth about 4 mm long. An- thers connate in tube, as long as corolla, free above and obtuse, with filiform basal appendages 0.5 mm long; filaments glabrous. Style short- hairy, exserted from corolla. Achenes oblong 4-5 mm long, weakly compressed, only at base slightly narrowed, weakly sulcate, with slightly raised longitudinal ribs, brownish or almost black; pappus deciduous, reddish, consisting of several rows of deciduous, toothed, scabrous hairs 4—S mm long. Flowering May to August. Alkaline soils, “solonetzes” [alkali soils], dry slopes, weedy places, fields, to mid-montane zone.—European Part: Volga-Kama, Middle Dnieper, Volga-Don, Trans-Volga, Bessarabia, Black Sea Region, Crimea, Lower Don, Lower Volga; Caucasus: Ciscaucasia, Dagestan, Eastern and Southern Transcaucasia; Western Siberia: Upper Tobol (southwestern part); Soviet Central Asia: Aralo-Caspian. Endemic. Described from Don River. Gmelin’s illustration is the type. Series 2. Dissectae Boriss. —Leaf lobes with cartilaginous cusp; outer involucral bracts finely floccose, with long spiniform cusp more or less deflected. 272 269 4. §. dissecta Ldb. Fl. alt. IV (1833) 40; Ldb. Fl. Ross. II, 2, 759 p. p.; DC. Prodr. VI, 669; O. and B. Fedtsch. Perech. Rast. Turk. IV, 292; Pavlov, Fl. Tsentr. Kazakhst. II], 316; Kryl. Fl. Zap. Sib. XI, 2940.—S. trichocephala Pall. in herb.—Ic.: Ldb. Ic. pl. Fl. Ross. V (1834) tab. 489. Perennial. Roots long, string-like. Rhizome oblique, short, thick. whole plant grayish, with stiff hairs. Stems solitary or 2, 15-40 cm high, branched only in upper part, straight, angular, leafy, covered with flexuous, articulate, scattered hairs, with fibrous remnants of old leaves at base. All leaves, except bracteal, pinnately divided, stiff; basal leaves often withering before flowering, oblong, 10—15 cm long, 1.5—3.5 cm wide; lower cauline leaves like basal with basally broad- ened, grooved petioles, middle and upper leaves remote, sparse sessile, 2-10 cm long, 1.0-3.5 cm wide, gradually reduced upward and less, incised; all leaves covered on both sides with scattered, long, articu- late and shorter bristly hairs; leaf lobes lanceolate or linear, 1—3(5) mm wide, pinnately lobed or toothed, sometimes undivided, tips of teeth with cartilaginous cusp. Capitula almost in corymbose-paniculate inflorescence. Involucre cylindrical, about 0.7—1.5 cm long, 0.5—1.0 cm in dia, round at base; involucral bracts mostly green, sometimes with brownish spot below cusp, puberulent; outer bracts broadly ovate, like shorter, oblong-lanceolate, gradually elongating, and narrower middle bracts with curved, more or less deflected, finely subulate cusp, 2-3 mm long; innermost bracts elongate-lanceolate, acuminate, light- colored. Corollas lilac-purple or pink, 14-16 mm long; corolla tubes about 7 mm long; limb lobed to half with linear lobes about 4 mm long, with very sparse small appressed hairs on outer side. Anther tubes often surpassing corolla; filaments with small papillae; anther appendages linear, 0.5-0.75 mm long. Achenes 5—6 mm long, elon- gate, yellowish or yellowish-reddish-brown, longitudinally finely sul- cate; pappus whitish, about 12 mm long, with short-toothed hairs. Flowering May to July. (Plate XV, Fig. 2). Stony and rubbly slopes of high steppes, low rounded, isolated hills, desert-steppe regions, clayey, sandy and alkaline steppes.—So- viet Central Asia: Aralo-Caspian, Lake Balkhash Region, Dzhungaria- Tarbagatai. General distribution: possibly in Dzhungaria-Kashgaria. Described from Arkaul Mountains and Dolon Kar (between Zaisan and Kurchum River. Type in Leningrad. Note. It is similar to S. angulata Kar. and Kir., from which it is distinguished by the pubescence and shape of the involucre and involucral bracts. 3 270 5. S. angulata Kar. and Kir. in Bull. Soc. Nat. Mosc. XIV (1841) 453; Ldb. FI. Ross. II, 2, 759; Pavlov, Fl. Tsentr. Kazakhst. III, 316.— S. dissecta var. angulata (Kar. and Kir.) Trautv. Enum. pl. Songor. (1866) No. 684; Sergievskaja in Kryl. Fl. Zap. Sib. XI (1949) 2941. Perennial. Root long, string-like. Rhizome oblique, short. Stem 15—40 cm high, angular scabrous, almost arachnoid-hairy. Leaves pin- nately incised with linear, undivided cuspidate lobes. Capitula numer- ous, in corymbose inflorescence, somewhat small. Involucres subglabrous, narrowly cylindrical, almost cuneately narrowed at base; involucral bracts imbricate, 4-rowed, outer ones with spine 3—5 mm long, usually erect and straight; inner bracts with elongate, acuminate appendage. Corollas pink or purple, about 15 mm long; corolla tube about 6 mm long, filiform; limb about 8 mm long, less than 1 mm wide, with linear lobes about 4 mm long. Anther tube somewhat shorter than corolla; style somewhat surpassing corolla. Achenes ovate ob- long, about 4 mm long and 2 mm wide, weakly scabrous-hairy. Flow- ering June to July. Sands.—Soviet Central Asia: Lake Balkhash Region. Endemic. Described from desert along Tonsyk River, near Ayaguz. Type in Leningrad. Note. This species is similar to S. dissecta Ldb., from which it is distinguished by a narrower cylindrical involucre that is cuneately narrowed to the base (and not rounded at the base), involucral bracts that are subglabrous, and longer, straight, hard spines. Section 4. Klasea (Cass.) DG. Prodr. VI (1837) 668.—Klasea Cass. in Dict. Sc. nat. XXXV (1825) 173, XLI (1826) 521.—Capitula medium-to large-sized, on more or less leafy peduncles; all florets similar, bisexual; achenes with pappus. Type of section: S. centauroides L. Series 1. Centauroides Boriss.—Stems well-developed, mostly strong; leaves pinnately incised, divided, or lobed, usually scabrous on margin, often on veins, often also on surface sometimes glabrous; capitula large- and medium-sized, mostly solitary, on leafless or leafy peduncles. This series combines similar, sometimes difficult-to-distinguish, geographically separated species. 6. S. centauroides L. Sp. pl. (1753) 820, ed. 2 (1763) 1148; DC. Prodr. VI, 668; Ldb. Fl. Ross. II, 2, 757; Popov, Fl. Sredn. Sib. II, 767.—S. pectinata Turcz. ex Herd. in Bull. Soc. Nat. Mosc. XLIII, 1 (1870) 94; Trautv. in Tr. Peterb. Bot. Sada, VIII, 509.—Klasea 274 271 Centauroides (L.) Cass. in Dict. Sc. nat. XXXV (1825) 173.—Ie.: Gmel. FI. sibirica, II (1749) tab. XVII. Perennial. Roots string-like, numerous, brown. Rhizome oblique, quite robust. Plants (10)30—80 cm high. Stems covered at base with fibrous remnants of petioles of old leaves, mostly angular, sulcate, more or less pubescent, simple, often branched, leafy to tip. Leaves pinnately incised or divided; leaf lobes oblong to linear, large-toothed, alternate and remote, at acute angle to rachis; lower leaves petiolate, pinnately incised, less often pinnately lobed, spiny-toothed in upper part, scabrous on margin and veins from short, curved hairs, less sca- brous on surface, with pubescent petioles; upper leaves short-petiolate. Capitula several, ovate, 2-3 cm in dia, narrowed at base, on peduncles, leafy to capitula. Involucres in upper part slightly constricted; involu- cral bracts imbricate, 5-rowed, yellowish, glabrous or subglabrous, greenish at tip, acute and with short cusp, about 0.5 mm long, with 5 dark veins, arachnoid-hairy; lower involucral bracts ovate, middle ones to oblong, inner ones lanceolate, longer than others, with scarious, spinescent appendage, innermost linear, with spiny-toothed, membra- nous appendage, tip and margin of appendage narrowly crisped-scari- ous. Corolla pink, narrowly tubular, tube about 8 mm long, limb 13 mm long, with linear lobes to 7 mm long; anthers almost as long as corolla, with obtuse, apical appendage about 1 mm long, and linear basal appendage; filaments papilliform, pubescent with hairs. Style 4— 5 mm, exserted from corolla, short-bilobate. Achenes oblong, about 5 mm long, constricted at tip and expanded crown-like below pappus, brownish, sulcate; pappus short, about 5 mm long, dense, of unequal shortly plumose bristles. Flowering June to August. Dry stony slopes, rocks, in mixed herb steppes, sandy steppes, and sands, limestones, sometimes in abandoned fields and crops.—Eastern Siberia: Angara-Sayans, Dauria. General distribution: Mongolia (north- ern Mongolia). Described from Siberia. Type in London. 7. S. komarovii Iljin in Izv. Glavn. Bot. Sada, XXVII, 1 (1928) 89.—S. centauroides auct. non L.: Kom. Fl. Manchzh. III (1907) 757; Kom. and Alis. Opred. Rast. Dalnevost. Kr. II, 1086.—Ic.: Kitagawa in Tokyo Bot. Mag. XLVIII (1934) 913. Perennial. Root string-like, numerous. Rhizome oblique, less often horizontal. Plants 80-150 cm high. Stems solitary or few, with fibrous remnants of old leaves at base, erect, simple, sometimes with few branches, purple in lower part and covered with crisped short hairs, subglabrous and leafless in upper part. Leaves all similar, oblong; basal and lower cauline leaves petiolate, usually pinnately divided, 275 272 with alternate lobes, often terminal lobe largest, less often lyrate; middle and upper cauline leaves sessile, pinnately divided, mostly with largest terminal lobe and oblong or lanceolate, toothed or, in upper part, entire lateral lobes with cartilaginous cusp; uppermost leaves reduced, brac- teal, entire; all leaves glabrous, subglabrous or mostly lower scatteredly hairy, ciliolate-hairy on margin. Capitula 2—3 cm in dia, solitary, less often 2, on peduncles 20-50 cm long. Involucres subglobose, con- stricted in upper part; outer and middle involucral bracts ovate or ovate-oblong, sulcate, often blackish in upper part, with narrow scari- ous margin, violet, with short, straight or slightly curved cusp and usually almost woolly, glabrous on remaining surface; innermost in- volucral bracts lanceolate, longer than others, with elongate, yellowish appendage, often reddish, very short-hairy on outer side, crisped-scari- ous at apex and margin. Receptacle covered with long smooth bristles. Florets reddish or violet; corolla 20-25 mm long; corolla tube narrow, 11-12 mm long, almost as long as limb of narrow linear, thin lobes 7-9 mm long. Anther tube shorter than or as long as corolla; basal appendages of anther about 1 mm long, almost undivided; filaments short-pubescent; style quite exserted from corolla, with short incon- spicuous branches. Achenes obconical, about 6 mm long and 2.5 mm wide, slightly constricted, scarcely toothed at tip, sulcate; pappus sor- did white with unequal, sulcate, deciduous bristles, 6-7 mm long. Flowering June. Dry stony slopes.—Eastern Siberia: Dauria; Far East: Zeya-Bureya, Ussuri. General distribution: China (north-eastern China). Described from Far East. Type (lectotype) in Leningrad. 8. S. radiata (Waldst. and Kit.) MB. Fl. taur.-cauc. III (1819) 545; DC. Prodr. VI, 668; Ldb. Fl. Ross. II, 2, 757; Boiss. Fl. or. III, 588; O. Hoffm. in Pflanzenfam. IV, 5, 326; Grossh. Fl. Kavk. IV, 194.—S. centauroides auct. non L.: MB. op. cit. II (1808) 264.—S. radiata ssp. bracteifolia Ijin ex Grossh. loc. cit.; Iljin in Fl. Yugo- Vost. VI, 414.—S. bracteifolia (Ijin ex Grossh.) Stank. in Stank. and Tal. Opred. Vyssh. Rast. (1949) 670; Dobrocz. in Vizn. Rosl. URSR, 571.—Carduus radiatus Waldst. and Kit. Descr. et icon. pl. rar. Hung. I (1802) 9.—Ie.: Gmel. Fl. Sibir. II (1749) tab. XVIII; Waldst. and Kit. op. cit. tab. 11; Rchb. Icon. Fl. Germ. XV, tab. 803; Fiori and Paol. Iconogr. FI. Ital. I, fig. 3708; Javorka and Csapody, Iconogr. Fl. Hung. 544; fig. 3893.—Exs.: Herb. norm. No. 5122. Perennial. Roots numerous, string-like, brown. Plants 30-100 cm high, scabrous, pubescent, or subglabrous. Stem with remnants of peti- oles of old leaves at base, virgate, angular-sulcate, erect, simple or weakly branched, with long branches, scabrous in lower part, setose. 276 273 Basal and lower cauline leaves long-petiolate, oblong-ovate, pinnately lobed, divided or incised, with linearly lanceolate or lanceolate, acute, entire or sparsely toothed lobes; leaves on branches smaller, undivided or pinnately divided; upper leaves sessile. Capitula on long peduncles, solitary, 1.0—1.5 cm in dia and about 1 cm long, subglobose. Involu- cral bracts glabrous, lustrous, coriaceous, yellowish-green, imbricate, 5—6-rowed, quite small; outer and middle ones ovate-triangular or oblong-ovate, with cartilaginous thin cusp, 1-2 mm long, more or less bent or almost curved; innermost involucral bracts elongate, lanceolate to linear, with long, membranous appendage, obtuse, and toothed-erose. Corolla pink, about 18 mm long; corolla tube very narrow, about 10 mm long, limb campanulate, about 9 mm long with linear, acuminate lobes about 5 mm long. Anther tube almost as long as corolla or shorter, anther appendages filiform, about 1 mm long; filaments with papillate hairs. Style exserted from corolla or as long, short-bilobate, its lobes about 0.5 mm long. Achenes 4—5S mm long, 1.5—2.0 mm wide, sometimes almost tetraquetrous, glabrous, finely sulcate, slightly curved, yellowish- or reddish-brown to brown, acute at base; pappus 5—7 mm long, yellowish, scabrous from very short, toothed hairs of equal length. Flowering July to August. (Plate XVII, Fig. 2). Dry montane forests, forest edges, shrub thickets, on limestones.— European Part: Upper Dnieper, Middle Dnieper, Volga-Don, Trans- Volga, Black Sea Region, Lower Don; Caucasus: Ciscaucasia, Dagestan, Eastern and Southern Transcaucasia. General distribution: Central Europe. Described from Hungary. Type in Budapest. Note. It is distinguished from the similar species S. centauroides L. by the longer cusps of the involucral bracts, the form and degree of incision of the leaves, and the pubescence. 9. S. biebersteiniana (Iljin ex Grossh.) Takht. in Takht. and Fed. Fl. Yerevana (1945) 323; Grossh. Opred. Rast. Kavk. 489; Ter- Khachaturova in Fl. Gruzii, VIII, 526.—S. radiata ssp. biebersteiniana Ijin ex Grossh. Fl. Kavk. IV (1934) 194; Iljin in Fl. Yugo-Vost. VI, 414; Agadj. in Fl. Azerb. VIII, 425. Perennial. Roots string-like, numerous. Rhizome oblique, thick; stems 30-60 cm high, densely, finely woolly, sulcate, with fibrous remnants of older leaves at base. Leaves very dense to capitula, ap- pressed to stem, all pinnately divided mostly with larger terminal lobe, coarsely toothed, teeth recurved, cuspidate, acute and with spinous cusp, hispid-scabrous, on veins and particularly on margin, pinnately incised in lower part, with decurrent lobes, rachis winged; lower cauline leaves sometimes undivided. Capitula globose or globose-ovoid, 1.0-— 1.2 cm in dia, elongated in upper part, broader at base. Involucral 278 274 bracts pale green, firmly imbricate, with 3, green, short stripes above, at tip with long, erect, often recurved, stiff, yellow, straight and car- tilaginous spines 2-5 mm long; outer involucral bracts ovate, arach- noid-hairy brown, scarious on margin; middle bracts oblong-ovate, inner oblong-lanceolate to linear, longer than others, pale green in lower part with 3 dark green streaks, with membranous, light yellow, apically toothed and acute appendage. Corollas pink or pale pink, 20-25 mm long; corolla tube narrow, about 8 mm long; limb narrow, about 13 mm long with thin, linear lobes 5-6 mm long. Anthers as long as or slightly shorter than corolla, about 8 mm long, subacute, with short- linear, basal appendage about 0.5 mm long; filaments about 4 mm long. Style exserted from corolla by 2 mm, or as long; style branches very short, inconspicuous, subacute. Achenes oblong, about 5 mm long, 1.5—2.0 mm wide, weakly sulcate or smooth, glabrous, brown; pappus shorter than corolla lobes, 6-8 mm long, not dense, comprising weakly scabrous hairs. Flowering June to August. (Plate XIV, Fig. 4). Stony dry steppe slopes, among shrubs, in mid- and high-montane zones.—Caucasus: Dagestan, Eastern and Southern Transcaucasia, Talysh. General distribution: Armenia and Kurdistan. Described from Caucasua. Type (lectotype) in Leningrad. 10. S. gmelinii Tausch in Flora, XI, 31 (1828) 485; Kryl. Fl. Zap. Sib. XI, 2937, non DC. (1837).—S. isophylla Claus in Beitr. Pflanzenk. Russ. Reichs, VIII (1851) 118, 301; Fedtsch. and Fler. Fl. Evrop. Ross. 1016; Iljin in Fl. Yugo-Vost. VI, 4141; Majevski, Fl. ed. 8, 608.—S. centauroides var. isophylla (Claus) Schmahh. F1. II (1897) 113.— Ic.: Gmel. FI. sibirica II (1749) tab. XVIII; Ijin, op. cit. Fig. 704. Perennial. Roots string-like, brown. Stems 60—100 cm high, soli- tary, erect or slightly assurgent at base, simple, leafy, sulcate, sub- glabrous or flexuous-hairy all over or only in upper part. Leaves simi- lar, pectinate-pinnately incised, with lanceolate or linearly lanceolate, acuminate, serrate, less often entire, sometimes with crescent-shaped lobes, coalescent in upper part of leaf with their bases; terminal lobe often larger and broader than laterals; basal and lower cauline leaves with long, thin petioles, obovate or ovate-oblong, 8—10 cm long, about 3.5 cm wide; middle cauline leaves sessile, not amplexicaul and not decurrent; upper leaves reduced, like lower. Capitula solitary, on long, mostly leafless peduncles, subglobose, (1.5)2—2.5 cm in dia, with broad base. Involucre yellowish-green, subglabrous, lustrous or with sparse appressed tomentum; involucral bracts herbaceous, sometimes on margin indistinctly scarious, imbricate, 5—6-rowed; outer involucral bracts tri- angularly ovate or oblong-ovate, with very short, soft, readily fragile curved cusp, blackish at tip; innermost bracts elongate, lanceolate to 275 ({Z (i ee = ZA ( &' _ Leer NS BZ CA Zan Plate XV. 277 1—Serratula chartacea Winkl.; 2—S. dissecta Ldb. 3—S. lyratifolia Schrenk; 4—S. algida l\jin. 279 276 linear, colored, with lanceolate, yellow membranous appendage. Corolla purple, 15-20 mm long, almost 2 times as long as pappus; corolla tube narrow, 8-10 mm long; limb about 10 mm long, with narrow lobes about 5 mm long. Anther tube longer or almost as long as corolla. Style far surpassing corolla, distinctly bilobate. Achenes 5—6 mm long, 1.5—2.0 mm wide, oblong, 3—4-angled, subacute at base, slightly curved, glabrous and finely sulcate, brownish; pappus about 7 mm long, 1/2 as long as corolla tube, whitish or yellowish, compris- ing stiff, thick, scabrous, toothed, unequal bristles. Flowering June to July. @ Meadow steppes, among thickets of steppe shrubs, along edges in open forests, in flooding meadows, on limestones, sometimes in old fields and salt-marsh meadows, in forest-steppe and steppe zones.— European Part: Upper Volga, Volga-Kama, Volga-Don, Trans-Volga, Lower Volga; Western Siberia: Upper Tobol, Irtysh. Endemic. Described from Western Siberia. Gmelin’s illustration is considered as the: type. Note. It is distinguished from S. centauroides L. by an always simple, mostly straight stem, with a single large capitulum and highly leafless peduncle; coalescent upper lobes of the leaves; a much larger terminal lobe; the shape of the leaves; and the finely serrate-scabrous pappus bristles. 11. S. tanaitica P. Smirn. in Byull. Mosk. Obshch. Isp. Prior. Otd. Biol. XLIX (1940) 92; Majevski, Fl. ed. 8, 609. Perennial. Roots numerous, almost horizontal, string-like, about 1 mm thick, brownish. Rhizome thin, vertical or oblique, 1—2 cm long. Stem 40-70 cm high, about 3 mm thick, with fibrous remnants of petioles of old leaves at base, glabrous, cylindrical, sulcate, with many leaves or almost leafless. Basal leaves petiolate, petioles almost as long as oblong lamina. Leaves pinnately divided, with 8—12 remote lobes; leaf lobes almost similar, linear, 2-4 cm long, 2.5—4.0 mm wide, almost at right angle to rachis, almost falcate, shorter and nar- rower in basal leaves, entire like upper leaves, sometimes with occa- sional or several teeth, finely scabrous-ciliate along margin, raised beneath along midrib, with short subulate cusp; rachis in upper part narrowly winged, more broadly winged above, at base unwinged; uppermost leaves entire, linear to subulate, 0.5—2 cm long, about 2 mm wide, sometimes absent. Capitula solitary on leafless peduncles. In- volucre globose, 1.5—1.7 cm in dia, attenuate in upper part, rounded at base; outer involucral bracts ovate, acute, middle ones oblong-ovate, inner ones oblong, longer than others, yellowish, with 3—S green veins and narrow scarious border, at tip sparsely woolly to subglabrous, with 280 277 scarious, curved, flat cusp 1.5—2.0 mm long; innermost bracts oblong to linear, with glabrous apical appendage about 7 mm long. Corolla pale, dry, violet, or pale, violet, about 18 mm long; corolla tube about 8 mm long; corolla limb about 10 mm long, with linear lobes about 4 mm long. Anther tubes as long as corolla or slightly longer. Style scarcely exserted from corolla, short-bilobate; ovary oblong, glabrous, finely sulcate. Achenes unknown. Flowering May. Chalky, more or less turfy, slopes.—European Part: Lower Don. Endemic. Described from mouth of Golubaya River in Volgograd Region. Type in Moscow; isotype in Leningrad. Note. This species is similar to S. radiata (Waldst. and Kit.) MB. and S. gmelinii Tausch. It differs from the first by having involucral bracts with a straight cusp, about 5 mm long, and from the second by having a glabrous stem, almost uniform, markedly narrower and entire leaf lobes, smaller capitula, and a pale corolla. Series 2. Heterophyllae Boriss—Stem straight, simple, with rela- tively few leaves. Basal leaves petiolate, undivided, large-toothed or pinnately lobed; cauline leaves mostly in lower part of stem, pinnately lobed to incised. Capitula large, solitary, on long leafless peduncles. S. caput-nejae Zahariadi, described from Romania and very close to S. heterophylla (L.) Desf., belongs to this series. 12. S. heterophylla (L.) Desf. Tabl. ecol. bot. Mus. hist. nat. Paris (1804) 93; DC. Fl. Fr. IV, 86; Prodr. VI, 669; Ldb. Fl. Ross. II, 2, 758; O. Hoffm. in Pflanzen fam. IV, 5, 326; Schmalh. Fl. II, 113; Fedtsch. and Fler. Fl. Evrop. Ross. 1016; Dobrocz. in Vizn. Rosl. URSR, 570.—S. nitida (Waldst. and Kit.) Bess. Enum. pl. Volhyn. (1822) 32.—Carduus heterophyllus L. Sp. pl. (1753) 824, ed. 2 (1763) 1154.—C. nitidus Waldst. and Kit. Descr. et icon. pl. rar. Hung. I (1802) 52.—Klasea heterophylla (L.) Cass. in Dict. Sc. nat. XXXV (1825) 173.—Ie.: Waldst. and Kit. loc. cit. tab. 52; Rchb. Icon. FI. Germ. XV, tab. 804; Fiori and Paol. Iconogr. FI. Ital. I, fig. 3709; Javorka and Csapody, Iconogr. Fl. Hung. 544, fig. 3891.—Exs.: Herb. norm. No. 5123; Fl. Cechosl. exs. No. 96. Perennial. Roots string-like, numerous. Whole plant sparsely papil- late-scabrous. Stem virgate, simple, straight, 50-80 cm high. Leaves relatively few, mostly in lower part of stem, pinnately incised to lobed, reduced above to narrowly linear; basal leaves petiolate, ovate, lobed or undivided, broad and large-toothed, cuspidate, sometimes pinnately divided in lower part, lyrate. Capitula solitary, on long leafless pe- duncles, medium- or large-sized, 1.5—2.0 cm in dia, subglobose, broader 278 toward base. Involucral bracts glabrous, coriaceous, imbricate, 6-rowed, outer ones ovate-triangular with five greenish-brown, narrow stripes, at tip narrowed into short cusp, with narrow membranous reddish- brown border; innermost bracts oblong, linearly lanceolate, acuminate and apically undulate, membranous. Corolla pink, about 20 mm long; corolla tube about 10 mm long, limb 10 mm long, 1 mm wide, with linear, acute lobes about 5 mm long. Anther tubes almost as long as corolla. Style considerably surpassing corolla. Achenes oblong; pap- pus about 6 mm long, comprising straight, scabrous bristles. Flowering June to July. Meadows and steppes, moist herb slopes, in scrubs, on limestones, in mountains at 900—1,000 m.—European Part: Upper Volga, Volga- Kama, Upper Dnieper, Middle Dnieper, Volga-Don, Trans-Volga, Bessarabia, Black Sea Region. General distribution: Atlantic, northern and Central Europe, Mediterranean. Described from northern Europe. Type in London. Series 3. Nitidae Boriss.—Leaves undivided, entire, or upper leaves with few large teeth or lobes, mostly glabrous or short-ciliate on mar- gin, basal leaves long-petiolate; capitula solitary or 2—7 on lateral branches of stem, large, about 3 cm in dia, or smaller, 0.8—1.5 cm in dia. 13. S. cardunculus (Pall.) Schischk. in Kryl. Fl. Zap. Sib. XI (1949) 2938; Grubov, Konsp. Fl. Mong. (1955) 279.—S. centauroides Spreng. Syst. veg. 3 (1826) 389, non L. nec MB.—S. nitida Fisch. ex Spreng. ibid., 390; DC. Prodr. VI, 668; Ldb. Fl. Ross. II, 2, 759; Schmalh. Fl. II, 114; Fedtsch. and Fler. Fl. Evrop. Ross. 1016; O. and B. Fedtsch. Perech. Rast. Turk. IV, 292; Iljin in Fl. Yugo-Vost. VI, 416; Pavlov, Fl. Tsentr. Kazakhst. III, 316, non Bess.—S. nitida a. typica Trautv. in Bull. Soc. Nat. Mosc. XXXIX, 2 (1886) 73; Kryl. Fl. Alt. III, 715.—Centaurea cardunculus Pall. Reise, I (1771) 500.—Ie.: Gmel. Fl. sibirica, II (1749) tab. XL; Zachariadi in Bull. Sect. Scient. XXVIII, 5 (1946) pl. IV, B. f. g. h. Perennial. Rhizome short, thick, with numerous string-like roots. Stem straight, 20-50 cm high, simple, less often branched in upper part, with brown remnants of old leaves at base, glabrous or sparsely hairy, sulcate. Basal leaves long-petiolate, elliptical or oblong-ellipti- cal to ovate, S—20 cm long, 1—5 cm wide, obtuse or short-acuminate, glabrous, finely ciliate on borders, glaucescent beneath, entire, with remote teeth or small-toothed, decurrent, sometimes broadly sinuate; cauline leaves sessile, smaller, fewer, lanceolate, entire, toothed to pinnately lobed; upper cauline leaves reduced, undivided, linearly 282 279 lanceolate or linear. Capitula on long peduncles, 2—7 on lateral branches of stem, less often solitary. Involucres 0.8—1.5 cm in dia, subglabrous, very sparsely hairy; involucral bracts greenish, sometimes with dull brownish spot at tip, imbricate, 5-6 rowed; outer bracts ovate, acumi- nate; inner ones linearly lanceolate, scarious at tip, middle ones ovate- lanceolate, with short, somewhat attenuate, cusp 0.5-1.0 mm long. Corolla pale, lilac-pink, about 15 mm long, with narrow tube, 4-5 mm long, gradually merging with deeply lobed limb about 9 mm long; limb lobes linear, about 5 mm long. Anthers with linear basal appendage, at tip obtusely short-lobate. Style somewhat surpassing corolla, short- bifid with divergent lobes. Achenes about 9 mm long, oblong, brown, finely sulcate, narrowed toward base, apically attenuate and crowned or small-toothed; pappus 8—9 mm long, whitish-yellowish, of unequal, short-plumose hairs. Flowering May to June. (Plate XVI, Fig. 2). Forest-steppe and steppe zone, steppe, often alkaline meadows, shores of salt lakes, in desert-steppes, now and then in old fields, shallow sands and sandy steppes.—European Part: Volga-Kama, Trans- Volga, Lower Don, Lower Volga; Western Siberia: Upper Tobol, Irtysh, Altai; Soviet Central Asia: Aralo-Caspian, Lake Balkhash Region, Dzhungaria-Tarbagatai. General distribution: Mongolia (northwestern part of Mongolia). Described from Volga. Type in London. 14. S. marginata Tausch in Flora, XI, 31 (1828) 484; Kryl. Fl. Zap. Sib. XI, 2939; Grubov, Konsp. Fl. Mong. 279.—S. glauca Ldb. in Mém. Acad. Sc. Pétersb. V (1812) 560; FI. alt. IV, 38 and FI. Ross. li, 2, 758 p. p.; DC. Prodr. VI, 668; Zahariadi in Bull. Sect. Scient. XXVIII, 5 (1946) 18, non L. (1753).—S. centauroides Ldb. FI. alt. IV (1833) 39, non L.—S. gmelini Ldb. ex DC. Prodr. VI (1837) 668 p. p. non Tausch (1828).—S. laxmanni Fisch. ex DC. ibid., 669, pro syn.— S. nitida B. glauca (Ldb.) Trautv. in Bull. Soc. Nat. Mosc. XXXIX, 2 (1866) 379; Kryl. Fl. Alt. H, 715; Schmalh. Fl. II, 114 p. p—Centau- rea uniflora Pall. Reise. III (1776) 252, 321, non L. (1767).—Carduus altaicus Patrin ex DC. op. cit. 669, pro syn.—C. uniflorus Turcz. ex DC. op. cit. 669, pro syn—lIc.: Gmel. Fl. sibirica, II (1749), tab. XXXIX, fig. 2; Ldb. Ic. pl. Fl. Ross. I, tab. 32. Perennial. Roots cord-like, numerous. Rhizome short, nodulose, oblique, ascending. Stems 15—70 cm high, solitary, simple, subglabrous, ribbed, glaucous, usually leafless above. Leaves glabrous, glaucous; basal and lower cauline leaves mostly long-petiolate; petioles almost as long as oblong or ovate lamina (1) 2—4 cm wide, undivided or some with few large teeth or lobes or pinnatifid, obtuse or short-acuminate, short-ciliate along margin; upper cauline leaves sessile, lanceolate, 4—9 cm long and 0.4—1.5 cm wide, coarsely and sharply toothed or 283 280 pinnatifid, sometimes with few teeth or undivided; uppermost leaves small, linear. Capitula solitary, quite large. Involucre semiglobose or ovate-semiglobose, 1.5—2.0 cm in dia, slightly arachnoid-hairy; outer involucral bracts ovate or ovate-oblong, gradually narrowed above and with spine 1-2 mm long, brownish or blackish along margin in upper part; inner bracts oblong, with scarious, obtuse or acute, fimbricate-ciliate, lacerate or toothed, dorsally appressed, short-hairy appendage. Corolla pinkish-violet, greatly exserted from involucre, about 20 mm long; corolla tube filiform-tubular, 10 mm long; limb about 10 mm long, with linear lobes, 5 mm long. Anther tube almost as long as corolla, in upper part with obtuse, free lobes, with filiform basal appendage about 1 mm long; filaments free, papillately hairy. Style surpassing corolla, very short-pubescent, bilobed, lobes about 1 mm long. Achenes oblong, 4 mm long, 2 mm wide, somewhat attenu- ated in upper part, narrowed toward base, finely sulcate; pappus about 15 mm long, yellowish with scabrous, very short-plumose hairs of varying length. Flowering June to July. Mountain steppes, dry stony slopes and rocks, in forest and sub- alpine mountain zones, in larch forests, along forest edges, in forest meadows and on meadow slopes, coastal gravels—Western Siberia: Irtysh, Altai; Eastern Siberia: Lena-Kolyma, Angara-Sayans, Dauria; Soviet Central Asia: Dzhungaria-Tarbagatai. General distribution: Dzhungaria-Kashgaria, Mongolia, China. Described from Siberia. Gmelin’s illustration is the type. Note. De Candolle (DC. op. cit. 670) recognizes S. glandulifera Turcz., described from the Khor steppe in Selenga Dauria from the collections of Turczaninow, and characterized by the unlobed, runci- nate leaves, and also a glandular stem. However, M.G. Popov (7. Sredn. Sib., U1, 768) considers this species an abnormal form of S. glauca Ldb. (= S. marginata Tausch) or S. centauroides L. 15. S. algida Ijin in Fedde, Repert. XXXV (1934) 357; Kryl. FI. Zap. Sib. XI, 2940; Czerneva in Fl. Uzbek. VI, 389. Perennial. Rhizome obliquely ascending or horizontal, with copi- ous, thin, string-like, fibrous roots. Plants 25—80 cm high, glaucescent or light green. Stems solitary, covered at base with fibrous remnants of old leaves, straight, simple, sulcate, glabrous or scatteredly pubes- cent, sometimes reddish at base, leafy. Leaves dull green on both sides, glabrous or scatteredly hairy on midrib, finely short-ciliate on margin or subglabrous; basal and lower cauline leaves 5.5—20.0 cm long, 0.7— 2.5 cm wide, oblong-ovate to oblong-lanceolate, long-petiolate, obtuse or with short, thin cusp, often withering at fruiting; cauline leaves sessile, small, oblong to lanceolate, uppermost narrowly linear; all leaves 284 281 entire, occasionally, almost toothed. Capitula solitary, 1.5-3.5 cm in dia, semi-globose, on long peduncles. Involucres 1.5—2.0 cm long and 1.5—3.0 cm in dia; involucral bracts imbricate, 6-rowed, almost coria- ceous, initially sparsely arachnoid-hairy, later subglabrous or with in- conspicuous pubescence, outer ovate or oblong-ovate, often black, middle ones broadly lanceolate all bracts acuminate or with 0.5—2.0 mm long cusp, gradually attenuate, with wide black stripe on margin and at tip; inner and innermost bracts oblong to lanceolate with elon- gated, stiff, almost scarious, acuminate or acute, undulate, irregularly, toothed and short-ciliate appendage. Corolla pinkish-reddish, 20-23 mm long; corolla tube 6-10 mm long, limb 9—13 mm long, with linear lobes 4.5—6.0 mm long; anther with short basal appendage 0.5—0.75 mm long; filaments papillose; style surpassing or as long as corolla, bifid, with long lobes. Achenes about 6 mm long, oblong, brownish, finely sulcate; pappus about 10 mm long, yellowish, finely barbate. Flowering June to August. Fruiting July to August. (Plate XV, Fig. 4). Rocks, stony and rubbly slopes, moraines, on herb slopes and passes, in cedar-larch and larch forests from alpine to forest zones, 1,500-3,600 m.—Western Siberia: Altai; Soviet Central Asia: Dzhungaria-Tarbagatai, Tien Shan. Pamiro-Alai. Endemic. Described from region of Kafarag Pass, descent from Anzob Pass. Type in Leningrad. 16. S. kirghisorum Iljin in Fedde, Repert. XXXV (1934) 338; Kryl. Fl. Zap. Sib. XI, 2939.—S. nidida var. sublyrata Kryl. ex Schischk. in Kryl. ibid., pro syn. Perennial. Roots numerous, fibrous, string-like. Rhizomes thick and short, almost horizontal. Plants 10-40 cm high, glaucescent. Stems almost always solitary, simple, straight, angular, glabrous or subglabrous. Leaves on both sides glabrous, scabrous and ciliate along margin with short dense cilia; basal leaves more or less numerous, like lower cauline leaves petiolate, ovate or oblong-ovate, almost lyrate or sinuate-toothed usually only in lower part, less often pinnately divided or undivided, obtuse, with very short cartilaginous cusp on teeth; middle cauline leaves sessile, mostly pinnately divided or lyrate, with oblong lobes or toothed; upper leaves reduced, linearly lanceolate, appressed to stem and mostly undivided. Capitula solitary, on more or less long peduncles. Involucres 1.2-1.5 cm long, 1.2—2.0 cm in dia, yellowish- green, lax arachnoid-tomentose at tip; outer involucral bracts ovate or oblong-ovate, middle ones broadly lanceolate, their cusp (1)2-3 mm long, blackish-brown at tip and with few dark veins; inner bracts often reddish, lanceolate, narrowed into scarious, linear appendage. Recep- tacle densely setose, bristles 3/4 as long as involucre. Corolla 15-19 28 N 282 mm long, pinkish-purple; corolla tube 7-9 mm long; limb with linear lobes 5-6 mm long. Anthers appendages 0.5—0.8 mm long filaments papillose. Style as long as or slightly surpassing corolla, short-bilo- bate. Achenes oblong, 5.0—-5.5 mm long, 1.5—2.0 mm wide, finely sul- cate, glabrous, lustrous, grayish or brownish-blackish, whitish at base, with small tooth at tip; pappus almost 1/2 as long as corolla, 7-9 mm long, yellowish, short-barbate. Flowering May to June. Desert-steppe zone, occasionally alkaline rubbly, clayey-stony and loamy slopes.—Western Siberia: Upper Tobol, Irtysh, Altai; Soviet Central Asia: Aralo-Caspian, Baltic Region, Dzhungaria-Tarbagatai. Endemic. Described from Chinghiz Mountains in region of Namas River in former Semipalatinsk Region. Type in Leningrad. 17. S. dshungarica Iljin in Fedde, Repert. XXXV (1934) 357. Perennial. Roots thin, string-like; rhizome woody, oblique, thick. Plants 30-75 cm high, glaucescent. Stems at base with remnants of old leaves, solitary or few, simple or weakly branched, angular, covered with long, beaded hairs. Leaves glabrous on both sides, less often scatteredly hairy beneath, ciliate on margin; basal leaves relatively few, like lower cauline leaves gradually narrowed into petiole, oblong, or oblong-ovate, pinnately divided, less often lyrate or undivided, with oblong-oval or oblong, obtuse lobes, cuspidate or sometimes acute; upper cauline leaves sessile, pinnately divided or toothed, with acumi- nate or acute lobes or teeth; apical leaves linear, almost entire. Ca- pitula 1-4 mostly solitary, on quite long peduncles, 2.5 cm in dia. Involucres 2.2—2.5 cm long, 1.5—2.2 cm in dia, arachnoid-hairy or tomentose; outer involucral bracts ovate or oblong-ovate, cuspidate, middle ones oblong-lanceolate, scarcely short-ciliate on-margin, at tip brownish-blackish, pubescent, gradually narrowed into brownish cusp 2—4 mm long; inner bracts lanceolate or linearly lanceolate short-hairy, at tip with membranous, oblong or linear, acuminate appendage, usu- ally hairy on margin and purple. Receptacle densely setose, bristles 1/3—3/4 as long as involucre. Corolla pinkish-purple, 19—25 (often 21— 22) mm long, corolla tube filiform, 8-11 mm long, limb 11-14 mm long, with linear lobes 5-8 mm long. Anther tube almost as long as corolla; anther appendages 0.5—1.0 mm long; filaments papillose. Style surpassing corolla, with 2 short lobes. Achenes 5—6 mm long and 2.0— 2.5 mm wide, oblong-ovoid, somewhat flat, sulcate, brownish or gray- ish, glabrous and lustrous, with small teeth at tip, somewhat narrowed at base; pappus 10-15 mm long, 1/2 as long as corolla, pale yellow, short-barbate. Flowering May to August. Common cattail—sheep’s fescue, often stony, steppe slopes, on tertiary clays, 1,100—1,800 m.—Soviet Central Asia: Dzhungaria- 286 283 Tarbagatai, Tien Shan. Endemic. Described from Dzhungaria Alatau, from vicinity of Sary-Dzhas. Type in Leningrad. 18. S. transcaucasica (Bornm.) Sosn. ex Grossh. Fl. Kavk. IV (1934) 194; Ter-Khachaturova in Fl. Gruzii, VII, 526; AgadJ. in FI. Azerb. VIII, 425.—S. glauca Ldb. FI. Ross. II, 2 (1846) 758 p. p.— S. haussknechtii Boiss. Fl. or. Il (1875) 589 p. p.—S. nudicaulis ssp. haussknechtii var. transcaucasica Bornm. in Vestn. Tifl. Bot. Sada, 32 (1914) 2.—S. haussknechtii var. transcaucasica (Bornm.) Grossh. and Schischk. Sched. ad Pl. or. exs. (1928) No. 224.—Exs.: Pl. or. exs. No. 224. Perennial. Roots numerous, string-like. Rhizome oblique. Stems 12-30 cm high, simple, sulcate, covered at base with fibrous remnants of old leaves. Leaves mostly at stem base, not coriaceous; basal leaves subglabrous, ciliate along margin, oblong, on short or more or less long, winged petiole; with thin, short spines at tip and on teeth, gradu- ally narrowed toward base, entire or in lower part with sparse, remote teeth; cauline leaves remote, few, linear, gradually reduced upward. Capitula small, 1.0-1.2 cm in dia, ovate-globose, solitary at tips of almost leafless stems. Involucral bracts subglabrous or pubescent in upper part, coriaceous, imbricate, 4—5-rowed; outer bracts triangularly ovate, with quite long, thin spines at tip; inner bracts much longer than outer, lanceolate, gradually narrowed into scarious appendage with short spine. Corolla pink, about 15 mm long; corolla tube 6-7 mm long; limb about 9 mm long with linear lobes 5—6 mm long. Anther tubes shorter than corolla; anthers with short, linear basal appendage about 0.5 mm long, obtuse, free; filaments short-papillose. Style surpassing corolla, short-bilobate. Achenes oblong-ovate, about 6 mm long, 2 mm wide, finely sulcate, brownish, somewhat narrowed toward base; pap- pus about 9 mm long, dense, of short unequal hairs. Flowering May to August. (Plate XIV, Fig. 3). Dry slopes, in middle and lower montane zones.—Caucasus: Dagestan, Eastern and Southern Transcaucasia, Talysh. General distri- bution: Asia Minor, Iran. Described from Caucasus. Type in Weimer. Note. S. haussknechtii Boiss. is a species extremely close to S. transcaucasica and possibly even identical to it. It differs in the shape of the more attenuate leaves, involucral bracts, plant height, range, and adaptation to the alpine zone of northern Iran. Series 4. Chartaceae Boriss.—Stems woody at base, branched or simple, sometimes reduced; leaves coriaceous or papery, undivided or pinnatifid, upper leaves oblong to lanceolate and linear. 284 19. S. chartacea Winkl. in Tr. Peterb. Bot. Sada, IX, 2 (1886) 524; O. and B. Fedtsch. Perech. Rast. Turk. IV, 293. Perennial. Root strong, brown, string-like. Rhizome woody, branched. Stem 30-40 cm high, almost woody at base, branched or simple, with cylindrical branches, sparsely pubescent, leafy almost to middle. Leaves thick, coriaceous, or papery, oblong, lyrate, 3-12 cm long, 1.5—5.5 cm wide, densely reticulate; lower ones narrowed into petiole, irregularly coarsely spiny-toothed and sinuate-toothed, along margin short-ciliate, glabrous above, sparsely pubescent beneath on veins, cuspidate; upper leaves gradually reduced, uppermost solitary, about 0.5 cm long, sessile, oblong-lanceolate, entire, cuspidate, subglabrous. Capitula solitary on glabrous peduncle, about 15 cm long, medium-sized 1.5—2.0 cm in dia, subglobose. Involucral bracts imbri- cate, densely below, sparsely pubescent above, outer ones coriaceous, triangular-ovate, elongate to lanceolate, almost spinescent at apex and brownish-blackish; inner ones linear, about 2 cm long, with scarious reddish cusp about 8 mm long, with undulate apical appendage. Re- ceptacle setose, bristles 8-9 mm long. Corolla red, about 22 mm long; corolla tube 8—9 mm long; limb campanulate, about 10 mm long, with linear, acute lobes 5 mm long. Anthers connate in middle, slightly shorter than corolla, free at tips, subobtuse, with basal linear append- age 1 mm long; filaments papillose. Style slightly exserted from co- rolla, shortly bilobate. Achenes oblong, 6 mm long, 2.0—2.5 mm wide, weakly attenuate, toward tip, weakly undulate on edges, densely sul- cate, glabrous; pappus dense, about 11 mm long, white, with short and thin plumose hairs. Flowering August to September (Plate XV, Fig. 1). Mountain slopes.—Soviet Central Asia: Pamiro-Alai. Endemic. Described from Pamiro-Alai. Type in Leningrad. Note. The form with entire leaves—f. integrifolia Iljin—has been reported from eastern Tadzhikistan. 20. S. procumbens Rgl. in Bull. Soc. Nat. Mosc. XL, 3 (1867) 165; O. and B. Fedtsch. Perech. Rast. Turk. IV, 294.—S. flexicaulis Rupr. in Mém. Acad. Sc. Pétersb. ser. VII, XIV, 4 (1869) 57; Trautv. in Tr. Peterb. Bot. Sada, VIII, 508.—S. depressa Rgl. and Schmalh. in Izv. Obshch. Lyubit. Estestv., Antrop. i. Etn. 34, 2 (1882) 52; Trautv. op. cit. non Stev.—Jurinea paulseni O. Hoffm. ex Pauls. in Kjoeb. Vidensk. Meddle. (1903) 155, (1909) 159. Perennial. Roots string-like, brown, numerous. Rhizome long, thick, creeping, with brown remnants of old leaves in upper part. Plants sub- glabrous or very short-pubescent, many-stemmed, many-headed, and woody at base. Stem reduced, 4—25 cm high, often procumbent, simple or with 2/4 branches; flowering stems as surgent, flexuous. Leaves 288 285 coriaceous, undivided, oblong-ovate to lanceolate, 5—8 cm long and to 2 cm wide; basal leaves narrowed toward base into short-winged peti- oles, apically attenuated, with widest part in upper third, irregularly sinuate and sharply serrate-toothed with straight or curved, short, car- tilaginous cusp; cauline leaves semiamplexicaul, upper ones gradually reduced, sessile, elliptical or oblong, in lower part sharply toothed, above almost undivided; uppermost leaves small, oblong to linear, entire. Capitula 1/3, solitary at tips of stems, large, 2-3 cm in dia, nodding, on straight, short peduncles, often with bracteiform apical leaves. Involucre 1.0—2.5 cm long, 1—2 cm in dia, light green, almost monochromatic, under lens short-woolly on margin, ovate-cylindrical, rounded at base, somewhat attenuated at tip, with many rows of imbri- cate bracts; outer involucral bracts triangular-ovate, sometimes with black cusp about 2.5 mm long, later oblong, green, with thin stripes, hairy on margin, tiny deciduous fluff, inner ones lighter in color, scari- ous, attenuate to linear, acuminate point, with scarious appendage; innermost violet in upper part, acute or subobtuse, lacking spine, longer than others, 2.0—2.5 cm long, linear, very finely pubescent on outside. Receptacle flat, long- and densely-setose, bristles white, almost as long as ovary. Florets red, pink, or pinkish-violet, violet when dry. Corolla about 23 mm long, narrowed over 9-10 mm; broader part of corolla tube about 12 mm long, with narrow, filiform, more intensely colored lobes 7-8 mm long. Anthers connate in middle, free above, obtuse, with linear basal appendage about 5 mm long. Style thin, long, far exserted from corolla or shorter, bilobed, with lobes 0.5—2.0 mm long. Achenes oblong, 5 mm long, and 1.5-2.0 mm_ wide, brown, indis- tinctly tetraquetrous, smooth or weakly sulcate; pappus about 12 mm long, many-rowed, short-barbate, stramineous, some bristles slightly shorter than corolla and shorter outer hairs longer than or as long as narrow part of corolla. Flowering June to September. (Plate XIV, Fig. 2). Dry stony and rubbly slopes of foothills and mountains, on steppe slopes, 1,700—4,000 m.—Soviet Central Asia: Tien Shan, Pamiro-Alai. Endemic. Possibly found in parts of China adjoining Pamir. Described from Tien Shan, Tekes. Type in Leningrad. 21. S. suffruticosa Schrenk. in Bull. phys.-math. Acad. Sc. Pétersb. III (1845) 110; Trautv. in Bull. Soc. Nat. Mosc. XXXIX, 2 (1866) 380; O. and B. Fedtsch. Perech. Rast. Turk. IV, 295. Perennial. Roots woody, strong, brown. Rhizome woody, often with numerous woody branches. Plants glabrous, at base woody, with brown remnants of petioles of old leaves. Stem branched above, with few long branches and solitary capitula. Leaves coriaceous, lower ones 289 286 oblong-lanceolate to linearly lanceolate, sometimes runcinate or pinnately incised into linear lobes, with winged petioles, sharply or spiny-serrate or sinuate-toothed; upper leaves linearly lanceolate to linear, entire. Capitula oblong or ovate, (1.5) 2.0-3.0 cm long, and 0.7—1.2 cm in dia, subglabrous. Involucral bracts greenish-stramineous, monochromatic, with very short deflected cusp; outer bracts ovate, cuspidate; inner bracts elongate to linear, almost scarious, acute, cus- pidate. Receptacle covered with scarious bristles shorter than achenes. Corolla pink, exserted from involucre, about 15 mm long; corolla tube about 5 mm long, bulged at base; limb campanulate, about 5 mm long, 1 mm wide, with narrow, filiform, acuminate, about 4 mm long lobes, somewhat attenuate at tip. Anthers somewhat shorter than corolla, fused over entire length of corolla, free at apex, with short basal appendage. Style not exserted from anther tube, linear, bifid, its lobes about 1 mm long. Achenes 5 mm long, and about 1.8 mm wide, oblong, glabrous, somewhat curved, with occasional furrows; pappus short, about 7 mm long, slightly longer than corolla tube; yellowish, with almost equal and almost smooth hairs. Flowering June. Stony slopes, on red gypsiferous sandstones and granites.—Soviet Central Asia: Lake Balkhash Region, Tien Shan, Dzhungaria- Tarbagatai. Endemic. Described from Tien Shan, Bektautu Mountains, Type in Leningrad. 22. S. aphyllopoda Iljin in Fedde, Repert. XXXV (1934) 359; Czerneva in Fl. Uzbek. VI, 390.—Ic.: Czerneva, ibid., Plate XL, Fuca . Perennial. Plants 20-30 cm high. Stems erect, almost leafless, sulcate, subglabrous, pubescent below capitulum, with one capitulum. Leaves oval or oblong-oval, obtuse, short-petiolate, sinuate-toothed, almost coriaceous, glabrous on both sides or noticeably scatteredly pubescent, mostly with prominent reticulate veins beneath; basal leaves numerous, 8-15 cm long and 3—7 cm wide; cauline leaves 2-3, re- duced, at base of stem. Capitulum medium-sized, about 1.5 cm in dia. Involucres glabrous, yellowish-green; outer involucral bracts ovate or ovate-oblong, middle ones oblong, with very short cusp, 0.5—1 mm long, blackish at tip, with 3 veins, inner ones linear with oblong, scari- ous, pale-purple appendage, attenuate and noticeably short-stiff-hairy basally on outer side. Corolla pale-purple, 20-24 mm long, narrowed over 10—12 mm, limb lobes 5.5—7.0 mm long. Basal appendages of anthers very short, about 0.25—0.5 mm long, flatish; filaments gla- brous. Immature achenes glabrous, mature achenes unknown; pappus whitish, densely barbate, with hairs almost as long as diameter of bristles. Flowering June to July. 290 287 Outcrops of gypsiferous rocks, in foothills —Soviet Central Asia: Tien Shan. Endemic. Described from Chatkal Range in area between Maili and Poneu. Type in Leningrad. Note. S. aphyllopoda Iljin is close to S. chartacea Winkl. and possibly even identical to it; additional material and observations in nature are needed. According to Iljin, S. aphyllopoda is close to S. alatavica C.A.M. and S. sogdiana Bge. but differs from them by the absence of the stem. Series 5. Sogdianae Boiss.—Leaves entire, incised-toothed to lyrate, with remote lobes; upper leaves sometimes deeply pinnately incised with few linear lobes, coriaceous or almost coriaceous, glabrous, lower ones short-petiolate, cauline leaves sessile. Capitula medium-sized, ovoid or oblong-ovate, with round base; involucral bracts with distinct cusp. 23. S. sogdiana Bge. Beitr. Kenntniss Russl. Stepp. Centr.-As. (1851) 191; Boiss. Fl. or. III, 588; O. and B. Fedtsch. Perech. Rast. Turk. IV, 293 p. p.; Czerneva in Fl. Uzbek. VI, 389. Perennial. Roots string-like. Rhizome thick, woody, branched. Stems 25—55 cm high, with adpressed, straight, thin and long branches, ribbed, glabrous, only on ribs scabrous, leafy. Basal and lower cauline leaves thin, coriaceous, oblong, mostly about 8 cm long, about 2.5 cm wide in middle, narrowed into short petiole, from base to middle in- cised-toothed, entire in upper part, with stipule-like, linear, deflected, straight, or curved auricles at base; middle cauline leaves and those on branches linearly lanceolate, about 3 cm long, gradually reduced, up- per ones almost linear, sessile, undivided or lyrate, with lanceolate terminal lobe and linear, short, lateral lobes, coriaceous, cartilaginous- spinescent, with acute, cartilaginous cusp. Capitula mostly solitary, 2.5 cm long, 1.5 cm in dia, obovate or ovate cylindrical, on more or less long peduncle, with several spinose, appressed scaly leaves. In- volucral bracts coriaceous, yellowish-green, glabrous, imbricate, out- ermost ovate, 4-5 mm long, gradually narrowed, with short, yellow, appressed cusp, on outside short-hairy; middle bracts oblong-lanceolate and lanceolate with deflected cusp 0.5—1.5 mm long, lower margin of cusp sparsely hairy, above base ciliate; innermost bracts linear, 20—22 mm long, scarious at tip and strong narrowed, ciliate almost to base, setose in middle on outside. Receptacle setose with smooth bristles about 1 cm long. Corolla purple or pink, about 16 mm long; corolla tube narrow, about 6 mm long; limb about 10 cm long, with linear lobes 5 mm long. Stamen filaments papillose; anther tube as long as corolla; anthers with obtuse, basal appendage not tailed. Style long, 291 288 longer than corolla, bilobed. Achenes reddish-brown, oblong-conical, about 5 mm long, narrowed at base, somewhat curved, almost smooth; pappus longer than corolla, about 10 mm long, with dense, gradually deciduous, very short and appressed plumose bristles. Flowering July to September; fruiting August to September (Plate XVI, Fig. 1). Mountain-steppe meadows, in juniper stands, on stony and rubbly slopes, screes, dry riverbeds and ravines, 2,500—3,000 m.—Soviet Central Asia: Tien Shan (Kirghizian Alatau), Pamiro-Alai (Gissar, Turkestan, Zeravshan and Alai ranges). Endemic. Described from Zeravshan. Type in Paris. 24. S. alatavica C.A.M. in Mém. Acad. Sc. Pétersb. sér. VII, XIV, 4 (1869) 56; Trautv. in Tr. Peterb. Bot. Sada, VIII, 508; Grubov, Konsp. Fl. Mong. 278.—S. dissecta var. asperula Rgl. and Herd. in Bull. Soc. Nat. Mosc. XL, 2 (1867) 116.—S. trautvetteriana Rgl. and Schmalh. in Tr. Peterb. Bot. Sada, VI, 2 (1880) 321.—S. sogdiana auct. p. p. non Bge.: O. and B. Fedtsch. Perech. Rast. Turk. IV (1911) 293.—Exs.: H.F.A.M. No. 498. Perennial. Rhizome thick, long, woody, with string-like roots. Stems 30-80 cm high, straight, angular and sulcate, branched, with long, cylindrical branches. Leaves glabrous, scabrous and spinous-toothed along margin; basal leaves oblong, undivided, triangular-toothed with short thin cusps; lower cauline leaves lyrate, with many, often almost linear, remote, lateral lobes, sometimes undivided, terminal lobe large, ovate; middle cauline leaves deeply pinnately incised, with few, al- most linear, deflected lobes; uppermost leaves elongated to linear, base of some lobes widened, sometimes subentire. Capitula solitary, some- times 2—3, about 2 cm long and 1.5 cm in dia, on leafless or more or less leafy peduncle. Involucral bracts with long, setaceous, deflected, cusp 3—5 mm long; outer bracts more or less pubescent mostly in young capitula; inner bracts greatly elongated, acuminate like bristle. Corolla pink, about 20 mm long; corolla tube about 6 mm long, almost as long as limb, 12 mm long and 1 mm wide; limb almost 1/2 as long as tube, with linear lobes about 5 mm long. Anther tube somewhat shorter than or almost as long as corolla. Style somewhat exserted from corolla. Achenes oblong-ovate, 4 mm long and 1.75 mm wide, sulcate, narrowed at base, slightly attenuate and undulate at apex, with black spots, brown; pappus almost as long as corolla tube, about 10 mm long, yellowish, of numerous unequal, very short-plumose (visible under lens!) hairs. Flowering June to September. Clayey, rubbly and stony steppe slopes, wormwood steppes and mountain meadows, among woody-scrub vegetation, 1,000—3,000 m.— Soviet Central Asia: Tien Shan, Pamiro-Alai. Endemic. Described from Alatau Mountains. Type in Leningrad. 292 289 Note. S. altavica C.A.M. was described from plants grown from seeds collected by Schrenk in the Alatau Mountains. S. trautvetteriana Rgl. and Herd. was described from the former Kokand Khanate (Shakhimardan). S. alatavica is a species similar to S. sogdiana Bge., but is distinguished from it by the large size of the plant as a whole, longer than 0.5—1.5 mm, recurved, not erect, spines of the involucral bracts. S. alatavica C.A.M. occurs in the more northern regions of the Kirgiz Alatau. Trans-Ili Alatau, and southern Kirghizia. S. sogdiana Bge. is confined mostly to Pamiro-Alai and to the Hissar, Turkestan, Zeravshan, and Alai ranges. 25. S. lancifolia Zak. in Fl. Uzbek. VI (1962) 515, 388. Perennial. Stems numerous, erect, 20-30 cm high, with thin, gla- brous, weakly pubescent below capitula, leafy, simple or branched in upper part, subramose, one-headed. Leaves thin-coriaceous, green on both sides, glabrous, entire, less often sinuate-toothed; basal and lower cauline leaves petiolate, lanceolate, subobtuse; middle cauline leaves gradually reduced, narrowly lanceolate, sessile, narrowed at both ends. Capitula about 2 cm long (with corolla) and 0.8 cm in dia, oblong- ovate, at base rounded. Involucral bracts numerous, 6-rowed, imbri- cate, coriaceous, obtuse, with very short cusp and 2-3 dark colored veins, arachnoid-hairy on margin; outer bracts ovate, inner ones lan- ceolate, erect, 15-16 mm long, puberulent on outside. Receptacular bristles smooth. Corolla pink, 15 mm long, corolla tube narrow, about 6 mm long; limb about 10 mm long and 1 mm wide, with linear lobes 4 mm long. Anther tube somewhat shorter than corolla. Style not ex- serted from corolla. Achenes 3 mm long and 1 mm wide, glabrous, with several furrows, brown, oblong-ovate, narrowed toward base, slightly attenuate at tip; pappus 7 mm long, slightly longer than corolla tube, of few, short-plumose hairs. Flowering July to August, fruiting August. Mid-montane zone.—Soviet Central Asia: Pamiro-Alai (Surkhan- Darya Region). Endemic. Described from Katarbulak on Kashka-Darya River. Type in Tashkent. Note. S. sogdiana Bge. is characterized by leaves that are undi- vided and entire in the upper part, very short cusps on the involucral bracts, and smaller capitula. 26. S. hastifolia Kult. and Korov. apud Ijin in Fedde, Repert. XXXV (1934) 360. Perennial. Rhizome short. Stem densely hairy at base, 30-50 cm high, solitary or less numerous angular-sulcate, glabrous and smooth, pale, often sordid-reddish, branched almost from base, branches long. 290 Leaves coriaceous, glabrous, smooth, undivided and entire, with promi- nent, mostly violet veins; basal leaves elliptical, gradually narrowed into petiole, obtuse or short-cuspidate; lower cauline leaves ovate- oblong, middle and upper ones ovate, all leaves sessile, auriculate, semiamplexicaul, obtuse, short-cuspidate; reduced toward stem tip. Capitula more or less numerous, on long peduncles, solitary terminal on branches. Involucres goblet-shaped, slightly constricted at tip, 1.2— 2.0 cm long and 1.2—1.5 cm in dia, yellowish-green, pubescent; involu- cral bracts coriaceous, outer and middle ones ovate or ovate-oblong, short-cuspidate, inner ones lanceolate, erect, gradually narrowed, re- curved on fruiting, sometimes with few teeth in upper part. Receptacle densely setose, bristles almost as long as involucre. Corolla pale-red- dish, 15-17 mm long, tube 5—7 mm, limb lobes 5—6 mm long. Basal appendage of anthers about 0.25 mm long, slightly broadened at tip and rounded; filaments papillose. Style exserted from corolla, more or less incised, with flat and recurved lobes. Achenes 5—6 mm long and about 2 mm wide, compressed, glabrous, smooth, dark, with some longitudinal streaks, broadened at tip and rounded; pappus scabrous with distant cilia. Flowering June to July. Soviet Central Asia: Tien Shan (western part). Endemic. Described from Talass Alatau (Karagoin). Type in Tashkent. Note. Described from the original diagnosis because of the ab- sence of herbarium material. Series 6. Coriaceae Boriss.—Leaves undivided or in lower part lyrate-pinnately lobed, coriaceos; capitula oblong or ovate-oblong, cuneately narrowed toward base; involucral bracts lacking distinct cusp or with very short deciduous cusp, yellow, or inner bracts violet. 27. S. coriacea Fisch. and Mey. in DC. Prodr. VI (1837) 668; Ldb. Fl. Ross. II, 2, 757; Boiss. Fl. or. III, 587; Grossh. Fl. Kavk. IV, 194; Agadj. in Fl. Azerb. VIII, 424.—Centaurea strictissima Boiss. and Buhse in Nuov. Mém. Soc. Nat. Mosc. XII (1860) 130.—Exs.: PI. or. exs. No. 374. Perennial. Root thick, woody. Stems 50-150 cm high, straight, sulcate, glabrous, in upper part paniculately branched, with long, cy- lindrical branches having few leaves. Leaves long- or short-petiolate, obovate or ovate-oblong, 9-25 cm long and (1.5) 4—9 cm wide, lower ones undivided, or lyrately pinnately lobed or pinnately divided in lower part, serrate or toothed, glabrous, lustrous, gradually cuneately narrowed toward base, winged; leaves on branches reduced, entire, upper ones pinnately lobed; lanceolate or oblanceolate, narrowed to- ward base. Capitula solitary, S—7 on plant, ovate, often oblong-ovate, 294 291 1.0—1.5 cm in dia, to 2.5 cm long, on subglabrous peduncle. Involucral bracts densely and firmly appressed, imbricate, 7—8 rowed, lacking spines, coriaceous, yellowish, almost woolly, later glabrous; outer bracts ovate, very short, obtuse, middle ones larger, oblong or oblong-lan- ceolate, with very short, straight, often deciduous cusp, innermost elon- gate, lanceolate-linear, pappery or scarious, undivided or entire, sca- brous along margin. Receptacle setose. Corolla purple, about 22 mm long; corolla tube narrow, about 9 mm long, limb about 12 mm long with linear lobes 5 mm long. Anther tube slightly shorter than corolla, free in upper part, with inconspicuous basal appendage. Style almost as long as corolla, short-bilobate. Achenes 5—6 mm long and 1.5 mm wide, almost triquetrous, narrowed toward base, slightly sulcate and with dark streaks, smooth; pappus about 10 mm long, yellowish, 1.5 times as long as achenes. Flowering July to October. (Plate XVI, Big. 3), Dry rubbly and stony slopes, in low- and mid-montane zones, to 1,800 m.—Caucasus: Southern Transcaucasia (Nakhichevan ASSR). General distribution: Iran. Described from Nakhichevan. Type in Leningrad. Series 7. Quinquefoliae Boriss.—Leaves compound, with 1-2 pairs, ovate or oblong, serrate, acuminate, wide lobes, and larger terminal lobe. 28. S. quinquefolia MB. Fl. taur.-cauc. II (1808) 264, III, 544; DC. Prodr. VI, 668; Ldb. FI. Ross. II, 2, 756; Boiss. Fl. or. III, 587; Schmalh. Fl. II, 112; Grossh. Fl. Kavk. IV, 193; Ter-Khachaturova in Fl. Gruzii, VIII, 525; Agadj. in Fl. Azerb. VIII, 423.—Klasea quinquefolia (MB.) Cass. in Dict. Sc. nat. XXXV (1825) 173.—Ie.: Bot. Mag. tab. 1871.—Exs.: GRF No. 465; Fl. cauc. exs. No. 146. Perennial. Plant branched, 50-120 cm, sometimes to 2 m high, subglabrous or weakly pubescent. Leaves compound, unpaired-pinnate, 1—2 paired, with 3-5 lobes often simple in upper part; leaf lobes ovate- oblong or oblong-lanceolate; lower leaves petiolate, with ovate or oblong, serrate, acuminate, broad lobes, narrowed toward base; lateral lobes often decurrent, terminal lobe larger; leaves on branches re- duced, fewer, petiolate, elliptical to oblong-lanceolate. Capitula on more or less leafy peduncles, numerous, medium-sized, 1.0—1.5 cm in dia, ovate. Involucral bracts numerous, firmly imbricate, membranous, cuspidate, pubescent and finely ciliate on margin, with dark stripes on outside; outer bracts ovate, middle ones lanceolate, inner ones linear, pubescent at tip, scarious, elongated, almost toothed. Corolla pink, about 17 mm long; corolla tube narrow, 9 mm long; limb campanulate, 295 292 Plate XVI. 1—Serratula sogdiana Bge.; 2—S. cardunculus (Pall.) Schischk.; 3—S. coriacea Fisch. and Mey. 296 293 8-10 mm long, with linear lobes 5 mm long. Anther tube almost as long as corolla; filaments papillose. Basal appendage of anthers less than 1 mm long, anthers free at tips. Style longer than corolla, bilobed, with short lobes. Achenes oblong, about 5 mm long, compressed, slightly narrowed toward base, slightly constricted at apex, sulcate; pappus reddish, deciduous, slightly longer than achenes, about 8 mm long, shorter than corolla tube, with somewhat scabrous hairs. Flow- ering June to October. (Plate XIV, Fig. 1). Mountain forests edges, scrubs, 300—1,000 m to mid-montane zone.—Caucasus: Ciscaucasia, Eastern, Western, and Southern Transcaucasia. General distribution: Iran, Balkans-Asia Minor. De- scribed from Caucasus. Type in Leningrad. Section 5. Iljinia Boriss. in Addenda XX VII, 606.—Low stemless plants or stems low, 2—20 cm high, but then leaves undivided; leaves in rosettes, basal and cauline leaves lyrate, more or less long-petiolate, with 2—5 pairs of lobes. Capitula 1.5—3.5 cm in dia, sessile or on more or less attenuate peduncles; involucral bracts 7-9 rowed. Type of section: S. lyratifolia Schrenk. Series 1. Lyratifoliae Boriss——Plants stemless or with very low stem to 2-8 cm high; basal and cauline leaves lyrate, pinnatifid or pinnately incised, 5-6 cm long and 1—2 cm wide, with 2-5 pairs of lobes, scabrous from short pubescence; capitula 1.5—2 cm in dia. 29. S. lyratifolia Schrenk. in Fisch. and Mey. Enum. pl. nov. I (1841) 45; Ldb. Fl. Ross. Ii, 2, 756; O. and B. Fedtsch. Perech. Rast. Turk. IV, 291; Czerneva in Fl. Uzbek. VI, 390. Perennial. Roots strong, string-like, brown. Rhizome oblique, in upper part covered with fibrous remnants of old leaves, petioles. Plants stemless or stem prostrate on debris in underground part, solitary, very short, 2-8 cm high, simple, considerably shorter than leaves, one- headed. Basal and cauline leaves 5—6 cm long and 1.5—2 cm wide, long-petiolate, lyrate, pinnatifid or pinnately incised with 2—S pairs of lobes, scabrous from scattered short bristle-like hairs; terminal lobe of leaf rounded or ovate, 3—4 cm long and 2.5—3 cm wide, occasionally narrower, lateral lobes considerably smaller than terminal one, coales- cent at base, broadly triangular, ovate, obovate, or oblong, all sharply serrate-toothed, teeth with white cartilaginous cusp; uppermost leaf at base of capitulum lanceolate, toothed. Capitula medium-sized, 1.5—2 cm in dia. Outer involucral bracts broadly ovate-triangular, obtuse, green; inner bracts elongate to lanceolate, almost woolley, at tip with herbaceous, nonspinous, thin, almost filiform, often contorted, straight, 2977 294 often deflected and bent apical appendage. Receptacle setose. Corolla purple, 20-25 cm long; corolla tube narrow, 10 mm long; limb 10-12 mm long, with narrow, almost filiform, acute, purple-violet lobes 5 mm long. Anthers shorter than corolla, connate in tube, free above, with linear, short, basal appendage; filaments papillose. Style slightly exserted from corolla or as long, with 2 very short branches. Achenes 5—6 mm long and about 2 mm wide, oblong, slightly narrowed toward tip but more toward base, grayish-brownish or almost black, smooth; pappus somewhat shorter than corolla, long, many-rowed, with hairs 17-20 mm long, yellowish. Flowering June to August (September). (Plate XV, Fig. 3). Dry rubble slopes, debris, and stony and rubbly peaks, to 3,500 m, to alpine zone of mountains.—Soviet Central Asia: Lake Balkhash Region (Chu-Ili Mountains), Dzhungaria-Tarbagatai, Tien Shan (Trans- Ili Alatau, Kirghiz Range), Pamiro-Alai (Turkestan and Hissar ranges). General distribution: Dzhungaria-Kashgaria (Kitakh-Sairam Moun- tains). Described from Trans-Ili Alatau Range in Kaktau region. Type in Leningrad. 30. S. tianschanica Sap. and Nik. in Bot. Mat. Gerb. Glavn. Bot. Sada, VI, 2 (1926) 31. Perennial. Roots string-like, thickened. Plants stemless, 4-6 cm high, with fibrous remnants of old leaves at base. Leaves in rosettes, thickish, oblong-lanceolate, 16 cm long, 1—1.5 cm wide, pinnately di- vided into irregular lobes, finely hairy on veins or all over on both sides, on wide petiole winged like rachis; leaf lobes almost opposite or alternate, ovate-triangular or roundish, obtuse, sometimes with very short, cartilaginous cusp, with sparse, unequal, irregular, short, acute teeth; terminal lobe similar to lateral or slightly larger, obtuse or sub- acute. Capitula solitary, in leaf rosette. Involucres ovate-cylindrical, 1.5—2 cm in dia; involucral bracts imbricate, 4—5-rowed, greenish, nar- rowly brownish-blackish on margin at a tip; outer bracts ovate, broad, acute or short-cuspidate, inner bracts oblong to lanceolate and linearly lanceolate, with scarious brownish-yellowish appendage, largest on innermost bracts. Receptacle alveolar, bristly on edges of alveoles. Corolla pinkish-violet, about 22 mm long; corolla tube 10—13 mm long; limb about 10 mm long, parted up to half into 5 narrow, linear, acute, violet lobes about 5 mm long. Anthers connate in middle, free above, with short basal appendage. Style exserted from corolla to 4-5 mm, with lobes 1.5—-2.0 mm long. Achenes oblong or oval, almost tetraquetrous, slightly compressed, 4.5 mm long and about 2 mm wide, glabrous; pappus slightly shorter than corolla tube, yellowish with scabrous, longer inner bristles. Flowering July to August. 298 300 295 Steep slopes, mountain meadows, and large bouldery, turfy mo- raines.—Soviet Central Asia: Tien Shan (eastern part), Dzhungaria- Tarbagatai (?). General distribution: Dzhungaria-Kashgaria. Described from Inylchek in Tien Shan. Type in Leningrad. Series 2. Turkestanicae Boriss.—Stems 10-20 cm high; leaves oblong, undivided, sometimes with 1-2 lobes in lower part, glabrous, 8—15 cm long and 2.5—4 cm wide; capitula (1.5)2—3 cm in dia, sessile or on more or less distinct peduncles. 31. S. modesti Boriss. in Addenda XXVII, 606. Perennial. Root string-like, brown. Rhizome strong, long; stem 10-20 cm high, simple, with remnants of old leaves at base. Basal leaves glabrous, 8—15 cm long and 2.5—4.0 cm wide, on (1)5—10 cm long petiole, sharply serrate, sometimes with 1-2 pairs of basal lobes, thick and coriaceous, subacute, with short, soft and thin cusp. Capitula 2.5—3.5 cm in dia, pedunculate, peduncles as long as leaves. Involu- cres 2—3 cm in dia, basally flatish, not narrowed, uniformly broad in upper part; involucral bracts 7—9-rowed, outer and middle ones ovate, gradually becoming longer, with narrow black border, green, with deciduous cusps; inner bracts lanceolate, with membranous append- age. Receptacle densely setose, bristles smooth, 5 mm long. Corolla purple, 25 mm long; corolla tube very narrow, about 13 mm long; limb narrow, campanulate, 13 mm long with linear lobes 7 mm long. Anther tube slightly shorter than corolla, free obtuse lobes in upper part, with linear, very short basal appendage; filaments free, sparsely papillose. Style slightly exserted from corolla, with 2 short branches. Achenes ovate, 5 mm long and 3 mm wide, almost smooth, very finely sulcate; pappus about 16 mm long, dense, yellowish, pappus hairs very short, appressedly plumose. Flowering June to July. Dry stony slopes and hills—Soviet Central Asia: Dzhungaria- Tarbagatai. Endemic. Described from Dzhungarian Alatau. Type in Leningrad. Note.- It is distinguished from S. lyratifolia Schrenk by the pres- ence of a developed stem, undivided leaves, and larger capitula and florets. Section 6. Leuzeopsis Boriss. in Addenda XXVII, 607.—Plants monoecious; leaves coriaceous, large, undivided or pinnately lobed and with prominent veins mostly in lower part, obtuse, and coarsely crenate-toothed. Capitula large, 3—4 cm in dia, subglobose, with bi- sexual florets; involucral bracts coriaceous, glabrous, with deflected, spinous appendage 5—6 mm long; achenes ovate, abruptly narrowed toward base; pappus 11-15 mm long, 2 times as long as achenes. 2995 296 Plate XVII. 1—Synurus deitoides (Ait.) Nakai; 2—Serratula radiata (Waldst. and Kit.) MB. —_ 297 Type of section: S. serratuloides (Fisch. and Mey.) Takht. A monotypic section. 32. S. serratuloides (Fish. and Mey.) Takht. in Takht. and Fed. Fl. Yerevana (1945) 323; Grossh. Opred. Rast. Kavk. 488; Agadzh. in Fl. Azerb. VIII, 428.—Leuzea serratuloides Fisch. and Mey. in DC. Prodr. VI (1837) 666; Grossh. Fl. Kavk. IV, 197—Phaeopappus freynii Hausskn. and Sint. in herb.—Ic.: Agadj. loc. cit. Plate XLII. Perennial. Root strong, woody. Stems angular or more or less straight, 10-30 cm high, usually short, less often somewhat surpassing lower leaves, strong, sulcate, weakly woolly pubescent. Leaves large, undivided or pinnately iobed, oblong, to 20 cm long and 4—10 cm wide, petiolate, thick, coriaceous, with prominent veins, subglabrous above, thinly hairy beneath mostly along veins, terminal lobe ovate or ovate-oblong, lateral lobes 2—3 pairs, smaller, decurrent on petioles, oblong, like terminal lobe irregularly crenate-toothed. Capitula solitary, terminal, almost globose or subglobose, large, 3—4 cm in dia. Involucral bracts 5—6-rowed; glabrous, coriaceous, narrowly membra- nous along margin, with dark green, longitudinal veins, subulate, often deflected spinous, apical appendage 5—6 mm long, 1/3-—1/2 as long as involucral bracts; outer bracts ovate, middle ones oblong, inner ones lanceolately-linear, acuminate. Receptacle setose. Corolla pink or light- pink, 30-33 mm long; corolla tube narrow, 16 mm long, clavately bulged at base; limb broad, campanulate, about 16 mm long, with linear lobes 7 mm long. Anther tube as long as or slightly shorter than corolla, about 11 mm long, with free obtuse lobes at tip, basal append- age about 0.5 mm long; filaments glabrous. Style about 36 mm long, exserted from corolla, usually short-bilobate. Achenes greatly compressed, ovate or oblong-ovate, 6-7 mm long and 3—5 mm wide, narrowed above, at base subacute, brown, weakly sulcate or smooth; pappus longer than achenes, 11-15 mm long, dense, white, of flat, plumose bristles at base connate in ring. Flowering May to July; fruit- ing July to August. Dry rubbly and rocky slopes, talus, in mid-montane zone, 1,500— 1,800 m.—Caucasus: Southern Transcaucasia (Nakhichevan ASSR, Armenia). Endemic. May be present in the neighboring parts of Iran. Described from southern Transcaucasia (Koshadar River Valley). Type in Leningrad. Section 7. Demetria Boriss. in Addenda XXVII, 607.—Plants mo- noecious; stems broad-winged; leaves large, decurrent, shallow sinu- ate-toothed and spinose. Capitula large, 2.5—3.5 cm in dia, globose or ovate; involucral bracts coriaceous, subglabrous, spines 7-10 mm long, straight; limb 1.5 times as long as corolla tube. 298 Type of section: S. caucasica Boiss. 33. S. caucasica Boiss. Fl. or. III (1875) 590; Grossh. Fl. Kavk. IV, 194; Agadj. in Fl. Azerb. VIII, 426.—S. cichoracea (L.) DC. Prodr. VI (1837).670 p. p.; Ldb. Fl. Ross. II, 2, 760.—Centaurea cichoracea Stev. in Mém. Soc. Nat. Mosc. IV (1813) 66; MB. FI. taur.-cauc. III, 595, non L. Perennial. Plant glabrous or subglabrous, 20-50 cm high. Stem robust, 4-5 mm thick, angular, simple, deeply sulcate, uniformly densely leafy and broad-winged, one-headed. Leaves 5—15 cm long and 2—7 cm wide, lanceolate or oblong, decurrent, irregularly finely, spinous-sinu- ate-toothed. Capitula 2.5—3.5 cm in dia, globose or ovate. Involucral bracts imbricate, 6-rowed; coriaceous, subglabrous, lanceolate or oblong, gradually attenuate above into almost straight, deflected spine 7-10 mm long; inner involucral bracts elongate, 2.5—3.0 cm long, softer and acuminate. Corolla purple or pinkish-purple, about 25 mm long; corolla tube narrow, about 10 mm long; limb narrowly campanulate, about 15 mm long, with linear lobes 9 mm long. Anther tube shorter than corolla, with free, subacute apical lobes, anthers with filiform, basal appendage about 1 mm long; filaments densely papillose. Style slightly exserted from corolla, short-pubescent (visible under lens!) with subacute lobes about 0.5 mm long. Achenes oblong, 9 mm long and 3 mm wide, finely sulcate above, brownish; pappus of yellowish, lax and short-plumose hairs. Flowering July to August. Dry slopes in subalpine mountain zone.—Caucasus: Dagestan, Eastern Transcaucasia (Azerbaidzhan). Endemic. Described from Caucasus. Type in Geneva. Note. It is distinguished from S. cichoracea (L.) DC. s. str. (Italy) by a shorter, broadly (and not narrowly) winged stem and thinner, almost straight (and not recurved) spines on the involucral bracts. With its large capitula and involucres, S. caucasica is closer to S. mucronata Desf. and S. flavescens Poir., differing from the latter by having de- current leaves. . GENUS 1608. Synurus Ijin" ” Iljin in Bot. Mag. Gerb. Glavn. Bot. Sada, VI, 2 (1926) 35. Capitula homogamous, large, few at tips of long branches. Involu- cral bracts arachnoid-hairy, stiff, triangular-lanceolate to linear, cuspi- 'Treatment by A.G. Borissova. "From the Greek words syn—together, and ura—tail; so named for the shape of the anther appendages. 299 302 date and acuminate, often with recurved, nodding long spine. Recep- tacle covered with smooth, more or less long bristles. Florets all tubu- lar, bisexual; corolla regular, 5-lobed. Anthers like filaments free, joined only by appendages, fused on inner side with filaments, leaving gap, or forming tube around filaments, sometimes fused with tube of neigh- boring anther. Style bifid, with short, more or less coalesced, obtuse, flat and glabrous lobes on inner side. Achenes slightly compressed, glabrous, with crown of apical teeth, surrounding base of pappus; pappus of scabrous unequal bristles, connate in ring. Large perennials with nodding, long, thin arachnoid-hairy branches; leaves large, long-peti- olate, deltoid-ovate with cordate base. Type of genus: S. deltoides (Ait.) Nakai. A monotypic genus occurring in China, Japan, Korean Peninsula, Mongolia, and the Far East. 1. S. deltoides (Ait.) Nakai, Kéry6 Sikenrin no Ippan (1927) 64; Kitagawa in Rep. first Sc. exp. Manch. IV, 2, 41, in nota; Lineam. FI. Mansh. 470; Kitam. in Mem. Coll. Sc. Kyoto Imp. Univ. ser. B, XIII 25 (Comp. Jap. I); Grubov, Konsp. Fl. Mong. 279.—S. atriplicifolius (Trev.) [jin in Bot. Mat. Gerb. Glavn. Bot. Sada, VI, 2 (1926) 35; Popov, Fl. Sredn. Sib. Il, 766.—S. hondae Kitagawa in Rep. fist Sc. exp. Manch. IV, 2 (1935) 40.—S. atriplicifolius var. hondae (Kitagawa) Kitagawa in Rep. Inst. Sc. Res. Manch. 1, 8 (1937) 324.—Onopordon deltoides Ait. Hot. Kew. III (1789) 146.—Cirsium ficifolium Fisch. in Mem. Soc. Nat. Mosc. II (1812) 69.—Carduus atriplicifolius Trev. Ind. Sem. hort. Vratisl. (1820); in Nova Acta phys.-med. Acad. Leopold. -Carol. Nat. Curios. XIII, 1 (1826) 198.—Silybum atriplicifolium (Trev.) Fisch. Ind. pl. Hort. Petrop. (1824) 64.—Rhaponticum atriplicifolium (Trev.) DC. Prodr. VI (1837) 663; Turcz. Fl. baic.-dahur. II, 1, 132; Regel, Tent. Fl. Ussur. 96; Fr. Schmidt, Fl. Amur. et Sachal. 52; Maxim. Prim. Fl. Amur. 175; Korsh. in Tr. Peterb. Bot. Sada, XII, 2, 361.— Stemmacantha ficifolia (Fisch.) Turcz. in DC. 1. c. pro syn.— Stephanocoma atriplicifolia (Trev.) Turcz. ex Ldb. Fl. Ross. li, 2 (1846) 751, pro syn.—Serratula atriplicifolia (Trev.) Benth. in Benth. and Hook. f. Gen. pl. II (1873) 475; Kom. Fl. Manchzh. II, 758; Kom. and Alis. Opred. Rast. Dalnevost. Kr. II, 1086; Miyabe and Miyake, Fl. Saghal, 281.—S. pungens Franch. and Sav. Enum. pl. Jap. II (1879) 416.—S. deltoides (Ait.) Makino in Tokyo Bot. Mag. XXIV (1910) 247.—S. atriplicifolia var. inciso-lobata Miyabe and Miyake, 1. c.— Centaurea atriplicifolia (Trev.) Matsum. Ind. pl. Jap. II (1912) 667.— Ic.: Ijin, op. cit. 35 (details of flower); Makino in Jap. Bot. Illustr. Album.(1933) 1368; Kitagawa in Rep. first Sc. exp. Manch. IV, 2 (1935) tab. XI. 30 Ww 300 Perennial. Rhizome thick, short. Plants to 1 m high, branched with rather strong stem and nodding, long, finely arachnoid-hairy branches. Basal and lower cauline leaves long-petiolate, deltoid-ovate, to 20 cm long, cordate, acute, toothed-lobed or sinuate-lobed, glabrous and green above, white-tomentose beneath; upper cauline leaves oblong-ovate, short-petiolate, almost entire. Capitula large, 3-4 cm in dia, nodding at tips of long lateral branches. Involucral bracts lanceolate to linear, weakly or densely arachnoid-hairy, gray, with dark brown, long, ap- pressed or recurved spines. Florets dark violet, numerous, bisexual, all tubular; corolla 27-30 mm long, almost regular, with narrow tubular part about 8 mm long, gradually broadened part about 15 mm long, with 5 linear lobes about 3 mm long, exserted from corolla; anthers about 7 mm long, free, joined only by appendages about 3.5 mm long. Style exserted from corolla, with short branches or almost equal to stamens. Achenes 7 mm long, and about 2.5 mm wide, ovate-oblong, tetraquetrous, slightly compressed, finely sulcate and dark-spotted, obliquely truncate at both ends, tip small-toothed; pappus of unequal, scabrous bristles 8—9 mm long. Flowering July to August. (Plate XVII, Fig. 1). Open forests, birch groves, forest edges, scrubs and herb mead- ows.—Eastern Siberia: Dauria; Far East: Zeya-Bureya, Ussuri, Sakhalin. General distribution: Mongolia, Japan, China, Korean Pen- insula. Described from cultivated plants. Type in London. GENUS 1609. Syreitschikovia PAVL.' ” Pavl. in Fedde, Repert. XXXI (1933) 192. All florets in capitulum tubular, bisexual; corolla lilac with very narrow tube and 5-parted limb. Anther tube with oblong, slightly acumi- nate, apical appendages and bifid, bristle-like, basal appendages; fila- ments glabrous. Style thickened at base of branches, with tuft of short bristles, with numerous papillae from basal thickening to branches; style branches appressed, slightly divergent only above. Pappus deciduous, with unequal toothed bristles, connate in ring, outer ones hair-like, inner ones thickened, acuminate; achenes oblong, compressed, glabrous, hilum basal, with toothed crown above enclosing pappophorus with nectary. Type of genus: S. tenuifolia (Bong.) Pavl. The genus includes two species, both found in the USSR. 'Treatment by M.M. Iljin and G.A. Semidel. 2Named in honor of D.P. Syreistschikov, author of “JIl. Fl. Mosk. Gub.” [Illus- trated Flora of Moscow Province]. 304 301 1. Leaves narrow, linear, spinose on margin and in lower part. Involu- cral bracts appressed, cuspidate, sometimes somewhat spreading...... ee Wkd Moka dite. ARMac Rae 1. S. tenuifolia (Bong.) Pavl. + Leaves oblong-ovate or oblong; cauline leaves sessile, semiamp- lexicaul, auriculate, with numerous spinules along margin. Involu- cral bracts with long arcuate apical prickles ............ ee eNO POMU OURAN WA AUN Te AO tN, OB 2. S. spinulosa (Franch.) Pavl. 1. S. tenuifolia (Bong.) Pavl. in Fedde, Repert. XXXI (1933) 192; Kryl. Fl. Zap. Sib. XI, 2934.—S. tenuis Botsch. in herb.—Serratula tenuifolia Bong. in Bull. Acad. Sc. Pétersb. VIII (1841) 340; Ldb. FI. Ross. II, 760; O. and B. Fedtsch. Perech. Rast. Turk. IV, 293.—Jurinea tenuis Bge. in Flora, XXIV (1841) 158.—Ic.: Bong. and Mey. in Mém. Acad. Sc. Pétersb. ser. VI, VI, 2, tab. 10 (1845). Perennial. Plants 5—40 cm high, densely caespitose with branched rhizome, densely covered with dark brown, scarious scales and large number of adventitious roots. Stems mostly several, straight, simple, weakly ribbed, spinose-arachnoid-hairy, with readily rubbed-off pu- bescence. Leaves linear, narrowly oblong, entire, acuminate, slightly narrowed toward base, at base along margin with tiny spine-like cilia, bright green above, somewhat lustrous, sometimes weakly arachnoid- hairy, white-tomentose beneath, 1.5—10.0 (13.0) cm long, 1-3 mm wide, with single longitudinal vein. Capitula solitary, with many florets. Involucres cylindrical or goblet-shaped, 8-15 mm long, green, gla- brous, weakly arachnoid-hairy at base; involucral bracts appressed, but with recurved spine-like tips; outer and middle bracts carinate, lan- ceolate, inner ones lanceolately acuminate, straight. Corolla 9-15 mm long; corolla tube very narrow, 5—7 mm long; limb with 5 linear lobes 3—4 mm long. Anthers purple, with acuminate apical appendage, basal appendages bifid into filiform bristles 2 mm long; filaments glabrous. Style with divergent branches. Pappus slightly shorter than corolla, 8— 12 mm long, its bristles toothed, unequal, innermost thickened, broad- ened upward, and long apical, bristles weakly connate in ring, decidu- ous on maturation; achenes glabrous, narrowly-oblong, dull, longitudinally striped, with dark purple spots, hilum basal, scarious- toothed crown at apex, enclosing cylindrical pappophorus with nectary in its center; mature achenes 6-7 mm long. Flowering June to July; fruiting July to September. High-mountain stony, steppe slopes.—Soviet Central Asia: Tien Shan, Dzhungaria-Tarbagatai. General distribution: western China. Described from Dashildag Mountains. Type in Leningrad. 305 302 2. S. spinulosa (Franch.) Pavl. in Fedde, Repert. XX XI (1933) 193.—Serratula spinulosa Franch. Pl]. du Turkest. Mission Capus (1883) 107; O. and B. Fedtsch. Perech. Rast. Turk. IV (1911) 295. Perennial. Plants 15—35 cm high. Stems angular-sulcate, straight, with spiny, arachnoid, deciduous hairs, usually few, occasionally branched from base, covered with brown scales, leafy to capitula. Leaves oblong-ovate or oblong, basal leaves gradually narrowed into petiole; cauline leaves sessile, semiamplexicaul, auriculate; all leaves undivided, subentire, margin slightly undulate with frequent spines, acuminate, gradually attenuate into spine, green and slightly lustrous above, at places with thin arachnoid pubescence, later glabrous, white-tomentose beneath with prominent ochreous midrib. Capitula solitary, urn shaped, with many florets, 25-40 mm in dia. Involucres about 20 mm long, thinly arachnoid-hairy; involucral bracts linear, with very prominent vein, gradually attenuate into thin spine, arcuately recurved. Corolla dark lilac when dry, with scattered glands, 15-18 mm long, gradually narrowed below over 7—8 mm; limb lobes 3—4 mm long. Anthers lilac- purple, their basal appendages bifid into filiform bristles; filaments glabrous. Style with short, slightly divergent branches; stigma entirely covered with numerous papillae. Pappus deciduous, 14—16 mm long, smoky; pappus bristles unequal, all connate in ring, small-toothed, outer ones filiform, inner ones thick, broadened above and terminating in long cusp; achenes glabrous, 6—8 mm long, and hilum basal, narrow- oblong, compressed, 4-angled, longitudinally striate, gray, with longi- tudinal violet spots, with irregularly toothed crown enclosing raised annular pappophorus with nectary in its center. Flowering July; fruiting August to September. High-mountain arboreal-shrub and subalpine zones.—Soviet Cen- tral Asia: Tien Shan (western). Endemic. Described from Ona Ulgan River Valley, near Chirchik, at 1,800 m. Type in Paris. GENUS 1610. Schumeria Iljin’? Iljin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XX (1960) 363.—Microlophopsis Czer. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XX (1960) 483. Capitula homogamous, few, medium-sized, with many florets, soli- tary, at tips of stems and their lateral branches. Involucres oblong, 306 about 1.5 cm wide; involucral bracts compactly imbricate, coriaceous, 'Treatment by A.G. Borissova. *Named after the ancient Sumeria lying between the Tigris and Euphrates rivers. 303 lacking appendage, with short, appressed, pointed tip, scabrous in up- per part, glabrous in lower, oblong-triangular. Receptacle densely cov- ered with long, lustrous, and smooth bristles. Florets all bisexual, simi- lar, tubular, with long tube, yellow, many-rowed; corolla limb with 5 equal, linear lobes. Anthers with subobtuse, rounded, apical append- age; basal appendages of anthers short, cuneate, almost undivided and scarious; filaments flattened, sparsely covered with papillae. Style gla- brous, exserted from corolla, with long lobes, on outer side covered with very short hairs. Achenes oblong, 8-10 mm long compressed, slightly sulcate or smooth, glabrous, hilum oblique; pappus large, 2 times as long as achenes, with several rows of deciduous bristles; pappus bristles barbate-toothed or plumose-barbate, gradually longer from outer to inner. Perennial or biennial glabrous herbs; leaves sessile, amplexicaul, short-decurrent, undivided and entire, sometimes lower and basal leaves pinnately incised. Type of genus: S: cerinthifolia (Sibth. and Sm.) I}jin. About 6 species, distributed in West Asia and Soviet Central Asia. Note. The genus Schumeria Iljin is subdivided into two sections: Schumeria (= sect. Oroboidea Ijin loc. cit.) and Serratulopsis jin (op. cit., 364). The genus Schumeria is similar to the genus Serratula L., from which it is distinguished by yellow florets, coriaceous short-acuminate involucral bracts, larger achenes, amplexicaul, undivided and entire cauline leaves, the lower sometimes pinnately incised, and by the habit of the plant. It approaches Serratula by the presence of an oblique lateral hilum and a pappus of bristles and not scales. The species of the genus Schumeria are close to those of the genus Microlophus Cass., particularly to M. behen (L.) Takht. (= M. alatus (Lam.) Cass. = Cen- taurea behen L. = C. alata Lam. = Piptoceras behen (L.) Cass.). De Candolle (Prodr. VI [1837] 567) referred the genera Microlophus Cass. and Piptoceras Cass. to section Microlophus (Cass.) DC. of the genus Centaurea L., where Centaurea cerinthifolia Sibth. and Sm. (= C. behen Lam. non L.) appears, which was referred by M.M. Iljin to the genus Schumeria. Additional material and further studies are needed to reach any definite conclusions. Possibly, the species of the genus Schumeria Ijin should be referred to the genus Microlophus Cass. 1. Plants glabrous; involucral bracts glabrous, only young plants SParsely Walry. ...so2.s00:s2.<0 ARO AN EY. 1. S. latifolia (Boiss.) Iljin + Plants scabrous from short barbate hairs; leaves with long, soft white hairs on veins mostly beneath, later becoming glabrous ........ Re ne Gh: Cee at on) OS aoa 2. S. litwinowii (Iljin) [jin 307 304 Section 1. Serratulopsis Iljin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XX (1960) 364.—All leaves undivided and entire; pappus plumose-barbate. Type of section: S. /atifolia (Boiss.) Ijin. 1. S. latifolia (Boiss.) Ijin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XX (1960) 364; Iljin in Fl. Turkm. VII, 254.—Serratula latifolia Boiss. Diagn. pl. or. ser. I, 10 (1849) 96; FI. or. III, 586; O. and B. Fedtsch. Perech. Rast. Turk. IV, 211; B. Fedtsch. Rast. Turk. 756.—Centaurea plumosa Aitch. and Hemsl. in Trans. Linn. Soc. ser. 2, Il (1888) 81; Parsa, Fl. Iran, III, 686, non Kern. (1872).— Microlophopsis plumosa (Aitch. and Hemsl.) Czer. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XX (1960) 488. Perennial. Plants glabrous, 25—90 cm high. Root long, strong, woody. Stems elongate, cylindrical, simple or weakly branched, with long, straight branches, whitish, with fibrous remnants of old leaves at base. Leaves coriaceous, lustrous, pale green, with thick, whitish mid- rib and reticulate venation, undivided; basal leaves 7-10 cm wide, 10— 13 cm long, ovate, on thick, short, grooved petioles, scarious at base along margin, amplexicaul; middle leaves sessile, oblong, subacute, with acute, cordate auricles at base, short-decurrent; leaves in lower part of stem and on branches reduced, erect, linear. Capitula solitary, about 3.5 cm long, on long peduncles. Involucres about 2.5 cm long, oblong-cylindrical, almost constricted at apex, lustrous, entirely gla- brous; involucral bracts firmly appressed, 6-rowed, lacking spines, almost lacking distinct veins, weakly sulcate, coriaceous, yellowish; outermost bracts ovate, with small cusp, innermost elliptical to linear, acuminate, gradually broadened. Receptacle densely covered with long, smooth bristles, almost as long as pappus. Florets lemon-yellow, all equal, bisexual, 1/3 longer than involucre; corolla about 33 mm long; corolla tube very narrow, about 18 mm long; limb narrowly campanu- late, about 14 mm long, with linear lobes about 6 mm long. Anthers connate in tube, slightly exserted from corolla with free, obtuse, apical appendage and very short, linear basal appendage about 0.5 mm long; filaments subglabrous. Style slightly exserted from corolla, 2-fid, with obtuse, oblong, grooved branches. Achenes somewhat compressed, about 6 mm long, sulcate; pappus about 15 mm long, with white, sordid-red, or yellowish, short-plumose hairs, basally connate in de- ciduous ring. Flowering May to August. Rubbly and loamy mountain slopes and foothills, mountain mead- ows.—Soviet Central Asia: mountainous Turkmenia (primarily Central Kopetdag). General distribution: Iran (Khorasan). Described from Iran. Type in Geneva. 308 305 2. S. litwinowii (Ijin) Iljin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XX (1960) 364; Iljin in Fl. Turkm. VII, 254.—Serratula litwinowii Iljin in Bot. Mat. Gerb. Glavn. Bot. Sada, V, 7 (1924) 112. Perennial. Root robust, woody. Stems with few branches, 25—70 cm high, sulcate, scatteredly short-hairy. Leaves oblong, undivided and entire, scatteredly hairy on both sides, mostly woolly beneath, along veins; with deciduous, quite long hairs; basal leaves petiolate, oblong, narrowed toward base; cauline leaves decurrent, upper ones reduced, with short-decurrent auricles. Capitula mostly large, 4.0—4.5 cm long, about 3 cm in dia at flowering. Involucral bracts yellowish- greenish; outer ones deltoid-ovate, glabrous or subglabrous, cuspidate; middle ones ovate-oblong; inner ones oblong-linear, acuminate, ap- pressed above, usually with short stiff hairs at tip. Receptacle setose, with long, smooth bristles. Corolla yellow, 30-38 mm long, corolla tube narrow, about 22 mm long; limb 8—10 mm long. Basal append- ages of anther short, 1-2 mm long, almost undivided; filaments some- what flat, finely papillose (visible under a hand lens!). Achenes com- pressed, smooth, glabrous, 8-9 mm long; pappus deciduous, plumose, with unequal bristle. Flowering June to July. Loamy mountain slopes in steppe zone, mountain meadows.— Soviet Central Asia: mountainous Turkmenia. Endemic. Described from Turkmenia. Type in Leningrad. Note. This species is similar to S. latifolia (Boiss.) Ijin, differing by having pubescence and larger capitula and achenes. GENUS 1611. Rhaponticum Adans.':? Adans. Fam. II (1763) 117.—Fornicium Cass. in Bull. Soc. Philom. Paris. (1819) 93 and in Dict. Sc. nat. XVII (1820) 249. Involucre many-rowed, imbricate; bracts coriaceous, yellowish- green at beginning of flowering, weakly stramineous outside in lower part, very strongly so inside, lustrous, with dry, papery, apical append- ages; outer involucral bracts very short, obtuse, ovate, later gradually elongating, innermost narrow, almost as long as capitulum; outer and middle bracts with broad and oval appendages, often fan-like, lacerate 'Treatment by Yu.D. Soskov. *From the Greek word Rha and Latin Pontus (Black Sea and its coast); translated into Russian it means Black Sea rhubarb. This name was already given by pre-Lin- naean systematists to one of the species of this genus, whose rhizome was used as a laxative and often substituted for rhubarb. In turn, the name rhubarb, Rheum (from the Greek word Rha), also originates from Rha—the ancient name of the Volga. 309 306 in some sections, innermost bracts with shorter, lanceolate or linear, always undivided appendages. Receptacle somewhat bulged, bristly. Florets all bisexual, similar, violet-purple, less often yellow R. integrifolium Winkl.), with tubular corolla, broadened in upper part, 5-fid into linear lobes. Stamen fila- ments covered with tiny papillae; anthers with small, broad, obtuse, continuous, often unequal basal appendages. Stigma bilobed, ciliate, with tuft of long, tasseled cilia at base, its branches somewhat diver- gent, obtuse. Pappus always longer than achene 2-many-rowed, usually free, of thick, barbate or short-plumose bristles, basally connate in ring, less often free; hairs on bristles not more than 10 times as long as thickness of bristles, bristles of inner row thicker and longer than those of outer rows; achenes glabrous, laterally compressed, oblong, rugose, more or less ribbed and tetraquetrous, basal hilum oblique. Rhizomatous herbs with simple, finely ribbed stem, at apex somewhat thickened, terminating into large, almost globose capitula. R. nanum Lipsky and R. namanganicum Iljin are stemless. In some species stem some times branched. Type of genus: R. scariosum Lam. Note. This genus comprises in all 17 species, distributed in a narrow, almost continuous belt in the northern hemisphere from the Atlantic Ocean to the shores of the Pacific Ocean, with the exception of one species in Australia (Soskov. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XTX [1959] 398). The genus Leuzea DC., repeat- edly reported for the flora of the USSR, is distributed in the Western Mediterranean, where it is represented by only three species. It is distinguished from the genus Rhaponticum by a long-plumose, soft, snow-white, many-rowed pappus, hairs on the bristles 20 and more times longer than the bristle thickness, and achenes that are brownish- barbate, subobtuse, and conically narrowed below. Economic Importance. Many species of Rhaponticum were intro- duced into botanical gardens already in the 18th and 19th centuries. Two of them—R. carthamoides (Willd.) Iljin and R. scariosum Lam.— were specially introduced into cultivation (the latter is now no longer in cultivation). All the species are highly decorative. It would be interesting to test the members of this genus (their seeds and under- ground organs) for their tonic and stimulant properties. 1. Sheaths of dead leaves covering stem base on inside with long silky-white hairs. Involucral bracts with brown spot or border below appendage; bract appendages thin-scarious, transparent. Plants stemless or with stem not higher than 50 cm ..............cccsceeeeseeeees a 310 10. 307 Sheaths of dead leaves covering stem base glabrous on inside. Plants RIStallyZSO2=080 cHathieh Waals SALE BRO RR 6. Pappus reddish, very stiff, about 3 cm long. Plants completely gla- brous; leaves with cartilaginous teeth ................ 8. R. nitidum Fisch. Pappus white, about 1.5 cm long. Plants more or less pubescent ..... Plantsiwith'sten 623 Orem higheh Lee en es 4. Plants stemless or almost stemless, 3—5 cm high .................eee 5. Plants whitish from dense, fine tomentum, 6-15 cm high; leaves Some whatichisped on margin RAL). ESE RE Mire RAP ROR Mate ORS, 10. R. karatavicum Rgl. and Schmalh. Plants more or less green, 15-40 cm high; leaves with scattered crisped hairs, leaf lobes scarious, acuminate or not acuminate ......... ee cca RD ROR, Bk UO RR 9. R. aulieatense Ijin Leaves pinnately divided into toothed lobes 1.0.0.0... ee eeeeceeeeeeeeeees ER RETR NUN ROL EY | MEN Sd BS. 11. R. namanganicum Ijin Leaves pinnately incised into pinnately divided or lobed lobes. Inner involucral bracts violet-purple under violet to reddish-brown appendages at the end of vegetative period .... 12. R. namum Lipsky Florets yellows; pappus persistent, its bristles not connate in a ring atjbaseseaves entirely undividedes!. un... ee Te Db SOM i SORE ON tre U8 4. R. integrifolium Wink] Plorets violet=purple;pappusi free ietere. AP EE ae Te Leaves mostly with undivided lamina ............. eee eeeeseeseeeeeeeees 8. Plants except for uppermost small, toothed leaves, with only pin- nately dividediorincised*leavesw22. i002 LR ER. 9. Basal leaves slightly lyrate. Appendages of involucral bracts yel- lowish-smoky, semitransparent. Plants of mountain valleys ........ MIA APE, AAR ED AT, SR EE pha Mega 3. R. lyratum Winkl. ex Ijin Basal leaves elliptical, acute, undivided or pinnately incised at base into 1-3 pairs of oblong lobes. Appendages of involucral bracts spatulate, with attenuate tip and light-brown band in middle nar- rowed to cusp. Plants of salt-marsh steppes ...........c:.cccccssssesseseseseneeeees ABA CU GIS. Bi vel es ita errs 89 2. R. serratuloides (Georgi) Bobr. Pappus short-plumose, 2-rowed, soft, reddish-brown to creamish. Mesophyllous plant of subalpine and alpine zones, (50)80—150(180) cm high, with horizontal branching rhizome and numerous roots .. dfn PAR BRS 8 ky i) 1. R. carthamoides (Willd.) iin Pappus barbate, 3-rowed, stiff. Distinct xerophyte with grayish- or whitish-tomentose stem (at least in lower paft) ............-.:eeceeeeee 10. Leaves white-tomentose beneath. Plants highly ornamental ........... cRIOLE iste ET RARE RMN IRS tanib le eS 5. R. pulchrum Fisch. and Mey. Leayesmotitomentose beneath \..:..5......05cc5e cas eccssseceeanndeeeeeaaee dott: 311 308 11. Stems 30-100 cm high, 0.5—1.0 cm thick, grayish tomentose below, later becoming glabrous; leaves of vegetative shoots large, 20-40 cm long, 7-10 cm wide, lyrate, with very large terminal lobe; cauline leaves shallowly pinnately divided into 4—6 pairs of oval-ovate OBES): Ng / NY Plate XVIII. 1—Amberboa glauca (Willd.) Grossh.; 2—A. bucharica jin. 331 328 yellow or yellowish-white, peripheral ones smaller (scarcely surpass- ing central florets), often relatively few; corolla 5—10-lobes, more or less hairy in middle. Achenes 4.5—5.0 mm long; pappus slightly shorter than achenes. Flowering May to June. Clayey and stony slopes, gravels, sands, alkali soils—European Part: Lower Volga; Caucasus: Eastern Transcaucasia; Western Sibe- ria: Upper Tobol (southern part), Irtysh (southern part), Altai (south- ern and southwestern foothills); Soviet Central Asia: All regions, but in mountains only in lower montane zone. General distribution: Iran- Afghanistan, Dzhungaria-Kashgaria. Described from western Kazakhstan (between Emba River and Ustyurt). Type in Leningrad. Note. As in the previous species, plants with a well-developed, erect stem (to which belongs the type of the species) predominate in the more northern parts of the range of A. turanica, while plants with the capitulum close to the base on a greatly reduced peduncle, as if lateral in relation to the other branches of the stem, predominate in the more southern and more eastern parts. Series 3. Flaviflorae Tzvel.—Relatively large biennial or annual plants. Involucres 9-24 mm in dia; appendages of inner involucral bracts relatively large, ovate or ovate-lanceolate. Florets yellow, corolla of peripheral florets 8—20-lobed. 6. A. amberboi (L.) Tzvel. comb. nova.—A. odorata (Cass.) DC. a. ambracea (Schkuhr) DC. Prodr. VI (1837) 560.—A. odorata a. floribus flavis Ldb. Fl. Ross. II, 2 (1845) 682.—A. moschata auct. non DC.: Boiss. Fl. or. III (1875) 605 p. p.—A. suaveolens (Willd.) Iljin in Izv. Bot. Sada Akad. Nauk SSSR, XXX, 1-2 (1932) 107; Grossh. Fl. Kavk. IV, 195 and Opred. Rast. Kavk. 489; Iljin in Fl. Turkm. VII, 255.—Centaurea moschata 8. amberboi L. Sp. pl. (1753) 909.—C. amberboi (L.) Lam. Encycl. meth. I (1783) 664.—C. suaveolens Willd. Sp. pl. III, 3 (1803) 2279.—C. ambracea Schkuhr, Bot. Handb. III (1803) 157.—Chryseis odorata Cass. in Dict. Sc. nat. IX (1817) 154.— Ic.: Schukur, loc. cit. tab. CLXI.—Exs.: Sintenis, It. transcasp.-pers. 1900-1901, No. 1616 (sub A. moschata). Annual or biennials. Plants 20-60 cm high. Stem erect, simple or more or less branched, more or less covered with short crisped hairs. Leaves more or less crisped-hairy to subglabrous; basal leaves quite long-petiolate, usually undivided but more or less toothed; middle and upper ones usually sessile or subsessile, more or less pinnately lobed or divided. Capitula solitary at tips of stem and lateral branches, on long leafless peduncles. Involucres broadly ovate, with florets 9-17 mm in dia and 12-17 mm long, covered with very thin arachnoid 332 829 tomentum; innermost involucral bracts with quite large, ovate or broadly lanceolate appendages. Florets yellow, peripheral much larger, usually 4-10 mm longer than central florets; corolla 8—20-lobed, more or less hairy in middle. Achenes 3.5—4.5 mm long; pappus usually slightly longer than achenes. Flowering May to July. Clayey and stony slopes of foothills and lower mountain zone, especially on gypsiferous rocks, sometimes also as introduced or es- caped plant, along roads and close to settlements.—Caucasus: Eastern and Southern Transcaucasia (very rarely and only as an introduced or escaped plant near large towns); Soviet Central Asia: mountainous Turkmenia. General distribution: Iran. Described from cultivated speci- mens originating, apparently, from Iran. Type in London. Note. I am restoring the prior epithet—amberboi—of this species, which does not exactly match with the generic name “Amberboa” and thus is entirely valid. Economic Importance. Like A. moschata, this species has long been cultivated as an ornamental plant, but mostly in the countries of the Orient. 7. A. bucharica Ijin in Izv. Bot. Sada Akad. Nauk SSSR, XXX, 1—2 (1932) 108 and in Fl. Turkm. VII (1960) 255.—A. odorata var. bucharica Winkl. ex O. and B. Fedtsch. Perech. Rast. Turk. [TV (1911) 267, nom nudum.—Centaurea bucharica Iljin, loc. cit. (1932) pro syn. Annuals or biennials. Plants 15-60 cm high. Stem erect, simple or more or less branched, more or less covered with short crisped hairs. Leaves more or less covered with short crisped hairs (sometimes also with very thin arachnoid-tomentum); basal and lower cauline leaves very long-petiolate, usually undivided, but more or less toothed; middle cauline leaves usually also petiolate, undivided, more or less toothed or shallowly-pinnately lobed; upper cauline leaves sessile or short- petiolate, toothed or pinnately lobed. Capitula solitary at tips of stem and lateral branches, on long leafless peduncles. Involucres broadly ovate, 16-24 mm in dia, 16-24 mm long, usually with very copious (but on fruiting more or less disappearing) arachnoid tomentum; inner- most involucral bracts with quite large, ovate-lanceolate or oblong, almost coriaceous appendages. Florets yellow, peripheral florets rela- tively small, only scarcely surpassing central florets; corolla 8-20 lobed, glabrous or weakly hairy in middle. Achenes 4—6 mm long; pappus slightly longer than achenes. Flowering May to July. (Plate XVIII, Fig. 2: Plate, XX, Fig..3). Stony and clayey slopes of foothills and lower montane zone (usually to 1,500 m), especially on outcrops of gypsiferous rocks.— 333 330 Soviet Central Asia: Tien Shan (southwestern part), Syr-Darya, Amu- Darya, Pamiro-Alai Region (western part), mountainous Turkmenia (eastern part). General distribution: probably Iran. Described from Syr-Darya River valley. Type in Leningrad. Economic Importance. Perhaps used as an ornamental plant. GENUS 1613. Grossheimia Sosn. and Takht.': 2 Sosn. and Takht. in Dokl. Acad. Nauk Arm. SSR, II, 1 (1945) 22. —Centaurea sect. Jacea*** Macrocephalae Boiss. FI. or. III (1875) 616. Capitula heterogamous, large, with many florets. Involucre ovate- cylindrical or globose-cuboid, 20-50 mm in dia, glabrous or arach- noid-hairy; involucral bracts many-rowed, densely imbricate, coria- ceous, smooth, greenish (yellowish when dry), from ovate outer ones to oblong and oblong-lanceolate inner ones, with large, membranous, light reddish-brown or blackish, irregularly fimbriate, deeply palmately- or pinnately parted, stiff, bristly appendage, not decurrent on bracts. Receptacle flat, densely setose. All florets tubular, yellow, surpassing involucre, heterogenous; corolla glabrous; central florets (disk florets) bisexual, in several rows; peripheral florets (ray florets) sterile, with 4—5-parted corolla, 1-rowed, not broadened. Anthers with short basal appendages, apical appendages acute; stamen filaments flattened, cov- ered with papillae. Style unbranched almost to tip, very short-woolly, with collar of hairs at base, entirely exserted from anther tube. All achenes alike, oblong, strongly compressed, 6—8 mm long, truncate above, with small-toothed or obtuse crown, smooth, lustrous, initially scatteredly fine-hairy, mature achenes subglabrous, hilum lateral; pa- ppus as long as or slightly longer than achenes, ochraceous, persistent, double, outer one many-rowed, of serrate bristles, gradually longer from short outer to inner, not connate into ring at base, inner pappus l-rowed, of short, serrate, acute scales. Perennial herbs with simple stems, more or less strongly bulged below capitulum. Type of genus: G. macrocephala (Muss.-Puschk.) Sosn. and Takht. Besides the two species growing in the Caucasus, one more— C. helenoides (Boiss. and Hausskn.) Sosn. and Takht.—is found in Asia Minor (Turkish Lazistan). 'Treatment by D.I. Sosnowsky. *Named in honor of the eminent Soviet botanist, Academician A.A. Grossheim. {***—Not explained in the text—General Editor]. 334 Soil 1. Involucre glabrous; appendages of involucral bracts light reddish- brown, membranous, lustrous, irregularly fimbriate ...................... EA ER. TE 1. G. macrocephala (Muss.-Puschk.) Sosn. and Takht. + Involucre arachnoid-hairy; appendages of involucral bracts black- ish, palmately or pinnately divided into long stiff bristles almost to base, CHG cee iok dere. 2. G. ossica (C. Koch) Sosn. and Takht. Series 1. Macrocephalae Sosn. and Takht. in Dokl. Akad. Nauk Arm. SSR, II, 1 (1945) 23.—Appendages of involucral bracts membranous, irregularly fimbriate, occasionally deeply palmately fimbriate. 1. G. macrocephala (Muss.-Puschk.) Sosn. and Takht. in Dokl. Akad. Nauk Arm. SSR, II, 1 (1945) 23; Grossh. Opred. Rast. Kavk. 489.—G. macrocephala var. minor Sosn. and Takht. op. cit. 24.— Centaurea macrocephala Muss.-Puschk. in Willd. Sp. pl. III, 3 (1803) 2298; Adams in Web. Mohr. Beitr. I, 73; MB. FI. taur.-cauc. II, 351, Il, 592; DC. Prodr. VI, 568; Ldb. Fl. Ross. I, 2, 638; Boiss. Fl. or. III, 632; Lipsky, Fl. Kavk. 364; Grossh. Fl. Kavk. IV, 210.— Phaeopappus macrocephalus (Muss.-Puschk.) Boiss. Diagn. pl. or. ser. I, 6 (1845) 124; C. Koch. in Linnaea, XXIV, 437.—Chartolepis macrocephala (Muss.-Puschk.) Takht. in Ind. Sem. hort. bot. erev. (1940) 8.—Ie.: Bot. Mag. XXXI (1809) tab. 1248.—Exs.: Hayek, Cent. exs. crit. fasc. II, No. 66. Perennial. Root neck densely covered with dark fibers. Stem to 1 m high, simple, erect, thick, sulcate, more or less densely covered with crisped hairs, densely leafy. Lower leaves oblong, narrowed into short petiole; cauline leaves gradually reduced, sessile, decurrent, upper- most linear; all leaves remotely small-toothed, sometimes almost en- tire, scatteredly pubescent on both sides and densely glandular with small sessile glands. Capitula large. Involucre ovate-globose, 30-50 mm in dia; appendages of involucral bracts membranous, almost rounded irregularly and usually shallowly fimbriate, occasionally deeply palmately fimbriate (var. minor Sosn. and Takht.), imbricate and en- tirely covering coriaceous part of bracts. Florets bright yellow. Flow- ering July to September. Glades in forests of upper montane zone and in tall-herb vegeta- tion.—Caucasus: Eastern and Southern Transcaucasia. General distri- bution: Asia Minor (Turkish Lazistan), Armenia and Kurdistan. De- scribed from Georgia. Type in Leningrad. Series 2. Helenioides Sosn. and Takht. in Dokl. Akad. Nauk Arm. SSR, III, 1 (1945) 24.—Appendages of involucral bracts deeply pal- mately or pinnately divided into stiff bristles. 335) 582 2. G. ossica (C. Koch) Sosn. and Takht. in Dokl. Akad. Nauk Arm. SSR, II, 1 (1945) 24; Kolak. Fl. Abkhazii, IV, 284; Grossh. Opred. Rast. Kavk. 490.—G. ossica var. tuba (Somm. and Lev.) Sosn. and Takht. loc. cit—Phaeo-pappus ossicus C. Koch. in Linnaea, XVII (1843) 40; XXIV (1851) 436.—Centaurea polyphylla Ldb. ex Nordm. in Bull. Acad. Sc. Pétersb. II (1837) 311; Ldb. Fl. Ross. II, 2, 712.— C. ossica (C. Koch.) Boiss. Fl. or. HI (1875) 632; Alb. Prodr. Fl. Colch. 148; Lipsky Fl. Kavk. 148; Grossh. Fl. Kavk. IV, 211.—C. tuba Somm. and Lev. in Tr. Peterb. Bot. Sada, XII, 1 (1892) 156, XVI (1900) 282; Lipsky, op. cit. 365; Grossh. Fl. Kavk. IV, 211.—le.: Somm. and Lev. loc. cit. XVI (1900) tab. XXIV. Perennial. Stem to 1.5 m high, simple or weakly branched, erect, thick, sulcate, usually sparsely covered with crisped hairs, densely leafy. Leaves oblong-lanceolate, lower ones narrowed into petiole, oth- ers sessiles, sometimes auriculately at base, not decurrent; all leaves remotely small-toothed or almost entire, sparsely short-pubescent on both sides, with very tiny sessile glands above. Capitula large. Involu- cre subglobose, 20-30 mm in dia, more or less densely arachnoid- hairy; appendages of involucral bracts imbricate, almost to base pal- mately divided into stiff usually recurved bristle, sometimes appendages pinnately incised; middle cilia much longer than lateral (var. tuba (Somm. and Takht.) Sosn. and Takht.). Florets bright yellow. Flower- ing July to September. Glades in forests of upper mountain zone.—Caucasus: Eastern, Western, and Southern Transcaucasia (western part). Endemic. De- scribed from Ossetia. Type in Berlin. GENUS 1614. Chartolepis Cass." * Cass. in Dict. Sc. nat. XLIV (1826) 36, L (1827) 247, LIV (1829) 492; Czer. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XX (1960) 460.—Centaurea sect. Chartolepis (Cass.) DC. Prodr. VI (1837) 568 p. p. excl. Centaurea macrocephala Muss.-Puschk.; Benth. in Benth. and Hook. f. Gen. pl. II (1873) 479; O. Hoffm. in Pflanzenfam. IV, 5 (1893) 329.—Centaurea subgen. Chartolepis (Cass.) Schmalh. FI. Yugo-Zap. Ross. (1886) 328; Fl. II (1897) 120.—Chartolepis subgen. Euchartolepis Jaub. and Spach. Ill. pl. or. III (1847-1850) 136. Capitula heterogamous, medium-sized or somewhat large, with many florets, solitary at tips of stem and lateral branches or in race- 'Treatment by S.K. Czerepanov. From the Greek words chartes—sheet, paper, and /epis—scale; probably named for the membranous appendages of the involucral bracts. 336 335 mose, racemose-corymbose or corymbose inflorescence, less often single on plant. Involucres oblong-ovate, ovate to subglobose (8)10— 30 mm in dia, glabrous; involucral bracts many-rowed, densely imbri- cate, coriaceous, smooth, greenish (yellowing when dry), from broadly elliptical and elliptical-ovate outer ones to oblong-linear and almost linear innermost; appendages of involucral bracts more or less large, slightly decurrent on bract, thinly-membranous, semitransparent, with dark reddish-brown, brown, or slightly brownish spot at base, almost rounded to ovate (in innermost bracts), erose-toothed, mostly lacerate. Receptacle bristly. All florets tubular, yellow, far surpassing involu- cre, heterogeneous, corolla glabrous; central florets (disk florets) bi- sexual, in many rows; peripheral florets (ray florets) sterile, lacking staminodes and rudiment of style, with 4—5-parted corolla, 1-rowed, not large. Anthers with very small basal appendages, apical append- ages acute; stamen filaments flattened, covered with papillae. Stigma very short-plumose, with collar of hairs connate at base (its branches connate to tip) or slightly bilobate, entirely exserted from anther tube. All achenes alike, oblong-elliptical or oblong, 5—6 mm long, truncate above, with small-toothed crown at apex, compressed, smooth, scatteredly hairy; hilum lateral; pappus longer than achenes, weakly ochraceous, persistent, double (simple in C. saligna C. Koch), outer one of few irregular rows of flat, plumose, filiform bristles, gradually longer from outer to inner ones and connate into ring at base; inner one much shorter than outer, 1-rowed, usually broader, in upper part irregularly ciliate, on outside brown, almost scarious bristles. Peren- nial herbs with erect, branched, winged stems, leafy throughout, cov- ered with arachnoid, occasionally woolley hairs; basal and lower cauline leaves petiolate, sometimes toothed or lyrately divided, others sessile, long-decurrent on stem, entire, all scabrous from short spiny hairs. Type of genus: C. glastifolia (L.) Cass. The genus comprises seven species distributed in the southern European part of the USSR, Transcaucasia, Kazakhstan, Kirghizia, the eastern part of Asia Minor, and Iran. C. lyrata Boiss. (Iran and the Herat Province of Afghanistan), C. umesaoi Kitamura (Afghanistan), and C. saligna C. Koch (eastern part of Turkey) are not found in the USSR. 1. Involucre globose-ovate or subglobose, 20-30 mm in dia, append- ages of involucral bracts completely overlapping coriaceous part of bracts, with dark reddish-brown basal spot .............:cssssscssseeseereeeeeees LG. ROTO) SET LNG LE ee Ee 1. C. glastifolia (L.) Cass. + Involucre ovoid or oblong-ovate, (8)10—15 mm in dia; appendages of involucral bracts partially overlapping coriaceous part of bracts... 334 2. Capitula in rather compact corymbose inflorescence; pppentaes of involucral bracts with dark reddish-brown spot at base. Stem more or less scabrous, broadly winged) ...::1).)es...c.ceeereecerasesresesrenes Beco e void. swage Noll a ei ieee leila 3. C. pterocaula (Trautv.) Czer. + Capitula in lax racemose or racemose-corymbose inflorescence; ap- pendages of involucral bracts with pale brownish spot at base or almost entirely sennitransparent,{).4<1./escne de ce ee as 3. 3. Stem more or less well proportioned, narrowly-winged; cauline leaves narrow, usually oblong-linear to narrowly linear (Transcaucasia) it..tetiee soe 4. C. biebersteinii Jaub. and Spach. + Stem rather scabrous, broadly-winged; cauline leaves broader (south- em European part of USSR, Kazakhstan and Kirghizia) ................... a Lies capieb whee eae hae et A 2. C. intermedia Boiss. 1. C. glastifolia (L.) Cass. in Dict. Sc. nat. LIV (1829) 492 p. p. quoad nom.; Czer. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XX (1960) 461; Karjagin in Fl. Azerb VIII, 435.—Centaurea glastifolia 337 L. Sp. pl. (1753) 915 p. p. quoad pl. ex. “Oriente,” ed. 2 (1763) 1294 p. p.; DC. Prodr. VI, 568 p. p.; Ldb. Fl. Ross. II, 2, 687 p. p. quoad pl. e Caucaso.—C. tournefortii (Jaub. and Spach) Walp. Ann. Bot. I (1848-1849) 448.—C. glastifolia var. tournefortii (Jaub. and Spach) Gugler in Ann. hist.-nat. Mus. Nat. Hung. VI (1908) 239.—Chartolepis tournefortii Jaub. and Spach, Ill, pl. or. III (1847-1850) 9; Boiss. FI. or. III, 696; Iljin in Izv. Bot. Sada Akad. Nauk SSSR, XXX, 3-4, 353; Grossh. Fl. Kavk. IV, 196.—Ie.: Jaub. and Spach, op. cit. tab. 207. Perennial. Root woody, oblique. Stem 40-80 cm high, erect, quite scabrous, finely ribbed, broadly winged, branched from middle or slightly above, weakly arachnoid-hairy or woolly, occasionally subglabrous, lateral branches upward spreading, more or less long. Leaves subacute, to short-acuminate, entire, scabrous from numerous pointed spinous hairs, often mixed with occasional arachnoid hairs and sessile, golden, punctate glands. Basal and lower cauline leaves ob- long-oblanceolate, elliptical-lanceolate, or almost lanceolate, sometimes weakly toothed, relatively short, to 5 mm wide, long-petiolate, others oblong-lanceolate to linear, sessile, long-decurrent on stem forming broad wings 2-7 mm wide. Capitula solitary at tips of stem and lateral branches, in racemose-corymbose or corymbose inflorescence. Involu- cre globose-ovate or subglobose, 23-30 mm long and 20-30 mm in dia; appendages of involucral bracts thinly membranous, completely overlapping coriaceous part of bracts and much longer than them. Achenes reddish-brown, 5.5-6.0 mm long and 2.0-2.3 mm wide; pappus 11-12 mm long. Flowering July to August. (Plate XIX, Fig. 2). 338 355 Meadow slopes, forest edges and glades; in mid- and upper-mon- tane zones, at 1,700-—2,200 m.—Caucasus: Eastern Transcaucasia (Lesser Caucasus), Southern Caucasia. General distribution: Armenia and Kurdistan, Asia Minor (eastern part). Described from Turkish Armenia from collections of Tournefort. Type in Paris. Note. Arguments in favor of using the name C. glastifolia (L.) Cass. for this species have been presented by us in an earlier published paper (Czerepanov, op. cit.). 2. C. intermedia Boiss. Diagn. pl. or. ser. II, 3 (1856) 64; FI. or. III, 696, in adnot.; Ijin in Izv. Bot. Sada, Akad. Nauk SSSR, XXX, 3-4 (1932) 352, 353; in Fl. Yugo-Vost. VI, 420; Kryl. Fl. Zap. Sib. XI, 2947; Grossh. Opred. Rast. Kavk. 490; Dobrocz. in Visn. Rosl. URSR, 571; Czer. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XX, 463.—C. biebersteinii Jaub. and Spach, Ill. pl. or. II (1847- 1850) 11 p. p.—Centaurea glastifolia L. Sp. pl. (1753) 915 p. p., ed. 2 (1763) 1294 p. p. quoad pl. sibir.; DC. Prodr. VI, 568 p. p.; Ldb. Fl. Ross. II, 2, 687 p. p.; Schmalh. Fl. II, 120 and auct. al. fl. Ross. (excl. pl. transcauc.).—C. glastifolia var. intermedia (Boiss.) Gugler in Ann. hist.-nat. Mus. Nat. Hung. VI (1908) 239.—Ic.: Iljin in FI. Yugo-Vost. VI (1936) Fig. 708; Dobroz. op. cit. Fig. 300; Stank. and Tal. Opred. Vyssh. Rast. ed. 2, Fig. 395. Perennial. Root woody, branched, thickened at root collar. Stem 50-100(150) cm high, erect, very scabrous, thinly ribbed, broadly winged, branched from middle or slightly above, weakly arachnoid- hairy, occasionally subglabrous, moderately covered with sessile, golden glands, lateral branches upward spreading, more or less long. Leaves subacte to short-acuminate, entire, scabrous from numerous, pointed, spiny hairs, usually with few arachnoid hairs, with sessile, golden, punctate glands. Basal and lower cauline leaves oblong- or elliptical- oblanceolate to almost lanceolate, sometimes weakly toothed, 2—6 cm wide, petiolate, petioles 4—20(25) cm long, others oblong-lanceolate, oblong, or oblong-linear, sessile, long-decurrent on stem forming broad, 2-10 mm wide wings. Capitula solitary at tips of stem and lateral branches, in relatively lax racemose or racemose-corymbose inflores- cence. Involucre oblong-ovate, 22—25 mm long and 10—15 mm in dia; appendages of involucral bracts thinly membranous, partially overlap- ping coriaceous part of bracts, with pale brownish spot at base or almost completely semitransparent. Florets yellow. Achenes brownish- creamish, 5.5—6.0 mm long, about 2.3 mm wide; pappus 8-10 mm long. Flowering July to August. Alkaline wet meadows and salt-marsh meadows, in cheegrass thick- ets, river valleys, and on lake shores.—European Part: Bessarabia 336 (extreme south), Black Sea Region, Middle Dnieper (southeastern part), Volga-Don (south), Lower Don, Lower Volga, Trans-Volga, Volga- Kama (southeastern part); Western Siberia: Upper Tobol (southern part), Irtysh (southern part), Altai (extreme southwest); Soviet Central Asia: Aralo-Caspian Region, Lake Balkhash Region, Dzhungaria- Tarbagatai, Tien Shan. General distribution: Dzhungaria-Kashgaria (Kuldzha, Kemerchik River in Mogolian Altai). Described from Cen- tral Russia, Siberia and Dzhungaria. Type in Geneva. 3. C. pterocaula (Trautv.) Czer. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XX (1960) 463; Karjagin in Fl. Azerb. VIII, 435.— C. biebersteinii auct. p. p. non Jaub. and Spach; Boiss. Fl. or. III (1875) 696; Grossh. Fl. Kavk. IV, 196; Opred. Rast. Kavk. 490.—C. finitima (Bordz.) Ijin in Izv. Bot. Sada Akad. Nauk SSSR, XXX, 3- 4 (1932) 353.—Centaurea pterocaula Trautv. in Tr. Peterb. Bot. Sada II (1873) 552, excl. syn. non Romel (1874).—C. finitima Bordz. in Tr. Bot. Sada Yurevsk. Univ. XIII (1912) 24. Perennial. Root woody, branched. Stem to 100 cm high, erect, scabrous, finely ribbed, broadly winged, branched above, weakly arach- noid-hairy, covered with scattered, sessile, golden glands, lateral branches upward-spreading, short. Leaves acute to short-acuminate, entire, scabrous from numerous, acute, spinous hairs, with few thin arachnoid hairs, and sessile, golden, punctate glands; basal and lower cauline leaves oblong-oblanceolate or lanceolate, sometimes weakly toothed, 2-3 cm wide, petioles, to 12 cm long, others oblong-lan- ceolate, lanceolate-linear, less often almost linear, sessile, long-decur- rent on stem forming wings 2—6(8) mm wide. Capitula solitary at tips of stem and short lateral branches, in compact corymbose inflores- cence. Involucre ovate, rarely oblong-ovate, 20 cm long and 10—13 mm in dia; appendages of involucral bracts thinly membranous, par- tially overlapping coriaceous part of bracts, with dark reddish-brown spot at base. Florets yellow. Achenes 5 mm long about 2 mm wide; pappus 8-10 mm long. Flowering July to August. Meadow slopes, forest edges and glades, in mid- and upper-mon- tane zones, to 2,500 m.—Caucasus: Southern Transcaucasia. General distribution: Armenia and Kurdistan,? Iran (Khvoy District). Described from Turkish Armenia (Kazikibaran). Type in Leningrad. 4. C. biebersteinii Jaub. and Spach, Il. pl. or. III (1847-1850) 11 p. p.; Boiss. Fl. or. II, 696 p. p. max.; Iljin in Izv. Bot. Sada, Akad. Nauk SSSR, XXX, 3-4, 353 p. p. excl. specim. nonnull.; Grossh. FI. Kavk. IV, 196 p. p. and Opred. Rast. Kavk. 490 p. p.; Czer. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XX, 464; Karjagin in FI. 340 337 Azerb. VIII, 435.—C. glastifolia auct. non Cass.; C.A.M. Verzeichn. (1831) 231, excl. syn.—Centaurea glastifolia auct. non. L.: MB. FI. taur.-cauc. II (1808) 355, excl. syn. Tourn.; Ldb. Fl. Ross. II, 2, 687 p. p.—C. biebersteinii (Jaub. and Spach) Walp. Ann. Bot. I (1848— 1849) 448.—C. glastifolia var. biebersteinii (Jaub. and Spack) Gugler in Ann. hist.-nat. Mus. Nat. Hung. VI (1908) 239.—Ic.: Jaub. and Spach, op. cit. tab. 208. Perennial. Root woody, branched. Stem 60—100(120) cm high, erect, very well proportioned, thinly ribbed, narrowly winged, branched from slightly above middle, weakly arachnoid hairy or subglabrous, sparsely glandular with sessile, golden glands, lateral branches upward spreading, quite long. Leaves acute to short-acuminate, entire, sca- brous from short spines or spiny hairs, fewer toward midrib, with occasional arachnoid hairs covered with sessile, golden, punctate glands; basal and lower cauline leaves almost oblong or lanceolate, to 2 cm wide, long-petiolate, others oblong- or lanceolately linear to narrowly linear, sessile, long, sometimes short, decurrent on stem forming nar- row wings 1-3 mm wide. Capitula solitary at tips of stem and lateral branches; in more or less lax racemose-corymbose inflorescence. In- volucre ovate, less often oblong-ovate, about 20 mm long and (8)10— 14 mm wide; appendages of involucral bracts thinly membranous, par- tially overlapping coriaceous part of bracts, with brownish spot at base or almost completely semitransparent. Florets yellow. Achenes 5.0— 5.5 mm long and 2.0—2.3 mm wide; pappus 10-11 mm long. Flowering July to August. Dry slopes; in mid-montane zone.—Caucasus: southern Transcaucasia (southern part), Talysh (Zuvant). General distribution: Iran-Afghanistan (northwestern part), Armenia and Kurdistan (Sarykamysh). Described from Caucasus. Type in Paris; isotype in Leningrad. GENUS 1615. Callicephalus C.A.M.'? C.A.M. Verzeichn. (1831) 66; Czer. in Fl. Turkm. VII (1960) 262.—Phalolepis Cass. in Dict. Sc. nat. L (1827) 248 p. p.—Centau- rea sect. Phalolepis (Cass.) DC. Prodr. VI (1837) 568 p. p.—Centau- rea sect. Callicephalus (C.A.M.) O. Hoffm. in Pflanzenfam. IV, 5 (1893) 327. "Treatment by S.K. Czerepanov. "From the Greek words kallos—beautiful, and kephalon—head. 342 338 Capitula homogamous, medium-sized, many-flowered, solitary at tips of stem and lateral branches, in more or less racemose or corym- bose-racemose inflorescence, less often single on plant. Involucres globose-ovate, 15-20 mm in dia, glabrous; involucral bracts many- rowed, imbricate, thinly-coriaceous, with 5—8 distinct, veins, greenish, oblong-elliptical outer ones to linear innermost; appendages of involu- cral bracts large, not decurrent on bracts, completely overlapping co- riaceous part of bracts, finely ciliate, semitransparent (in outer and often middle bracts), slightly brownish to dark lilac-purple (in middle and innermost bracts), lustrous, bulged on outside, roundish and roundish-rhombic to ovate, undivided, usually somewhat lacerate, with small brownish cusp, mostly at base of small apical notch. Recep- tacle setose. All florets tubular, pale lilac-pink, surpassing involucre, homogeneous, bisexual; corolla glabrous. Anthers with small basal ap- pendages; apical appendages subobtuse; stamen filaments flat, covered with papillae. Stigma short-plumose, with collar of hairs at base, bi- lobed, included in or exserted from anther tube by its lobes. All achenes similar, 4-angled-cuneate, 4.0-4.5 mm long, truncate above, with scarcely toothed crown at apex, rugose-tuberculate, glabrous, hilum lateral; pappus longer than achene, ochraceous, persistent, simple, of numerous serrate, elastic bristles, longer from outer to inner, connate in ring at base; 4 inner pappus bristles noticeably longer and broader than outer ones. Annual herbs with erect, branched, throughout leafy stem and scabrous, pinnately incised, sessile leaves not decurrent on stem. A monotypic genus. 1. C. nitens (MB.) C.A.M. Verzeichn. (1831) 66; Boiss. FI. or. III, 703; Grossh. Fl. Kavk. IV, 197; Opred. Rast. Kavk. 490; Czer. in Fl. Turkm. VII, 262; Agadj. in Fl. Azerb. VIII, 436.—Centaurea nitens MB. in Willd. Sp. pl. III, 3 (1803) 2305; MB. FI. taur.-cauc. II, 354, III, 593; DC. Prodr. VI, 568; Ldb. Fl. Ross. II, 2, 668.—Serratula nitens (MB.) Spreng. Syst. veg. 3 (1826) 389.—Ic.: Jaub. and Spach, Ill. pl. or. III (1847-1850) tab. 219; Czer. in Fl. Turkm. VII (1960) Plate XL.—Exs.: Herb. Fl. cauc. No. 147; Fl. cauc. exs. No. 25; Pl. or. exs. No. 124; Ed. Hort. Bot. Petri Magni, No. 100; Aucher-Eloy, Herb. d’Orient. No. 4840; Sintenis, It. transcasp.-pers. 1900-1901, No. 1880. Annual. Whole plant green, scabrous- from pointed, spiny hairs. Stems 20-80 cm high, erect, branched from middle or above, with thin, virgate, upward spreading long branches. Leaves pinnately in- cised, sessile, not decurrent on stem; leaf segments narrowly linear, terminating into small bristle, entire, convolute. Capitula in more or Plate XIX. 1—Cheirolepis persica Boiss.; 2—Chartolepis glastifolia (L.) Cass. 343 340 less lax racemose or corymbose-racemose inflorescence, occasionally single on plant. Involucre globose-ovate, 15—20 mm long and as much in dia; appendages of outer and middle involucral bracts semitranspar- ent or slightly brownish, roundish, and roundish-rhombic, weakly lac- erate, with small notch at tip; appendages of inner involucral bracts brownish to dark lilac-purple, ovate, not lacerate, lacking notch at tip. Florets pale lilac-pink. Achenes 4.0—4.5 mm long and about 1.5 mm wide; pappus 12—15(17) mm long. Flowering June to July. (Plate XXI, Fig. 1). Dry stony and rubbly slopes, on screes and rocks; in lower and mid-montane zones.—Caucasus: Dagestan (south), Eastern and South- ern Transcaucasia; Soviet Central Asia: mountainous Turkmenia (Kopetdag Range). General distribution: Armenia and Kurdistan (former Artvin District), Iran-Afghanistan (northwestern part). Described from Caucasus. Type in Leningrad. Note. In Flora Turkmenia (Vol. VII, Plate XL) there is an error in the illustration of the pappus of the achenes. It is that, in reality, the four inner pappus bristles are narrower in the upper part, setaceous, and longer than the outer bristles, but not to the extent shown in the plate. GENUS 1616. Acroptilon Cass.’ ? Cass. in Dict. Sc. nat. L (1827) 464; Czer. in Fl. Turkm. VII (1960) 257.—Centaurea sect. Acroptilon (Cass.) Benth. in Benth. and Hook. f. Gen. pl. II (1873) 478; O. Hoffm. in Pflanzenfam. IV, 5 (1893) 327.—Centaurea subgen. Acroptilon (Cass.) Schmalh. Fl. Yugo- Zap. Ross. (1886) 328; Fl. If (1897) 118. Capitula homogamous, very small or medium-sized, with many florets, solitary at tips of stem and lateral branches, in racemose-corym- bose or paniculate inflorescence. Involucres ovate, oblong-ovate or subglobose, 5—13(15) mm in dia; glabrous; involucral bracts many- rowed, imbricate, thinly coriaceous, with inconspicuous veins, green- ish (yellowing when dry), from more or less round and semielliptical outer ones to linearly lanceolate innermost; appendages of involucral bracts scarious, in outer and middle bracts as broad, semitransparent, short-hairy border, in innermost bracts very narrow, linearly lanceolate, densely long-pilose. Receptacle bristly. All florets tubular, pink or 'Treatment by S.K. Czerepanov. "From the Greek words acron—pointed, tip, and ptilon—wing; named for the pointed appendages of the involucral bracts. 344 341 lilac-pink, 1.5 times as long as involucre, homogeneous, bisexual; corolla glabrous. Anthers with small basal appendages, apical append- ages obtuse; stamen filaments glabrous. Stigma very short-woolly, with collar of hairs at base, bifid almost to middle, entirely or only their branches exserted from anther tube. All achenes similar, ellipsoidal- obovate, 3—4 mm long, sharply rounded above, compressed, with in- distinct longitudinal veins, glabrous, hilum slightly oblique; pappus usually 3 times as long as achene, whitish, very readily deciduous, with simple or serrate, flattened bristles, longer from short outer ones to inner, not connate into ring at base; innermost pappus bristles short- hairy above. Perennial herbs with erect, strongly branched stems, densely leafy throughout; leaves arachnoid-hairy, scabrous on margin, sessile, not decurrent on stem, pinnately divided or with remote teeth. Type of genus: A. repens (L.) DC. The two species of this genus are distributed in the southeastern European part of the USSR, extreme south of Western Siberia, through- out Soviet Central Asia, West Asia and farther in the east up to Mongolia inclusively. 1. Involucres 13—18(23) mm long and 7—13(15) mm in dia; append- ages of outer and middle involucral bracts acuminate, completely overlapping and sia i surpassing coriaceous part of ntaictisusts, cleugeeetty.siennets. bape eto eah eee on 1. A. australe Ijin + Involucres 10-13 mm long and 5—7 mm in dia; appendages of outer and middle involucral bracts obtuse to acute, partially over- lapping and usually not surpassing coriaceous part of bractsyemndh cD pee trie are enters aes 2. A. repens (L.) DC. 1. A. australe Iljin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, VII, 3 (1937) 59; Czer. in Fl. Turkm. VII, 258.—Ic.: Iljin op. cit. Fig. 4.—Exs.: Bornm. It. Pers.-turc. 1892-1893, Nos. 4065, 4066, 4067, 4069, 4089; Sintenis, It. transcasp.-pers. 1900-1901, Nos. 359, 359c, 698, 698b. Perennial. Root string-like, long, with lateral, more or less hori- zontal or obliquely rising, long suckers. Stem 1 or few (2-3), 30-70 cm high, erect, ribbed, strongly branched almost from base or slightly above, grayish-arachnoid hairy, underground part of stem blackish- brown with blackish or brownish scales. Leaves stiff, oblong to almost linear, usually with small, whitish, cartilaginous cusp, pinnatifid and remotely toothed to entire upper cauline, mostly weakly arachnoid- hairy, scabrous or spinulose along margin, with sessile punctate glands, sessile, not decurrent on stem. Capitula solitary at tips of stem and lateral branches, in lax racemose-corymbose or paniculate inflores- 345 342 cence. Involucre oblong-ovate, ovate, or subglobose, 13—18(23) mm long and 7—13(15) mm in dia; appendages of involucral bracts scari- ous, completely overlapping in outer and middle bracts and much sur- passing coriaceous part of bracts, acuminate, entire, pubescent, in in- nermost bracts very narrow, linear-lanceolate, densely pilose, slightly recurved, 6-12 mm long. Florets lilac-pink. Achenes (3.0)3.5—4.0 mm long and 1.5—2.0 mm wide; pappus 8—12(13) mm long. Flowering May to August. Clayey, sandy, and stony mountain slopes, up to 2,100 m, along sandy-gravelly banks of rivers and lakes; as weed in crops, in old fields, etc.—Soviet Central Asia: mountainous Turkmenia, Kara-Kum (Krasnovodsk), Aralo-Caspian Region (chalky mountains near Ak- Chaimak in Mangyshlak), Pamiro-Alai (Alai Range, vicinity of Gulcha), Tien Shan (Issykkul Depression, Ketmen Range, Trans-Ili Alatau Range; vicinity of Alma-Ata), Lake Balkhash Region (Ili-Borokhudzir cross- ing), Dzhungaria-Tarbagatai (southwestern part of Dzhungarian Alatau). General distribution: Iran, Dzhungaria-Kashgaria (Uchturfan, “Cherny” [Blackj Irtysh River above mouth of Kran River, Tekes River), Mongolia (western part). Described from Kopetdag (Kizil-Imam on Chandar River). Type in Leningrad. Note. It is distinguished from A. repens (L.) DC. by larger involu- cres, acuminate appendages of the outer and middle involucral bracts that entirely overlap the bracts and are much longer than the coria- ceous part of the inner involucral bracts, and by the longer appendages of the inner involucral bracts. Forms with intermediate features are commonly found where the ranges of the two species come in contact. The absence of similar intermediate forms in regions where these species do not grow to- gether leads us to assume a hybrid origin. Economic Importance. A noxious weed of crops. 2. A. repens (L.) DC. Prodr. VI (1837) 663; Grossh Opred. Rast. Kavk. 490; Kryl. Fl. Zap. Sib. XI, 2945; Czer. in Fl. Turkm. VII, 257; Agadzh. in Fl. Azerb. VIII, 437.—A. obtusifolium, subdentatum, serratum and angustifolium Cass. in Dict. Sc. nat. L. (1827) 465, 466. A. picris (Pall ex Willd.) C.A.M. Verzeichn. (1831) 67; DC. op. cit. 662; Ldb. FI. Ross. II, 2, 750; Boiss. Fl. or. III, 612; O. and B. Fedtsch. Perech. Rast. Turk. IV, 289; Grossh. Fl. Kavk. IV, 197; Dobrocz. in Vizn. Rosl. URSR, 572; Iljin in Fl. Yugo-Vost. VI, 418; Stank. and Tal. Opred. Vyssh. Rast. ed. 2, 412.—Centaurea repens L. Sp. pl. ed. 2 (1763) 1293.—C. picris Pall. Tabl. phys. et topogr. taur. (1795) 58 and ed. 2 (1796) 141, nomen; Willd. Sp. pl. III, 3, 2302, descr.; MB. Fl. taur.-cauc. II, 352; Schmalh. FI. II, 118.—Serratula picris (Pall. ex. 346 343 Willd.) MB. Fl. taur.-cauc. III (1819) 546; Ldb. Fl. alt. IV, 41.—Ic:: Jaub. and Spach, Ill. pl. or. I (1842-1843) tab. 10, 11; Malkov, Glavn. Sornyaki Turkm. SSR (1936), fig. 7; Iljin in Fl. Yugo-Vost. VI, Fig. 706; in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, VII, 3 (1937) Fig. 4A; Stank. and Tal. loc. cit. Fig. 394.—Exs.: GRF Nos. 1881, 1983a, 1983b; Herb. Fl. cauc. No. 148; D6rfler, Herb. norm. No. 4150. Perennial. Root string-like, very long (to 6 m), with lateral, more or less horizontal or obliquely rising, long (to 1 m and more) suckers. Stem single or few (2-3), 15—60(75) cm high, erect, ribbed, strongly branched almost from base or slightly above, grayish-arachnoid-hairy, occasionally subglabrous; underground part of stem blackish-brown, covered with brown or blackish scales. Leaves stiff, oblong to almost linear, with very small cusp, pinnatifid and with remote teeth to entire upper cauline, grayish arachnoid-hairy, sometimes almost lacking arach- noid pubescence, usually scabrous only along margin from acute, short, spiny hairs or obtuse spines, covered with punctate sessile glands, sessile, not decurrent to stem. Capitula solitary at tips of stem and lateral branches, in lax racemose-corymbose or paniculate inflores- cence. Involucre oblong-ovate or ovate, 10-13 mm long and 5—7(8) mm in dia; appendages of involucral bracts scarious in outer and middle bracts, partially overlapping and not surpassing coriaceous part of inner bract, obtuse to acute, often weakly lacerate, short-hairy, narrow in innermost bracts, densely long-hairy, somewhat recurved, 3.0—4.5 mm long. Florets pink. Achenes (2.5) 3.0—4.0 mm long and (1.5) 2.0 mm wide; pappus 8—11 mm long. Flowering May to August. Clayey, sandy, and stony steppes, on alkaline meadows and salt marshes, cheegrass thickets, sandy-gravelly and rocky shores of rivers and lakes, stony and clayey mountain slopes, from lowlands to 2,600— 2,800 m; as weed in crops, in old fields, roadsides, in residential areas, etc.—European Part: Crimea, Black Sea Region (in the west to Kherson Region inclusively), Lower Don, Volga-Don (vicinity of Voronezh, introduced), Lower Volga, Trans-Volga (south); Caucasus: Ciscaucasia, Dagestan, Eastern and Southern Transcaucasia, Talysh (Zuvant); West- ern Siberia: Upper Tobol (southern part), Irtysh (southern part), Altai southwestern part); Soviet Central Asia: all regions (absent in Kopetdag). General distribution: Mongolia, Dzhungaria-Kashgaria, Iran (mostly northern part), Armenia and Kurdistan, Asia Minor (mostly northern part, in west up to Esk). Described from “Orient.” Type in London. Note. This species is highly variable in appearance over its vast range, but no geographic pattern could be found in its variation. Cassini described some forms of A. repens (L.) DC. without discrete ranges, as separate species (cf. synonymy), which quite rightly are not recog- 347 344 nized even by De Candolle (loc. cit.). Kuntz (Tr. Peterb. Bot. Sada, X, 1 (1887) 201) studied polymorphism in this species. The majority of the infraspecific taxa described by him do not merit, in my opinion, separate names. Economic Importance. A noxious, hardy weed of crops. GENUS 1617. Oligochaeta C. Koch’ * C. Koch in Linnaea, XVII, (1843) 42; Czer. in Fl. Turkm. VII (1960) 259.—Serratula sect. Oligochaeta (C. Koch) DC. Prodr. VI (1837) 671.—Microlonchus auct. p. p. non Cass.: Boiss. Fl. or. III (1875) 700.—Centaurea sect. Microlonchus (Cass.) O. Hoffm. in Pflanzenfam. IV, 5 (1893) 329 p. p.—Cousinia subgen. Oligochaeta (C. Koch) Winkl. in Tr. Peterb. Bot. Sada, XIV, 2 (1898) 188, 217. Capitula homogamous, with many-florets, 2—5(7), aggregated, oc- casionally solitary at tips of stem and axillary. Involucres oblong or oblong-ovate, 4—5 mm in dia, whitish-arachnoid-hairy; involucral bracts many-rowed, imbricate, semicoriaceous, with indistinct veins, greenish (yellowing when dry), from triangular or oblong-lanceolate outer ones to lanceolate and lanceolate-linear innermost, with rather long, strongly recurved and light-colored (in outer and middle bracts) or straight and violet-carmine (in inner bracts) awn-like spine. Receptacle bristly, more or less deeply laveolar. All florets tubular, lilac-pink, homogeneous, bisexual; corolla glabrous. Anthers with small basal appendages; sta- men filaments flattened, covered with papillae. Stigma very short-hairy, with collar of hairs at base, bifid to collar of hairs, with divergent branches, fully or partly exserted from anther tube. Achenes variable or similar, oblong-ellipsoidal, scarcely narrowed toward base, 2.0—2.5 mm long, truncate above, with obtuse or scarcely toothed corona at tip, slightly compressed, glabrous; peripheral achenes transversely rugose, dark-smoky, sometimes weakly tetraquetrous, occasionally, obtuse above, with very light-colored, entirely deciduous pappus or like inner ones; inner achenes smooth, dark reddish-brown or dark sandy, narrower, with persistent, slightly longer pappus; hilum basal, punctate; pappus as long as achenes or longer, white, double: outer one with relatively few, gradually longer from outer to inner, serrate- scabrous hair-like bristles, connate into ring at base; inner ones with 1 (rarely 2) serrate-scabrous, broadened below, stiff bristles, longer 'Treatment by S.K. Czerepanov. *From the Greek words oligis—less, few, and chaete—hairs; named for the achenes that are scarcely toothed above. 348 345 than outer pappus, and 3—5 small, unequal scales, projecting from upper edge of ring. Annual herbs with divaricately branched stem and arachnoid-hairy or arachnoid-tomentose, sometimes eee un- divided or lyrately pinnately incised leaves. Type of genus: O. divaricata (Fisch. and Mey.) C. Koch. The genus includes three species distributed in Transcaucasia, Soviet Central Asia, and Iran. 1. Florets much longer than involucre; corolla 12—15 mm long; achenes vaniable( Transcaucasia) sa). west eee chee Dae eee Lee 2). + Florets barely longer than involucre; corolla 7.5 mm long; all achenes alike; plants low or dwarf (Soviet Central Asia) ...........:cscseeee ma) eral in. Sy, sboumd tains. ffm. 3. O. minima (Boiss.) Briq. 2. Plants weakly arachnoid-hairy or subglabrous; leaves remotely ioothe ducanelylyratels..ulleiucds: ores Ivcoalciel ne eee de bald wee x ue ae rane Eee es 1. O. divaricata (Fisch. and Mey.) C. Koch + Plants arachnoid-hairy-tomentose; leaves lyrate ............cceeeeees ROM A. _.2yrs)AtaSh. cha) ARIE Sok) i ctec 2. O. tomentosa Czer. 1. O. divaricata (Fisch. and Mey.) C. Koch in Linnaea, XVII (1843) 43; Ldb. Fl. Ross. II, 2, 723; Grossh. Opred. Rast. Kavk. 490; Agadzh. in Fl. Azerb. VIII, 440.—Serratuia divaricata Fisch. and Mey. Ind. Sem. hort. Peterop. I (1835) 37; DC. Prodr. VI, 671.— Microlonchus oligochaetus Boiss., Fl. or. II (1875) 701; Grossh. FI. Kavk. IV, 198.—Cousinia Massalskyi Winkl. in Tr. Peterb. Bot. Sada, XIV, 2 (1898) 217.—Ie.: Iljin in Sov. Bot. 5 (1935) Fig. 1 and p. 123; Agadj. loc. cit. Plate XLIV.—Exs.: Pl. or. exs. No. 49. excl. syn. Annual. Stem (5)10—40 cm high, usually very strongly divaricately branched almost from root collar, with weakly white-arachnoid-hairy or subglabrous, remote, spreading, lower usually more or less divaricate, ribbed, branches sparsely leafy throughout. Leaves glabrous or with scattered hairs above, weakly white-arachnoid-hairy or subglabrous beneath, sometimes mixed with spiny hairs, oblong or oblong-lanceolate, with small, thin, white cusp, remotely toothed, less often lyrate, short- petiolate, upper ones subsessile. Capitula 2—5(7), aggregated at tips and axillary, usually many on plant. Involucre oblong or oblong-ovate, 8—12 mm long and 4—5 mm in dia; involucral bracts with quite long, awn-like spine. Florets lilac-pink, much surpassing involucre; corolla 12-15 mm long. Achenes variable: peripheral ones transversely rug- ose, dark-smoky, 2.0—2.2 mm long and about 0.9-1.0 mm wide; inner ones smooth, dark reddish-brown, 2.2 mm long and about 0.8 mm wide. Pappus 2-3 mm long; bristles of inner pappus scarcely longer, slightly broadened toward base. Flowering May to July. 349 346 Dry, stony, stony-sandy, and clayey slopes up to lower mountain zone.—Caucasus: Southern Transcaucasia (Araks River Basin). Gen- eral distribution: Armenia and Kurdistan (Kagyzman District), Iran (northwestern part). Described from northwestern Iran (Khuoy). Type and isotype in Leningrad. Note. It differs from the closely related O. tomentosa Czer. by having weakly arachnoid pubescence on the entire plant, mostly undi- vided leaves, and somewhat different distribution. From another closely related species, O. minima (Boiss.) Briq., it is well distinguished by larger florets that are much longer than the involucre, variable achenes in the capitulum, a higher growth in general, the pubescence and shape of the leaves, and the well delimited range. In 1957, N.N. Tzvelev and I collected O. divaricata (Fisch. and Mey.) C. Koch near the southwestern edge of Ekhegnadzor (Daralgyez). Most likely, it was introduced incidentally at the reported locality from the Araks River valley. 2. O. tomentosa Czer. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX (1959) 455. Annual. Stem 30-40 cm high, strongly divaricately branched much above root collar, with remote, spreading, ribbed, more or less long branches, sparsely leafy to tip. Leaves weakly arachnoid-hairy or subglabrous above, like stem white-arachnoid-hairy tomentose beneath, lyrate or lyrati-pinnately incised, short-petiolate, upper subsessile, sometimes undivided, deeply toothed, lateral leaf segments 2—4 on each side, terminating in small, thin, white cusp. Capitula 2—3 or soli- tary at tips and axillary, rather numerous on plant. Involucre oblong or oblong-ovate, 10-12 mm long and 4—5 mm in dia; involucral bracts with rather, long awn-like spine. Florets lilac-pink, far surpassing in- volucre; corolla 12-13 mm long. Achenes variable, peripheral ones transversely rugose, dark-smoky, 2 mm long and about 0.9 mm wide, inner ones smooth, dark reddish-brown, 2.2 mm long and about 0.8 mm wide; pappus 2 mm long, inner pappus bristles scarcely longer, basally slightly broadened. Flowering June to July. Dry steppes.—Caucasus: Eastern Transcaucasia (Naftalan, Gean steppe). Endemic. Described from Naftalan, Azerbaidzhan SSR. Type and paratype in Leningrad; isotype in Baku. 3. O. minima (Boiss.) Brig. in Archiv. Sc. Phys. et Nat. Geneve, 5, XII (1930) 113 n. v.; Iljin in Sov. Bot. 5, 121; Czer. in Fl. Turkm. VII, 260.—Microlonchus minimus Boiss. FI. or. II (1875) 701; O. Fedtsch.-in Izv. Bot. Sada, VI, 29; O. and B. Fedtsch. Perech. Rast. Turk. IV, 280; Bornm. in Beih. Bot. Centralbl. LIX B (1939) 305; 350 347 Parsa, Fl. Iran, III, 703.—Jurinea korolkowi Rg]. and Schmalh. in Tr. Peterb. Bot. Sada, VI, 2 (1880) 322.—Centaurea minima (Boiss.) B. Fedtsch. Rast. Turk. (1915) 757; Pavlov, Fl. Tsentr. Kazakhst. III, 322, 323.—C. vvedenskyi M. Pop. in Tr. Sredneaz. Gos. Univ. Ser. VIIIb, Bot. 3 (1928) 111 (Sched. ad Herb. Fl. As. Med. fasc. XX).— Exs.: Bornm. It. Pers.-turc. 1892-1893, No. 4048; H.F.A.M. No. 500. Annual. Plant low or dwarf. Stem very much reduced, terminating into single capitulum, divaricately branched almost from root collar with remote, spreading or more or less divaricate, ribbed branches, sparsely leafy to tip, usually densely crisped-hairy, with sordid-white hairs. Leaves usually not densely arachnoid-hairy above, later subglabrous, scabrous beneath from more numerous, articulate, weakly flexuous, stiff hairs, mostly mixed with occasional, thin, flexuous or intertwined hairs, often white-arachnoid-hairy along margin, oblong- obovate or oblong-lobate, like teeth on leaves with small, thin, white cusp, sinuate to spiny-toothed, very short-petiolate, upper ones subsessile, smaller; apical leaves densely crowded below capitulum. Capitula aggregated 2—3(5), rarely solitary at stem tips and axillary, less numerous on plant. Involucre oblong, 10-13 mm long and 4-5 mm in dia; involucral bracts with quite long, awnlike spine. Florets lilac-pink, almost not surpassing involucre; corolla 7.5 mm long. All achenes alike, smooth, dark sandy, usually with reddish-brown spots, (2.5) 2.8-3.0 mm long and 0.9-1.0 mm wide; pappus 3.5—4.0 mm long, inner pappus bristles 5-6 mm long, strongly thickened at base. Flowering April to May. (Plate XIII, Fig. 2; Plate XX, Fig. 2). Compacted sandy, rubbly-sandy places in depressions and hollows of sand ridges atop outlier ridges in desert and on gentle gypsiferous slopes in foothills.—Soviet Central Asia: Kara-Kum, Kzyl-Kum (areas bordering Amu-Darya), Amu-Darya (Karakul, Charshanga), Pamiro- Alai Region (Termez). General distribution: Iran. Described from Iran (area between Kashan and Nasrabad, vicinity of Kerman, area between Shakhrud and Nishapur). Type in Geneva. Note. A comparison published by M.G. Popov of isotypes of Cen- taurea vvedenskyi M. Pop. with authentic specimens of O. minima (Boiss.) Briq. has shown that the former species is very close to the latter and can hardly be considered a separate species. The reference to the whitish color of the florets in C. vvedenskyii M. Pop. needs to be verified. O.V. Czerneva (Fl. Uzbek. IV (1962) 412, pl. XXXVIII, Fig. 2) recognizes O. vvedenskyii (M. Pop.) Tschern. as a separate species and indicates that it differs from O. minima (Boiss.) Briq., which, according to her, does not grow in Soviet Central Asia, by having larger achenes, a longer pappus, and whitish florets. Ng NYA CaAW ZA 2 WN Wane YN on ) 5 A Plate XX. 351 1—Schischkinia albispina (Bge.) Ijin; 2—Oligochaeta minima (Boiss.) Briq.; 3—Amberboa bucharica Ijin. 352 349 A form has been reported with lyrately divided leaves: f. lyratisecta Czer.—foliis lyratisectis (Irania, inter oppida Tegeran et Kerman, Naugumbez. 19. IV. 1906, No. 84, K. Adamova). GENUS 1618. Schischkinia Ijin’? Iljin in Fedde, Repert. XXXVIII (1935) 73; Nikitin in Fl. Turkm. VII (1960) 260. Capitula heterogamous, small, with few florets, at tips of greatly shortened branches, on very short peduncles or sessile. Involucres oblong-cylindrical, 4—5 mm in dia, glabrous; involucral bracts many- rowed, imbricate, herbaceous, with inconspicuous veins, greenish, from ovate-lanceolate outer to linearly lanceolate innermost, with white membranous border, terminating in stiff, scarious, nonspiny, subulate cusp; innermost involucral bracts often lilac at tip. Receptacle more or less deeply alveolate; alveoles bordered with thin short cilia. All flo- rets tubular, yellow, not surpassing involucre, heterogeneous, central florets (disk florets) bisexual with 5-fid corolla, basally covered with few, short glandular hairs; peripheral florets (ray florets) sterile, lack- ing staminodes and rudiment of style, 1-rowed, with 4—S-fid glabrous corolla. Anthers with oblong, undivided and obtuse basal appendages, apical appendages obtuse; stamen filaments flattened, covered with papillae. Stigma very short-woolly, with collar of hairs at base, not exserted from anther tube, bilobed, lobes not divergent, short. Achenes oblong-obovate or obovate, narrowed toward base, 3.5—4.0(4.5) mm long, truncate above, with toothed crown at tip, compressed, glabrous, smooth, lustrous, peripheral immature, transversely rugose; hilum lat- eral, deep slit-like; pappus longer than achene, white or slightly ochre- ous, persistent, double, outer one of more numerous, gradually longer from outer to inner serrate-scabrous hair-like bristles, connate in ring at base, inner one comprising single serrate-scabrous, stiff, scarious bristle somewhat thicker below, surpassing outer pappus bristles and arising from mostly irregularly toothed upper edge of ring. Annual herbs with very short stem, usually branched from base, or stemless and with undivided leaves, bearing white, thin, stiff bristles on margin. A monotypic genus. 1. S. albispina (Bge.) Iljin in Fedde, Repert. XX XVIII (1935) 73; Nikitin in Fl. Turkm. VII, 262.—Microlonchus albispinus Bge. Delect. 'Treatment by S.K. Czerepanov. ?Named in honor of the eminent Soviet botanist, florist, and systematist, B.K. Schischkin (1886-1963). 353 350 Sem. hort. Dorpat. (1843) 8; O. and B. Fedtsch. Perech. Rast. Turk. IV, 281.—Centaurea albispina (Bge.) B. Fedtsch. Rast. Turk. (1915) mS Annual. Plants small, with hemispherical crown, stemless or almost stemless, with short stems, usually branched from base. Stem and its branches whitish, finely white-pubescent. Leaves oblong- lobate, obtuse, narrowed into petiole, entire or crenate, with white, thin, stiff, mostly palmately branched bristles on margin, glabrous on both sides or often with scattered tiny whitish hairs beneath. Capitula aggregated in small numbers at tips of strongly reduced branches, very short-pedunculate or sessile, surrounded by few apical leaves, far surpassing capitulum. Involucre oblong-cylindrical, 12-14 mm long and 4—5 mm in dia; involucral bracts herbaceous, white-membranous along margin and at tip, terminating in stiff, membranous, nonspiny, subulate cusp; innermost bracts lilac at tip. Florets yellow. Inner achenes 3.5-4.0 (4.5) mm long and 2.0—2.5 mm wide; pappus 6—7 mm long; inner pappus bristles to 9 mm long. Flowering May. (Plate XX, Fig. 1). Deserts along lower mountain slopes.—Soviet Central Asia: Kara- Kum (southern Ustyurt, Kryk-Sekiz collective farm), Amu-Darya, Kyzyl-Kum, Pamiro-Alai Region (western part). Aralo-Caspian Re- gion (extreme southeast), Lake Balkhash Region (Muyun-Kums). Gen- eral distribution: Iran-Afghanistan (Iran, Afghanistan and Baluchistan). Described from Lehmann’s collections from desert in Aral Sea region. Type and isotypes in Leningrad. Note. Oligochaeta leucosmerinx Rech. f. and KGie (in Dan. Biol. Skr. 8, 2 [1955] 184, Fig. 126 (symb. Afgh. Il) is, apparently, simply a synonym of S. albispina (Bge.) Ijin. GENUS 1619. Aetheopappus Cass." * Cass. in Dict. Sc. nat. L (1827) 250; LI (1827) 53.—Centaurea sect. Aetheopappus (Cass.) DC. Prodr. VI (1837) 574; O. Hoffm. in Pflanzenfam. IV, 5 (1893) 330 p. p. Capitula heterogamous, rather large, with many florets, solitary at tip of stem. Involucre ovate-globose or hemispherical, 15—25 mm in dia; involucral bracts many-rowed, imbricate, outermost entirely mem- branous, silver-white to dark reddish-brown, fimbriate-ciliate, more or less densely short-pubescent like appendages of outer and middle bracts, others.coriaceous, smooth, greenish (yellowing upon drying) from ovate 'Treatment by D.I. Sosnowsky. *From the Greek words aethes—unusual, and pappus—beard. 351 and oblong-ovate middle to oblong-linear innermost; appendages of middle and inner bracts large, not decurrent but entirely overlapping bracts, membranous, reddish-brown to dark reddish-brown, ovate or broadly-ovate, fimbriate-ciliate. Receptacle setose. Florets bright-pink (in USSR species) less often yellow, variable; corolla glabrous; central florets (disk florets) bisexual, tubular, many-rowed, slightly surpassing involucre, with deeply 5-fid corolla, peripheral florets (ray florets) sterile, lacking staminodes and rudiment of style, tubular-infundibuli- form, one-rowed, strongly broadened, 2 times as long as involucre, with deeply (5) 6—8-fid corolla. Anthers with small obtuse basal ap- pendages, apical appendages obtuse; stamen filaments flattened, cov- ered with papillae. Stigma short-bilobed, almost entirely exserted from anther tube. All achenes alike, oblong, about 6 mm long, truncate above, with small-toothed crown, compressed, smooth, glabrous, hi- lum lateral, pappus longer than achene, ochraceous, persistent, simple, of numerous, short, hairy bristles, gradually longer from short outer to inner ones, connate into ring at base. Perennial herbs with undivided or pinnately incised leaves. 354 Type of genus: A. pulcherrimus (Willd.) Cass. This genus comprises at least five [sic., three] species, which grow in the Caucasus and Asia Minor. in Root collar covered with undivided, rather broad remnants of peti- oles of basal leaves; appendages of involucral bracts one-colored; dark reddish-brown or reddish-brown. .............:.ccceeeceesceeneeeeeeeeeeteees Maateiva el. $8 un aes reed baaek Bein, 2ihe 1. A. vvedenskii (Sosn.) Sosn. Root collar covered with fibrous remnants of petioles of basal leaves; appendages of involucral bracts bicolorous: outer silver- white or light stamineous, middle and inner reddish-brown ......... 2 Leaves, except some basal and uppermost, pinnately incised, with 3-7 pairs of ovate-oblong or ovate-roundish, decurrent lateral seg- ments, divergent at rightyangle to rachisy:.3.25.0... 222%. Poe Leaves lanceolately linear or linear, undivided or some basal and lower cauline leaves pinnatifid with fewer lanceolate or linear, acuminate lobes, divergent at acute angle to rachis, less often all cablinesleaves pinnatelydivided) 2:02 88k. LEE. es: 1. A. vvedenskii (Sosn.) Sosn. in Fl. Gruzii, VIII (1952) 550; in Zam. po. Sist. i Geogr. Rast. Tbil. 17 (1953) 15.—A. pulcherrimus var. peduncularis Somm. and Lev. in Tr. Peterb. Bot. Sada, XVI (1900) 273.—A. raddeanus (Trautv.) Sosn. ex Grossh. Opred. Rast. Kavk. 355 352 (1949) 493 p. min. p.; Kolak. Fl. Abkhazii, IV, 283.—A. fedorovii Charkevicz in Bot. Zhurn. SSSR, XXXVI, 4 (1951) 400.—Psephellus vvedenskii Sosn. in Grossh. Fl. Kavk. IV (1934) 201 in Zam. po Sist. i Geogr. Rast. Tbil. 14, 13; Grossh. Opred. Rast. Kavk. 492.—Exs:.: Pl. Abchaz. exs. No. 259. Perennial. Rhizome oblique, thin; root neck covered with dark brown, undivided, rather broad, remnants of petioles of basal leaves. Stem 40-50 cm high, erect, simple, arachnoid-hairy, leafy to middle. Leaves on both sides, particularly beneath, densely floccose-arach- noid-hairy; basal leaves undivided, linearly lanceolate or lanceolate, long-petiolate, about 15 cm long, to 2.5 cm wide, petiole to 8 cm long; cauline leaves lyrately pinnately parted, with oblong, acuminate termi- nal lobe considerably larger than lateral ones; lateral lobes 1—3 pairs, oblong, acuminate, coalescing mutually and with terminal lobe, re- duced toward leaf base. Capitula large. Involucre hemispherical, to 2.5 cm in dia; appendages of involucral bracts broadly ovate, all reddish- brown or dark reddish-brown. Florets violet-pink. Flowering July to August. Meadows and stony slopes of subalpine zone.—Caucasus: Ciscaucasia (Main Range), Western Transcaucasia (Abkhazia). En- demic. Described from Great Bambak Mountain. Type in Tbilisi; isotype in Leningrad. 2. A. caucasicus Sosn. in Zam. po Sist. i Geogr. Rast. Tbil. 17 (1953) 14, descr. latina; in Fl. Gruzu, VIII (1952) 550, descr. georgica; Karjagin in Fl. Azerb. VUI, 448.—A. pulcherrimus var. foliosus Somm. and Lev. in Tr. Peterb. Bot. Sada, XIV (1900) 272.—A. pulcherrimus auct. p. p. non Cass.: Boiss. Fl. or. III (1875) 603; Grossh. Fl. Kavk. IV, 204; Opred. Rast. Kavk. 493.—Exs.: GRF No. 320. Perennial. Rhizome more or less thick, long; root collar densely covered with brown fibers. Stem 7—20(35) cm high, simple, erect or assurgent at base, tomentose, usually leafy throughout. Leaves short- petiolate, on both sides, especially beneath, densely grayish-arachnoid- tomentose, occasionally subglabrous, leaves of nonflowering shoots and some basal leaves sometimes undivided, ovate-lanceolate others pinnately incised, lanceolate, with 3—7 pairs of ovate-oblong or ovate- roundish, indistinctly smooth-toothed or sinuate-toothed, decurrent, lateral segments at right angle to rachis, terminal lobe somewhat larger. Capitula large. Involucre ovate-globose, to 2.5 cm in dia; outer involu- cral bracts silver-white, membranous, short fimbriate-ciliate, middle and inner ones with reddish-brown, membranous, fimbriate-ciliate ap- pendage. Florets bright pink. Flowering July to August. 355 Rocks and rubbly places of alpine and subalpine zones.—Caucasus: Ciscaucasia, Eastern and Western Transcaucasia (area of Main Caucasus Range and Adzharo-Imeretian Range). Endemic. Described from Kazbeg District of Georgian SSR. Type in Tbilisi. 3. A. pulcherrimus (Willd.) Cass. in Dict. Sc. nat. LI (1827) 54; Boiss. Fl. or. III, 603 p. p.; Grossh. Opred. Rast. Kavk. 493 p. p.; Sosn. in Zam. po Sist. i Geogr. Rast. Tbil. 17, 15; Karjagin in FI. Azerb. VIII, 449.—A. pulcherrimus var. denticulatus C. Koch in Linnaea, XXIV (1851) 422.—A. pulcherrimus var. raddeanus (Trautv.) Bordz. in Vestn. Tifl. Bot. Sada, Nov. Ser. 5 (1931) 65; Grossh. FI. Kavk. IV, 204.—A. raddeanus (Trautv.) Sosn. ex Grossh. Opred. Rast. Kavk. (1949) 493 p. p.—Centaurea pulcherrima Willd. Sp. pl. Ill, 3 (1803) 2298; DC. Prodr. VI, 574; Ldb. Fl. Ross. II, 2, 694.—C. raddeana Trautv. in Tr. Peterb. Bot. Sada, II (1873) 475.—C. pulcherrima var. intermedia Trautv. op. cit. [V (1876) 381.—Ic.: Rchb. Pl. crit. VIII (1838) tab. 716; Karjagin, op. cit. Plate XLV, Fig. 2. Perennial. Rhizome oblique, thin or more or less thick; root neck densely covered with brown fibers. Stem to 60 cm high, erect or as- surgent at base, simple or sometimes with 1-2 branches, occasionally stem short, 10-15 cm high. Leaves green or grayish-green above, slightly arachnoid-hairy, sometimes later becoming glabrous, densely grayish-arachnoid-hairy-tomentose beneath, occasionally glabrous on both sides, lanceolately linear or linear, serrate-toothed or almost entire, very often some basal and lower cauline leaves lyrately di- vided, with large, lanceolate, acuminate terminal lobe and few, small, linearly lanceolate or lanceolate lateral lobes, at acute angle to rachis, occasionally almost all cauline leaves pinnately parted. Capitula large. Outer involucral bracts silver-white, membranous, short-fimbriate-cili- ate; middle and inner bracts with reddish-brown, membranous, ovate, fimbriate-ciliate appendages. Florets bright pink, very rarely white. Flowering July to August. Meadows in subalpine and alpine zones.—Caucasus: Dagestan, Eastern Transcaucasia (Lesser Caucasus), Southern Transcaucasia. General distribution: Armenia and Kurdistan, Asia Minor (northeast- ern part). Described from “Orient.” Type in Berlin. 35) ee) 354 GENUS 1620. Cheirolepis Boiss. ? Boiss. Diagn. pl. or. ser. I, 10 (1849) 106; Czer. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XX (1960) 466.—Centaurea sect. Mesocentron (Cass.) DC. Prodr. VI (1837) 592 p. p. min.—Chartolepis subgen. Cheirolepis (Boiss.) Jaub. and Spach. Ill. pl. or. III (1847-— 1850) 136.—Centaurea sect. Seridia (Juss.) Benth. in Benth. and Hook. f. Gen. pl. II (1873) 481 p. p. non DC.—Phaeopappus sect. Cheirolepis (Boiss.) Boiss. § 1. Eucheirolepidei Boiss. Fl. or. III (1875) 594.— Centaurea sect. Cheirolepis (Boiss.) O. Hoffm. in Pflanzenfam. IV, 5 (1893) 330. Capitula heterogamous, medium-sized, with many florets, solitary at tips of stem and few lateral branches. Involucre ovate or oblong- ovate, 8-20 mm in dia, subglabrous; in volucral bracts many-rowed, densely imbricate, coriaceous, smooth, greenish (yellowing when dry), from triangular and ovately triangular outer ones to oblong-linear in- nermost; appendages of outer and middle involucral bracts small, not decurrent on bracts, coriaceous, brownish, with palmately divergent spines, of which middle spine usually longer than lateral ones; append- ages of inner bracts small, membranous, irregularly toothed. Recep- tacle bristly, variable; corolla glabrous; central florets (disk florets) bisexual, many-rowed; peripheral florets (ray florets) sterile, lacking staminodes and rudiment of style, with 4—5-fid corolla, one-rowed, not enlarged. Anthers with small basal appendages and acute apical ap- pendages; stamen filaments flattened, covered with papillae. Stigma very short-woolly, with collar of hairs at base, united (its branches connate to tip) or scarcely bilobed, entirely exserted from anther tube. All achenes alike, oblong-ellipsoidal, 4-7 mm long, truncate, with small- toothed crown at apex, compressed, glabrous, hilum lateral; pappus longer than achenes, whitish or ochraceous, persistent, simple, of many irregular rows of flattened, plumose, hair-like, bristles, gradually elon- gated from outer to inner, connate in ring at base. Perennial herbs, with erect, arcuately ascending, or assurgent stem, simple or with few branches, not winged, leafy to tip, covered with arachnoid hairs or almost tomentose pubescent. Basal and lower cauline leaves petiolate, others sessile, not decurrent on stem, all entire to sinuate-pinnately lobed, scabrous or scabrous-lanate from spiny or long-articulate hairs to finely arachnoid tomentose. Type of genus: C. drabifolia (Sibth. and Smith) Boiss. Of the nine species in this genus belonging to two phylogenetic 'Treatment by S.K. Czerepanov. "From the Greek words cheir—wrist, and /epis—scale; named for the palmately divided appendages of the involucral bracts. 2352) Cyaan at x é Af \\ i Aw ‘)} 1 Se AWA Sag \\ Ne S Sue | \\ Yj My = “yg | a ! : g Rife NY Vis ; \ 4 W, (77 <= J SE — 497 11. 12: 13. eit AN. ce AR RRS eg 104. C. salonitana Vis. Middle involucral bracts at apex with spine, more than 1 cm long, lateral’ cilia usually longer (16 mium)\:..22 eee ere eee. 6. Spine of middle involucral bracts 1.5—3.5 cm long, thick (near base 2.5—6.0 mm thick), concave above, longitudinally grooved; involu- ere 203 Ommmin diag ete. Meas. 107. C. sosnovskyi Grossh. Spine of middle involucral bracts 1—3 cm long, thinner (usually to 1.5—2.0 mm thick at base), flat or convex above; involucre 15—23 TAM NG ay; coats year ace an Nae es 106. C. reflexa Lam. Floretstdark redieeiun.. 287. Sey hereafter ee Ley 8. Florets pink of various shades, less often reddish ...................... Su Involucral bracts with very large (almost entirely covering them), blackish-brown appendages; capitula always solitary; pappus simple Xe seen A hee ak rien hanced: ean Dre ei Suits 109. C. kotschyana Heuff. Appendages of involucral bracts in form of very narrow brownish border; stems usually many-headed; pappus double ..................eeeee adahces AU, ca Meese cats SLATE nt Meda ued netic: 105. C. rubriflora Illar. All leaves undivided and entire, less often with 1—2 small lobes neambase ir. BeBe. esc hk A. 111. C. integrifolia Tausch. Leaves pinnately parted or lobed, less often lower ones entire .... Appendages of middle involucral bracts at apex with recurved spine, f-40hmm lomeibrownishe.ui! Bi.94 We ae eee Oe Appendages of involucral bracts at apex lacking spine or with very short spine to S—7 mm long, usually dark brown or almost black .... Apical spines of middle involucral bracts 7-16 mm long, thinner, flatcondconvexabove: (Trahseauicasia)) is.) 2222s ieee econ 2p Tes asses ssc lino eee oa cso celas 117. C. carduiformis DC. Apical spines of middle involucral bracts 15—40 cm long, thick, usually concave above, longitudinally grooved ..............cceseeseeeeeee Beaune mene 2 DEMON ica), Lea AS 118. C. sintenisiana Gand. Appendages of middle involucral bracts rather large, (1.8)2.0— 4.0(6.0) mm long, excluding part decurrent on bract margin; pe- ripheral florets distinctly enlarged ..............ccccecscseseesesseeseeeeeeees 1: Appendages of middle involucral bracts very small, to 1.5(1.8) mm long, excluding part decurrent on bract margin and apical spinule; peripheral florets not surpassing middle ...............cccsceeeseeseeneeeees 1S. Appendages of middle involucral bracts at apex with short recurved spine (or spinule) 2.5—6.0 mm long (Transcaucasia) ..............:06 Dear Weta ene ck aan a ay OE OT LS, Se 116. C. glehnii Trautv. Appendages of middle involucral bracts at apex lacking spine, rarely 498 with straight spinule to 2 mm Jong ............ceceeceeneeeeeeeeeeeeeeeeeees 14. 14. | Achenes 5—6 mm long, with pappus 4.5—6.0 mm long (Caucasus) PPE Rc I RET wd tet MaGEE ase hee ae BAER 112. C. ossethica Sosn. + Achenes 3.5—4.5 mm long, with pappus 4—5 mm long (forest zone of European part of the USSR and Siberia) ............ ce eeeeeeeeees DAIEN: RINIERRE ODEs ES. LIOR AS SIO Ee eal 110. C. scabiosa L. 15. | Appendages of middle involucral bracts at apex with short recurved spine or spinule 2—6 mm long; achenes 5.0—6.5 mm long, with pappus 6-8 mm long (Southern Transcaucasia) ...............cccceeeees bY, Pan. ROROEEES, AOE EG ol 115. C. pseudoscabiosa Boiss. and Buhse + Appendages of middle involucral bracts at apex lacking spine, sometimes with spinule to 2 mm long, but then achenes smaller (3554 Senna PO ae it ee 16. 16. Involucral bracts at apex lacking or with very small appendage to 0.8(1.0) mm long, usually with few very short cilia along margin (steppe and semi-steppe zone) ...............4 114. C. adpressa Ldb. + Involucral bracts with apical appendage, 0.8-1.5 mm long, with fairly long cilia along margin (mostly plants of forest-steppe zone) TE TAS TEE ACRE US ER 113. C. apiculata Ldb. Section 1. Acrocentron (Cass.) DC. Prodr. VI (1837) 586. p. p.— Centaurea sect. Acrocentron *Euacrocentrae flaviflorae Boiss. FI. or. III (1875) 618.—Florets yellow, less often dark red; involucral bracts with apical appendages in form of narrow, more or less pectinate- ciliate border, often terminating in cusp or spine; achenes with more or less equally long double pappus. Type of section: C. collina L. 104. C. salonitana Vis. in Flora, XII, 1 (1829) 23; Boiss. FI. or. III, 666; Schmalh. Fl. II, 127; Grossh. Fl. Kavk. IV, 217 and Opred. Rast. Kavk. 499.—C. centauroides auct. non L.: MB. FI. taur.-cauc. II (1808) 359; Ldb. Fl. Ross. II, 2, 708.—C. latisquama DC. Prodr. VI (1837) 589.—C. pelia auct. non DC.: Stank. and Tal. Opr. Vyssh. Rast. (1949) 676.—Acrocentron salonitanum (Vis.) A. and D. Léve in Bot. Notis. CXIV, 1 (1961) 44.—Ie.: Vis. Fl. Dalm. II (1847) tab. 13; Rchb. 1c. Fl. Germ. XV (1852) tab. 793; Fedtsch. and Fler. Fl. Evrop. Ross. Fig. 1050; Prod. Fl. Roman II (1923) tab. 35; Stojan. and Stef. Fl. Bulg. (1925) 1187; Javorka and Csapody, Iconogr. Fl. Hung. (1933) 550, fig. 3941.—Exs.: Fl. Dalm. exs. No. 89; Schultz. Herb. norm. nov. ser. No. 1595; Fl. exs. austro-hung. No. 235; Hayek, Cent. exs. crit. No. 10; GRF No. 572. *Asterisk not explained in the original—General Editor. 499 496 Perennial. Plants to 80—100 cm high, with short, crisped and papil- liform hairs, often mixed with arachnoid-hairy tomentum, more or less scabrous, green. Stems solitary or few, upright, more or less branched. Leaves bipinnately parted to pinnately lobed, lower ones rather long- petiolate, usually with longer and denser hairs, uppermost leaves subsessile, sometimes with undivided and entire lamina; lateral seg- ments or lobes oblong to almost linear, terminal segment usually not larger than lateral ones. Capitula usually aggregated in corymbs. In- volucre 13—20 mm in dia and 18—25 mm long, glabrous or subglabrous; outer and middle involucral bracts short-ciliate in upper half (usually cilia to 1 mm long), obtuse, rounded, with short spine or with recurved spine to 1 cm-long; innermost involucral bracts with almost round or kidney-shaped membranous apical appendage. Florets yellow, periph- eral ones not enlarged. Achenes sparsely short-hairy, 4-5 mm long; pappus whitish or smoky, 7—9 mm long, scale-like bristles of inner pappus 1/4—1/3 as long as longest outer pappus bristles. Flowering June to August. Steppes, slopes, often as weed in wastelands, near roads.—Euro- pean Part: Bessarabia, Black Sea Region, Lower Don (western part), Crimea; Caucasus: Ciscaucasia (western part), Western Transcaucasia (in Novorossiisk area). General distribution: Central Europe (south eastern part), Balkans. Described from Yugoslavia (Dalmatia). Type in Padua. Note. On the southern coast of Crimea (especially near Sudak) and on the Taman Peninsula, together with the typical form with rela- tively short spines or entirely without them, one finds plants with longer spines.—C. salonitana var. macracantha (DC.) Boiss. and Heldr. in Boiss. Diagn. Pl. or. ser. I, 3 (1856) 78. This variety apparently is a weakly separated, more southern, ecogeographical race, intimately related to typical C. salonitana. The known hybrids are: C. salonitana x C. orientalis and C. salonitana Xx C. adpressa. 105. C. rubriflora Illar. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVIII (1957) 309.—C. salonitana var. purpurea Ferm. ex Stojan. and Stef. Fl. na B’lg. ed. 3 (1948) 1203. Perennial. Plants to 20-30 cm high, with short, crisped and papil- liform hairs, more or less scabrous, green. Stems solitary, upright, with few lateral branches. Leaves pinnately parted to lobéd, uppermost subsessile, undivided, lower ones fairly long-petiolate; lateral segments or lobes oblong, entire, terminal lobe usually not larger than lateral ones. Involucre 15—22 mm in dia and 22—24 mm long, glabrous or subglabrous; outer and middle involucral bracts short-ciliate in upper half, at apex with short, recurved spine; innermost bracts with almost 500 497 round, short, and irregularly fimbriate apical appendage. Florets dark red. Achenes sparsely short hairy, 4.5—5.0 mm long, pappus 5—9 mm long. Flowering June to August. Stony slopes, along roads, up to lower montane zone.—European Part: Crimea (southeastern part). General distribution: Balkans (north- eastern part). Described from Crimea (Karadag Mountain). Type in Leningrad. Note. Possibly this is a color variant of the previous species and needs confirmation on the basis of more extensive material. 106. C. reflexa Lam. Encycl. meth. I (1783) 675; MB. FI. taur.- cauc. II, 359; DC Prodr. VI, 589 p. p.; Ldb. Fl. Ross. II, 2, 708 p. p.; Boiss. Fl. or. III, 667 p. p.; Grossh. Fl. Kavk. IV, 218 and Opred. Rast. Kavk. 499. Perennial or biennial. Plants to 70-80 cm high; with short, crisped and papilliform hairs, often mixed with arachnoid tomentum, more or less scabrous, green or grayish-green. Stems solitary, erect, branched mostly in upper part. Lower leaves rather long-petiolate, pinnately parted, less often lobed, upper fewer, strongly reduced, subsessile; lateral segments or lobes linear to oblong, more or less pinnately lobed, less often undivided; terminal segment not larger than lateral ones. Capitula often aggregated in corymbs. Involucre 15—23 mm in dia, 18— 27 mm long, glabrous or subglabrous; outer and middle involucral bracts in upper half pectinate-ciliate (with 1-4 mm-long cilia), termi- nating in recurved spine, 1-3 cm long and up to 1.5—2 mm thick (at base) in middle bracts; upper (inner) surface of spine convex or flat; innermost involucral bracts oblong, with almost round membranous, long-fimbriate apical appendage. Florets yellow, peripheral ones not enlarged. Achenes 4-5 mm long, with rather dense, connate hairs; pappus 7-9 mm long, whitish or weakly smoky; scale-like inner pap- pus bristle 1/4—1/3 as long as longest outer pappus bristle. Flowering June to July. Steppes, on stony slopes; up to lower montane zone.—Caucasus: Dagestan (southern part), Eastern Transcaucasia (Kura River basin), Western Transcaucasia. Endemic. Described from “East” (“Levant”) without precise mention of locality, probably from Georgia. Type in Paris. Note. Apparently, it splits up into several, still inadequately stud- ied ecogeographical races, differing mostly in the form and nature of the pubescence of the leaves, as well as the length of the cilia of the involucral bracts. The hybrid C. reflexa x C. adpressa, which usually shows the normal development of achenes, is, known from Dagestan. 50 —_ 498 107. C. sosnovskyi Grossh. in Beih. Bot. Centralbl. XLIV (1928) 246; Fl. Kavk. IV, 218 and Opred. Rast. Kavk. 499.—C. reflexa auct. non Lam.: C.A.M. Verzeichn. (1831) 65; DC. Prodr. VI, 589 p. p.; Ldb. FI. Ross. II, 2, 708 p. p.; Boiss. Fl. or. III, 667 p. p.—C. reflexa var. macracantha C. Koch in Linnaea, XXIV (1851) 429.—C. reflexa var. grandiflora Grossh. in Tr. Tifl. Bot. Sada, Nov. Ser. 1 (1920) 41. Perennial. Plants to 70-80 cm high, with crisped and papilliform hairs, sometimes mixed with arachnoid tomentum, more or less sca- brous, green or somewhat grayish. Stems solitary, upright, usually with few lateral branches, often simple. Lower leaves rather long-petiolate, pinnately parted, upper ones fewer, strongly reduced, sessile or sub- sessile; lateral segments linear to oblong, pinnately lobed or undi- vided; terminal segment not longer than laterals. Capitula solitary or aggregated in lax corymb. Involucre 20-30 mm in dia and 20-28 mm long, glabrous or subglabrous; outer and middle involucral bracts in upper half pectinate-ciliate (cilia 2-6 mm long), terminating in long spine, 1.5—3.5 mm long and 3—6 mm wide (at base) in middle bracts; upper (inner) surface of spine concave; innermost bracts ovate-oblong, with almost round, pectinate-fimbriate appendage. Florets yellow, pe- ripheral ones not large. Achenes 5-6 mm long, with rather dense, appressed hairs; pappus 8—10 mm long, whitish, scale-like inner pap- pus bristles 1/4—1/3 as long as longest outer pappus bristle. Flowering June to July. (Plate XXV, Fig. 2.) Stony slopes, rocks; up to lower montane zone.—Caucasus: East- ern Transcaucasia (southern part), Southern Transcaucasia, Talysh. General distribution: Armenia and Kurdistan. Described from Nakhichevan ASSR (between Ordubad and village of Magri). Type in Tbilisi. Note. It is not ruled out that the name C. reflexa Lam. pertains to this and not the previous species. Section 2. Orientales (Hayek) Tzvel. comb. nova.—Centaurea sect. Acrocentron D. Orientales Hayek, Centaurea-Art. Osterr.-Ung. (1901) 53.—Centaurea sect. Acrocentron subsect. Orientales (Hayek) Tzvel. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX (1959) 433.—Capitula yellow, less often dark red; involucral bracts with very large, lanceolate, ovate or roundish, weakly decurrent apical append- age, ciliate or fimbriate, and often terminating in a short cusp; achenes with more or less as long pappus; pappus double or simple (because of reduction of inner pappus). Type of section: C. orientalis L. 502 499 108. C. orientalis L. Sp. pl. (1753) 913; MB. FI. taur.-cauc. I, 351 and III, 592; DC. Prodr. VI, 587; Ldb. Fl. Ross. II, 2, 707; Boiss. Fl. or. Ill, 666; Schmalh. FI. II, 127; Grossh. Fl. Kavk. IV, 217 and Opred. Rast. Kavk. 499.—C. coelolepis Fisch. and Mey. ex Heynh. Nomen. bot. hort. II (1846) 130, nom. nud.—Acrocentron orientale (L.) A. and D. Léve in Bot. Notis. CXIV, 1 (1961) 44.—Ie.: Rchb. Ic. Fl. Germ. XV (1852) tab. 790; Fedtsch. and Fler. Fl. Evrop. Ross. Fig. 1048; Stojan. and Stef. Fl. Bulg. 1183; Javorka and Csapody, Iconogr. Fl. Hung. (1933) 550, fig. 3942.—Exs.: Fl. exs. austro-hung. No. 656; Fl. cauc. exs. No. 175. Perennial. Plants to 80-100 cm high, with very short papilliform hairs, sometimes mixed with thin arachnoid-hairy-tomentum, more or less scabrous, green. Stems solitary or few, upright, usually with few lateral branches. Leaves pinnately parted or lobed, sometimes some bipinnately incised, except uppermost rather long-petiolate; lateral leaf segments oblong to narrowly linear, sessile or basally narrowed into petiolule, entire or irregularly toothed, subacute, sometimes with short spinule; terminal segment often longer than lateral ones. Capitula soli- tary or less numerous at tips of stems and lateral branches. Involucre 15-24 mm in dia and 17-25 mm long, glabrous; appendages thin- coriaceous, brownish-yellow or yellowish, weakly decurrent, almost covering bract, broadly lanceolate to almost round in outer bracts, pectinate-ciliate (cilia shorter than width of appendage), usually with upto 2-3 mm-long apical spinule, considerably smaller in innermost bracts, almost round, short-fimbriate, lacking spinule. Florets yellow, peripheral ones almost not enlarged. Achenes 4—5 mm long, hairy; pappus almost as long as achenes, usually with brownish-gray (smoky) tinge; inner pappus bristles scale-like, 1/5—1/4 as long as longest outer pappus bristle. Flowering June to September. Steppes, stony slopes; up to lower montane zone.—European Part: Middle Dnieper, Volga-Don (southern part), Bessarabia, Black Sea Region, Lower Don, Crimea; Caucasus: Ciscaucasia. General distri- bution: Centrral Europe (southeastern part), Balkans (northern part). Described from European part of USSR (“hab. in Sibiria’”). Type in London. Note. Two varieties of this species are known: C. orientalis var. brachylepis Fisch. and Mey. Ind. Sem. hort. Petrop. VII (1840) 44— with obtuse appendages on the lower and middle involucral bracts and a terminal spinule no longer than the lateral cilia, and C. orientalis var. macrolepis Fisch. and Mey. loc. cit. 44—with elongated, more or less broadly lanceolate appendages on the lower and middle involucral bracts and a prickly terminal spinule distinctly longer than the lateral cilia. Although their ranges almost overlap, the second variety, on the 503 500 average, is more southern and typical of arid localities, constituting a separate but morphologically very weakly distinguished, ecogeographi- cal race. C. orientalis easily hybridizes with other species of the genus, especially with C. apiculata Ldb., C. pseudomaculosa Dobrocz., and C. salonitana Vis. The more common fertile hybrids of C. apiculata Ldb. x C. orientalis L. (cf. EM. Lavrenko in Bot. Mat. Gerb. Glavn. Bot. Sada, IV, 1923, 33-38) are known under the following names: C. rigidifolia Bess. in DC. Prodr. VI (1837) 587.—C. rubescens Bess. in DC. op. cit. 587.—C. calocephala auct. fl. Ross. non Willd.—C. orientalis L. var. calocephala Schmalh. FI. Yugo.-Zap. Ross. (1886) 335.—C. orientalis L. var. rubescens (Bess.) Schmalh. op. cit. 335.— Stem and leaves as in C. orientalis but pubescence usually much stron- ger. Involucre 15-25 mm wide; appendages of outer and middle in- volucral bracts more or less triangular-lanceolate, pectinate-ciliate, with apical spine, various shades of brown but usually darker than in C. orientalis and lighter than in C. apiculata; florets pale yellow, more rarely pinkish (C. rubescens Bess.); achenes as in C. orientalis but smaller, 3.5—4 mm long; pappus 4—5 mm long. 109. C. kotschyana Hevuff. in Flora XVIII, 1 (1835) 245; Dobrocz. in Vizn. Rosl. URSR, 581.—C. heuffelii Rchb. fil. in Rchb. Ic. FI. Germ. XV (1852) 27.—Acrocentron kotschyanum (Heuff.) A. and D. Léve in Bot. Notis. CXIV, 1 (1961) 44.—Ic.: Rchb. Ic. Fl. Germ. XV (1852) tab. 775 and 801; Prod. Fl. Roman. I (1923) 1095 and Cent. Roman. (1930) tab. 3; Stojan. and Stef. Fl. Bulg. (1925) 1183; Javorka and Csapody, Iconogr. Fl. Hung. (1933) 550, fig. 3945.—Exs.: FI. exs. austro-hung. No. 3428; Hayek, Cent. exs. crit. fasc. II, No. 65; Fl. exs. reip. Bohem.-Sloven. No. 383; Fl. Cechoslov. exs. No. 97. Perennial. Plants to 80-100 cm high, with short papilliform hairs mixed with crisped hairs and thin arachnoid-hairy-tomentum, weakly scabrous, green. Stems usually solitary, upright, simple, less often with 1—2 lateral branches. Leaves pinnately parted or lobed, lower ones rather long-petiolate, sometimes undivided and then toothed, upper leaves sessile; lateral segments or lobes oblong to lanceolately linear, usually more or less toothed or lobed, acuminate; terminal segment often larger than lateral ones. Capitula usually solitary. Involucre 22-35 mm in dia and 20-27 mm long, subglabrous; appendages of outer and middle involucral bracts very large, entirely covering bracts, thin-coriaceous, weakly decurrent, oblong-ovate, blackish-brown, long fiunbriate-ciliate (cilia to 6-7 mm long, usually much lighter in color than appendages). Florets dark red, peripheral ones almost not en- larged. Achenes about 5 mm long, sparsely hairy (to subglabrous); 501 pappus 3-4 mm long, consisting of numerous irregular bristles of identical structure. Flowering June to August. (Plate XXV, Fig. 1). Mountain meadows, in upper montane zone.—European Part: Up- per Dniester. General distribution: Centrral Europe (southeastern part), Balkans (northern part). Described from Carpathian Mountains. Type in Vienna. Note. This species very closely approaches C. orientalis through another high-mountain centaury species, C. atripurpurea Willd. Section 3. Lopholoma. —Centaurea sect. Lopholoma (Cass.) DC. Prodr. VI (1837) 580 p. p.—Centaurea sect. Acrocentron C. Lopholomae Hayek. Centaurea-Art. Osterr.-Ung. (1901) 34.—Florets various shades of purple; involucral bracts with apical appendages of different sizes and forms (from broadly ovate to very narrow, some- times almost lacking border) but always strongly decurrent on bracts, pectinate-ciliate, terminating in short or long spine; achenes with more or less as long double pappus. Type of section: type of subgenus. Series 1. Scabiosiformes Dobrocz. in Ukr. Bot. Zhurn. XIX, 1 (1962) 43, descr.; in Bot. Zhurn. Akad. Nauk URSR, IV, 3-4 (1948) 78, nomen.—Polycarpic plants surviving few years, with not thick roots. Appendages of involucral bracts lacking spine, less often with spine to 5 mm long. 110. C. scabiosa L. Sp. pl. (1753) 913; MB. FI. taur.-cauc. II, 350 p. p.; DC. Prodr. VI, 580 p. p.; Ldb. Fl. Ross. II, 2, 700 p. p.; Boiss. Fl. or. III, 655 p. p.; Schmalh. Fl. II, 126 p. p.; Grossh. Fl. Kavk. IV, 218 p. p.; Kryl. Fl. Zap. Sib. XI, 2960 p. p.—C. coriacea Waldst. and Kit. Descr. et icon. pl. rar. Hung. II (1805) 214.—Acrocentron scabiosa (L.) A. and D. Léve in Bot. Notis. CXIV (1961) 44.—Ic.: Waldst. and Kit. loc. cit. tab. 195; Rchb. Ic. Fl. Germ. XV (1852) tab. 774; Fedtsch. and Fler. Fl. Evrop. Ross. Fig. 1046; Syreistsch. Ill. Fl. Mosk. Gub. III (1910) 315; Prod. Fl. Roman. I (1923) 1095; Hegi, Ill. Fl. VI, 2 (1929) Taf. 273, fig. 3; Javorka and Csapody, Iconogr. Fl. Hung. (1933) 550, fig. 3950.—Exs.: Fl. Bohem. and Morav. exs. No. 1164; Fl. exs. austro- hung. No. 972; Pl. Finl. exs. No. 995; GRF No. 1619. Perennial. Plants to 80-100 cm high, with short papilliform hairs mixed with crisped hairs and thin arachnoid-hairy-tomentum, more or less scabrous, green. Stems solitary or few, upright, more or less branched mostly in upper part. Leaves pinnately parted or lobed, some- times some bipinnately incised, lower rather long-petiolate, upper ones strongly reduced, partly sessile; lateral segments oblong to linear, 504 502 terminal one usually not much larger than lateral ones. Involucre (11) 13—22(25) mm in dia and 17—23 mm long, glabrous or thinly arach- noid-hairy tomentose; appendages of outer and middle involucral bracts thin-coriaceous, obtuse triangular, (1.8)2—6 mm long (excluding bor- der, decurrent on bract), blackish-brown or almost black, pectinate- ciliate, with 0.8-2.5 mm long cilia, usually lighter in color than ap- pendage, often with apical spine or cusp; appendages of innermost bracts almost round, membranous, brownish, fimbriate-ciliate. Florets various shades of pink, peripheral ones slightly large. Achenes 3.5—4.5 mm long, sparsely short-hairy; pappus 4-5 mm long, whitish with brownish-gray tinge, scale-like inner pappus bristles 1/5—1/4 as long as largest outer pappus bristles. Flowering June to September. Meadows, scrubs, along roads; up to mid-montane zone.—Euro- pean Part: Karelia-Lapland (southern part), Dvina-Pechora, Volga- Kama, Baltic Region, Ladoga-IImen, Upper Dnieper, Upper Volga, Upper Dniester, Middle Dniester (northern part), Volga-Don (rarely), Trans-Volga (rarely); Western Siberia: Ob (southern part), Upper Tobol, Irtysh, Altai; Eastern Siberia: Angara-Sayans. General distribution: Atlantic Europe, Scandinavia, Centrral Europe, Balkans (northern part). Described from northern Europe. Type in London. Note. It varies greatly in the shape of the leaves, pubescence, and size of the appendages of the involucral bracts (the appendages, as a rule, gradually reduce from northwest to southeast), which has led to the description of a whole series of varieties and forms of this species. These, in our opinion, do not merit special names. However, mention must be made of C. scabiosa var. malenocephala Rupr. (in Beitr. Pflanzenk. Russ. Reichs. IV [1845] 81), from the northwest of the European part of the USSR (vicinity of Leningrad, Lake Onega), with especially large, long-ciliate appendages almost covering the bracts. This variety resembles C. alpestris Hegetschw. (in Hegetschw. and Heer, Fl. Schweiz. [1840] 854), found in the Alps and the Carpathians (Tatry), and is close to C. scabiosa; however, it can hardly be iden- tified with it. On the limestones of the Velikaya River near Pskov, there are plants with almost entire leaves that approach the next species. 111. C. integrifolia Tausch in Flora, XI, 31 (1828) 485; Tzvelev in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX, 435.—C. scabiosa var. integrifolia Korsch. Tent. Fl. Ross. Or. (1898) 242; Kryl. Fl. Zap. Sib. XI, 2961, non Lindem. (1872).—Ic.: Gmel. FI. sibirica, II (1749) tab. 42, fig. 1. Perennial. Plants to 100-120 cm high, with short papilliform hairs mixed with crisped hairs and thin arachnoid-hairy-tomentum, green. 505 503 Stems solitary or fewer, upright, more or less branched in upper part. Leaves undivided and entire, oblong; lower and middle ones gradually narrowed toward base into rather long petiole, acuminate or subobtuse, upper ones sessile or subsessile, sometimes with 1—2 small lobes at base of lamina. Involucre 12—20 mm in dia and 17—22 mm long, gla- brous or finely arachnoid-hairy-tomentose; appendages of outer and middle involucral bracts thin-coriaceous, obtuse, triangular, 1.8—4.0 mm long (excluding border decurrent on bracts), blackish-brown or almost black, with brownish cilia 0.8—1.5 mm long, usually lacking apical spine. Florets pink, peripheral ones slightly large. Achenes 3.5— 4.5 mm long, sparsely short-hairy; pappus 4—5 mm long, usually with brownish-gray tinge, scale-like inner pappus bristles 1/6—1/4 as long as largest outer pappus bristles. Flowering June to September. Birch forests, glades, meadows; up to mid-montane zone.—Euro- pean Part: Volga-Kama (southeastern part), Trans- Volga (eastern part); Western Siberia: Ob Region (southwestern part), Upper Tobol. En- demic. Described from Siberia. Type in Berlin. Note. A distinct Ural-Siberian race, possibly a mutant but with a very well marked range. A quite large number of specimens is avail- able in the Herbarium of the Botanical Institute of the Academy of Sciences of the USSR. 112. C. ossethica Sosn. in Addenda XXVII, 614; in Fl. Gruzii, VIII (1952) 568, diagn. georgica.—C. glehnii auct. non Trautv.: Grossh. Fl. Kavk. IV (1934) 218 p. p.—C. buschiorum Sosn. ex Grossh. Opred. Rast. Kavk. (1949) 500; diagn. rossica, non Psephellus buschiorum Sosn. (1948). Perennial. Plants to 80-100 cm high, with short papilliform hairs mixed with crisped hairs and thin arachnoid-hairy-tomentum, weakly scabrous, green. Stems solitary or few, upright, simple or with few lateral branches. Leaves pinnately parted or lobed, lower ones quite long-petiolate, upper ones sessile or subsessile, sometimes some of them entire, with 2—5 lateral segments on each side, oblong, entire or more or less toothed, terminal segment usually larger than laterals. Involucre 19-23 mm in dia and 14-25 mm long, glabrous or subglabrous; appendages of outer and middle involucral bracts thin- coriaceous, obtuse-triangular, 2.0—-3.5 mm long (excluding border de- current on bracts), blackish-brown, with whitish cilia 2-3.5 mm long, with very short (to 1.0-1.5 mm long) cusp; innermost bracts with al- most round, deeply fimbriate appendages. Florets pink, peripheral ones slightly large. Achenes 5—6 mm long, sparsely short-hairy; pappus 4.5— 6.0 mm long, scale-like inner pappus bristles 1/3—2/5 as long as larg- est outer pappus bristle. Flowering July to September. 506 504 Meadows, stony slopes; in mid- and upper montane zones.— Caucasus: Ciscaucasia (southern part), Dagestan (southern part), East- ern Transcaucasia (northern and western parts), Western Transcaucasia (northern and eastern parts), Southern Transcaucasia (northern part), Talysh. Endemic. Described from southern Ossetia. Type in Tbilisi. Note. It is an ecogeographical race very similar to C. scabiosa L., and replaces this northern species in the upper montane zone of the Caucasus. It also considerably resembles C. glehnii Trautv. and is linked to this species through populations with intermediate charac- ters. 113. C. apiculata Ldb. Ind. Sem. hort. Dorpat. (1824) 3; DC. Prodr. VI, 581.—C. scabiosa auct. non L.: MB. FI. taur.-cauc. II (1808) 350 p. p.; Grossh. Fl. Kavk. IV, 218 p. p.; Kryl. Fl. Zap. Sib. XI, 2960 p. p.—C. scabiosa var. apiculata (Ldb.) Ldb. and var. coriacea Ldb. Fl. Ross. II, 2 (1845) 701; Schmalh. Fl. II, 126.—C. coriacea auct. fl. Ross. non Waldst. and Kit—C. pseudocoriacea Dobrocz. in Bot. Zhurn. Akad. Nauk URSR, IV, 3—4 (1948) 78 and Vizn. Rosl. URSR, 576.— Exs.: Fl. exs. reip. Bohem.-Sloven. No. 891 (sub C. scabiosa L.); F1. polon. exs. No. 843 (sub C. scabiosa var. adpressa Ldb.). Perennial. Plants to 80-100 cm high, with short papilliform hairs mixed with crisped hairs and thin arachnoid-hairy-tomentum, weakly scabrous, green. Stems solitary or few, upright, branched mostly in upper part. Leaves pinnately parted or lobed, sometimes partly entire, lower ones rather long-petiolate, upper ones sessile or sub-sessile; lateral segments oblong to almost linear, entire, more or less toothed or lobed; terminal segment usually not larger than laterals. Involucre 11-17 mm in dia and 17-22 mm long, glabrous or thinly arachnoid-hairy-tomen- tose; appendages of outer and middle involucral bracts thin-coriaceous, obtusely triangular, very small (0.5)0.8—1.5(1.8) mm long (excluding very narrow, to 0.5 mm wide, margin decurrent on bracts), blackish- brown, with darker cilia 0.5—1.2 mm long, usually terminating in spine to 2 mm long; appendages of innermost bracts almost round, membra- nous, brownish, fimbriate-ciliate. Florets various shades of pink; pe- ripheral ones slightly large. Achenes 3.5—4.5 mm long, sparsely short- hairy; pappus 3.5—4.5 mm long, inner pappus bristles 1/7—1/5 as long as largest outer pappus bristles. Flowering June to September. Steppes, scrubs, meadows, old fields, up to mid-montane zone.— European Part: Upper Dniester (southern part), Volga-Kama (south- eastern part), Upper Volga (southern part), Upper Dniester, Middle Dnieper, Volga-Don, Trans-Volga, Bessarabia, Black Sea Region, Lower Don, Lower Volga (northern part), Crimea; Caucasus: Ciscaucasia, Dagestan; Western Siberia: Upper Tobol, Irtysh, Altai. 507 505 General distribution: Centrral Europe, Mediterranean, Balkans. Described from Crimea (between Simferopol and Belogorsk). Type in Leningrad. Note. Apparently, in the synonymy of this species it is also nec- essary to include C. fritschii Hayek, described from southeastern Eu- rope (in Denkschr. Math.-naturw. Klasse Akad. Wien, LXXII [1901] 623; Taf. II, fig. 9; Centaurea-Art. Osterr.-Ung. 39), and C. spinulosa Rochel (f7. Banat. rar. [1828] 76, Tab. 36), which differ from C. apiculata only by having weaker (on the average) leaf pubescence of leaves. Plants with longer spines at the tips of the appendages of the involucral bracts, often identified as C. spinulosa and in reality very similar to typical specimens of this species, are scattered almost through- out the range of C. apiculata, but are especially widespread in the Podolsk Upland, Donetsk Ridge, and Ciscaucasia. They hardly merit recognition as a separate ecogeographical race. C. apiculata is closely linked through intermediate populations with C. scabiosa, C. adpressa, C. ossethica, and C. pseudoscabiosa, as a result of which the morpho- logical boundaries between all these species are not all that distinct. Possibly, it would be better to treat them as subspecies of a single polymorphic species. C. apiculata often forms hybrids with C. orientalis L. (cf. note to that species) and with C. pseudomaculosa Dobrocz. 114. C. adpressa Ldb. Ind. Sem. hort. Dorpat. (1824) 3; Grossh. Opred. Rast. Kavk. 500; Kryl. Fl. Zap. Sib. XI, 2962.—C. scabiosa auct. non L.: MB. Fl. taur.-cauc. II, 350 p. p.—C. scabiosa var. angustata Ldb. FI. alt. IV (1833) 48.—C. scabiosa var. adpressa (Ldb.) DC. Prodr. VI (1837) 580; Ldb. Fl. Ross. II, 2, 701; Schmalh. FI. II, 126; Grossh. Fl. Kavk. IV, 218.—C. rigida Andrz. in Rogowicz. Obozr. (1869) 303, nom nud. non Willd. nec Russ.—Exs.: Hayek, Cent. exs. crit. fasc. II, No. 63. Perennial. Plants to 80-100 cm high, with short papilliform hairs mixed with crisped hairs and thin arachnoid-hairy-tomentum, weakly scabrous, green. Stems solitary or few, upright, branched mostly in upper part. Leaves pinnately parted or lobed, sometimes some of them entire, lower leaves quite long-petiolate, upper ones sessile or subsessile; lateral segments oblong to linear, usually entire, terminal segment larger than laterals. Involucre 9— 15 mm in dia and 15-22 mm long, thinly arachnoid-hairy-tomentose or glabrous; outer and middle involucral bracts with small (to 1 mm long) dark brown or brownish apical appendage and few, very short (to 0.5, less often to 1 mm long), brownish cilia along margin, sometimes almost lack- ing appendages and cilia, terminating in very short (to 1 mm) spinule; innermost involucral bracts with almost round, fimbriate appendage. Flo- rets various shades of pink, peripheral ones almost not enlarged. Achenes 508 506 4—6 mm long, sparsely short-hairy; pappus almost as long. Flowering June to September. Steppes, stony slopes, sands, old fields, meadows; up to mid- montane zone.—European Part: Volga-Don (southern part), Trans- Volga (occasionally), Bessarabia, Black Sea Region, Lower Don, Lower Volga, Crimea; Caucasus: Ciscaucasia, Dagestan, Eastern Transcaucasia (northwestern part); Western Siberia: Upper Tobol (southern part), Irtysh (southern and eastern parts); Soviet Central Asia: Aralo-Caspian (northern part), Lake Balkhash Region, Dzhungaria-Tarbagatai, Tien Shan, Syr-Darya. General distribution: Balkans, Dzhungaria-Kashgaria. Described from Odessa Region of Ukrainian SSR. Type in Leningrad. Note. This species replaces the previous one in the zone of arid steppes and semideserts; however, in the mountains of Soviet Central Asia, it is found in comparatively mesophytic communities. Hybrids of this species with C. orientalis L., C. reflexa Lam., and C. salonitana Vis. are known. 115. C. pseudoscabiosa Boiss. and Buhse. in Nouv. Mém. Soc. Nat. Mosc. XII (1860) 131; Grossh. Opred. Rast. Kavk. 500.—C. scabiosa var. spinulosa (Rochel) Boiss. Fl. or. III (1875) 656 p. p.— C. spinulosa auct. non Rochel: Grossh. Fl. Kavk. IV (1934) 218 p. p.—Exs.: GRF No. 4400. Perennial. Plants to 80-100 cm high, with short papilliform hairs mixed with crisped hairs and thin arachnoid-hairy-tomentum, weakly scabrous, green. Stems solitary or few, upright, branched mostly in upper part. Leaves pinnately parted or lobed, sometimes some of them entire, lower leaves quite, long-petiolate, upper ones sessile; lateral segments oblong to almost linear, entire, less often shart-lobed; termi- nal segment usually not larger than lateral ones. Involucre 12-18 mm in dia and 18-25 mm long, finely arachnoid-hairy-tomentose or gla- brous; outer and middle involucral bracts usually pectinate-ciliate al- most to middle (cilia 1.2—-2.5 mm long) but with very narrow (to 0.5 mm wide), decurrent part of appendage (or without it), with small obtusely triangular blackish-brown apical appendage (excluding de- current part) to 1.5—2 mm long, terminating in recurved spinule or spine (1.5)2—6(8) mm long; appendages of innermost, oblong involu- cral bracts almost round, pectinate-fimbriate. Florets various shades of pink, peripheral ones almost not enlarged. Achenes 5.0—6.5 mm long, sparsely short-hairy; pappus 6-8 mm long, scale-like inner pappus bristles 1/4—1/3 as long as longest outer pappus bristles. Flowering June to August. Stony slopes, steppes, scrubs; up to mid-montane zone.—Caucasus: Eastern Transcaucasia (southwestern part), Southern Transcaucasia 507 (eastern part). General distribution: Asia Minor, Armenia and Kurdistan, Iran-Afghanistan (north-western part). Described from Iran (in the region of Tabriz). Type in Geneva; isotype in Leningrad. Note. This species is very close to C. apiculata Ldb.; however, the larger achenes and the well separated range permit us to consider it as a separate, more southern, ecogeographical race. Populations intermediate between these species are known from Dagestan and many areas of Azerbaidzhan. It is also linked through intermediate forms with C. glehnii Trautv., which, being a more alpine and, correspond- ingly, more mesophilic race, often replaces C. pseudoscabiosa in the mid-montane zone. 116. C. glehnii Trautv. in Tr. Peterb. Bot. Sada, IV, 1 (1876) 382 p. p.; Boiss. Fl. or. Suppl. (1888) 314 p. p.; Grossh. Fl. Kavk. IV, 218 p. p. and Opred. Rast. Kavk. 500. Perennial. Plants 50, less often to 70 cm high, with short papilli- form hairs mixed with long crisped hairs and fine arachnoid-hairy- tomentum, weakly scabrous, green. Stems solitary or few, upright, branched in upper part. Leaves pinnately parted, lobed, or some of them undivided; lower leaves quite long-petiolate, upper ones sessile or subsessile; lateral segments broadly ovate to almost linear, usually entire; terminal segment often larger than lateral ones. Involucre 12— 23 mm in dia and 14—25 mm long, glabrous or finely arachnoid-hairy- tomentose; appendages of outer and inner involucral bracts thin-coria- ceous, obtusely triangular, 2—3.5 mm long (excluding border decurrent on bracts), blackish-brown, with whitish or brownish 1.5—3.5 mm long cilia along margin, terminating in recurved spinule or 2—6 mm-long spine; appendages of innermost bracts almost round, fimbriate. Florets pink, peripheral ones almost not enlarged. Achenes 5—6 mm long, sparsely short-hairy (hairs longer only near hilum); pappus 7-8 mm long, usually with brownish-gray tinge, scale-like inner pappus bristles 1/6—1/4 as long as longest outer pappus bristle. Flowering June to August. Stony slopes, meadows; mostly in mid-montane zone.—Caucasus: Southern Transcaucasia. General distribution: Armenia and Kurdistan. Described from Armenian ASSR (between Leninakan and village of Mastara). Type in Leningrad. Note. Apparently split up into several morphologically very weakly separated ecogeographical races more or less approximating the closely related species C. ossethica, C. pseudoscabiosa, and C. apiculata. The most typical of them is the very low-growing (usually to 30 cm high) race distributed in the region of Lake Sevan and Aragats Mountain. 510 508 Populations approaching this species have been placed tentatively in C. apiculata, also known from montane Dagestan. Series 2. Carduiformes Tzvel.—Biennials or short-lived monocarpic perennials with strongly thickened string-like root. Appendages of involucral bracts with more than 5 mm-long apical spine. 117. C. carduiformis DC. Prodr. VI (1837) 590; Boiss. FI. or. III, 659; Grossh. Fl. Kavk. IV, 219 and Opred. Rast. Kavk. 500.—C. glehnii var. bicolor Trautv. in Tr. Peterb. Bot. Sada, IV, 1 (1876) 382. Biennial. Plants to 70—80 cm high, with short papilliform and longer crisped hairs, mixed with finely arachnoid-hairy tomentum, more or less scabrous, green or somewhat grayish. Stems solitary or few, up- right, usually with few lateral branches. Leaves pinnately parted, lobed or some of them undivided, lower leaves rather long-petiolate, upper ones sessile or subsessile; lateral segments oblong to almost linear, entire or more or less irregularly toothed; terminal segment usually not larger than lateral ones. Involucre 15—27 mm in dia and 20-27 mm long, terminating in brownish or dark-brown, thin-coriaceous triangu- lar appendages and then in attenuate recurved spine, 6—16 mm long in middle involucral bracts, shorter in outer ones, spines up to 1.5 mm thick at base, flat or convex above, with few stiff cilia along margin near base; appendages of innermost bracts round or kidney-shaped, long-fimbriate. Florets usually pale pink, peripheral ones not enlarged. Achenes 4—5 mm long, more or less hairy; pappus 6—8 mm long, often with brownish-gray tinge, scale-like inner pappus bristles 1/6—1/4 as long as largest outer pappus bristles. Flowering May to August. Stony slopes, steppes, up to mid-montane zone—Caucasus: West- ern Transcaucasia (southern part), Southern Transcaucasia (western part). General distribution: Asia Minor (northeastern part), Armenia and Kurdistan, Iran (northwestern part). Described from northwestern part of Turkey. Type in Paris. Note. Like the previous species, it splits up into several, still in- adequately studied, narrowly endemic ecogeographical races, differing in the length of the spine and cilia on the involucral bracts, size of the capitula, color of the florets, and shape and pubescence of the leaves. Possibly some of these races are the ones described by C. Koch from northeastern Turkey as species: C. recurvata C. Koch (in Linnaea, XXIV (1851) 428), C. bicolor C. Koch (op. cit. 428), and C. albiflora C. Koch (op. cit. 430). However, in the absence of the type material of these species and also of C. carduiformis, it is difficult to say anything definite in this regard. On the border with Turkey, one can find still another species close to C. carduiformis, C. cyrtolepis Ldb. (FI. Ross. II, 2 (1845) 709), differing from it by having larger (20—40 51 —" 509 mm wide) capitula, spines strongly broadened at the base, and stems woolly tomentose at the base. 118. C. sintenisiana Gand. in Bull. Soc. Bot. France LXV (118) 37.—C. carduiformis DC. var. sintenisiana (Gand.) Ijin ex Tzvel. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX (1959) 437, pro syn.—Ic.: Czer. in Fl. Turkm. VII (1960) Plate XLIII. Biennial. Plants to 70-80 cm high, with short papilliform and longer crisped hairs mixed with fine arachnoid-hairy tomentum, more or less scabrous, green. Stems solitary or few, upright, usually with few lat- eral branches. Leaves as in previous species, but their terminal seg- ment usually larger than lateral ones. Involucre 15—28 mm in dia and 20-28 mm long (excluding spines), glabrous or subglabrous; outer and middle involucral bracts in upper part pectinate-ciliate, with whitish cilia 2-5 mm long, terminating in brownish thin-coriaceous append- ages and then in recurved spine, spine 15—40 mm long in middle bracts, shorter in outer, 1.5—3 mm thick at base, above more or less concave, grooved, with sparse stiff cilia along margin; appendages of innermost bracts almost round, long-fimbriate. Florets pink, peripheral ones not enlarged. Achenes as in previous species. Flowering May to July. Stony slopes, rocks; up to mid-montane zone.—Soviet Central Asia: mountainous Turkmenia. General distribution: Iran-Afghanistan (north- ern part). Described from Kopetdag (in Kyzyl-Arvat region). Type in Paris; isotype in Leningrad. Note. In Iran, it is linked with C. carduiformis through a series of intermediate populations (or narrowly endemic ecogeographical races?). Section 4. Stereophyllae (Tzvel.) Tzvel. comb. nova.—Centaurea sect. Acrocentron subsect. Stereophyllae Tzvel. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX (1959) 437.—Florets pale pink, less often pale yellow; involucral bracts with broadly lanceolate, weakly decurrent, ciliolate, brownish, apical appendage, usually terminating in short spine; pappus double, 1/3-1/2 as long as achenes. Type of section: C. stereophylla Bess. 119. C. stereophylla Bess. Enum. pl. Volhyn. (1822) 35; DC. Prodr. VI, 580; Boiss. Fl. or. III, 656; Schmalh. Fl. II, 127; Grossh. Opred. Rast. Kavk. 500; Sosn. in Fl. Gruzii, VIII, 567.—C. scabiosa var. stereophylla (Bess.) Ldb. Fl. Ross. II, 2 (1845) 701.—Ic.: Fedtsch. and Fler. Fl. Evrop. Ross. (1910) Fig. 1047; Prod. Fl. Roman. I (1923) 1095. Perennial. Plants 80-100 cm high, with short and rather stiff, papilliform hairs mixed with thin arachnoid-hairy-tomentum, scabrous, Sila 510 dark green. Stems solitary or few, upright, branched mostly in upper part. Leaves pinnately lobed or parted, sometimes some of them undi- vided and entire; lower leaves rather long-petiolate, upper ones sessile; lateral segments or lobes oblong, entire, obtuse, but usually with short cusp; terminal segment usually larger than lateral ones. Involucre 9— 14 mm in dia and 15—22 mm long, thinly arachnoid-hairy-tomentose or subglabrous; appendages of outer and middle involucral bracts thin- coriaceous, brownish or dark brown, broadly lanceolate, 1.5—4.0 mm long, short decurrent on bracts, with lighter colored, to 1.0—1.5-mm- long cilia along margin, usually with up to 2 mm-long spinule at apex; appendages of innermost bracts almost round, membranous, short-pec- tinate-fimbriate. Florets pale pink, less often whitish or light yellow; peripheral ones not enlarged. Achenes 3.5—5.0 mm long, sparsely short- hairy; pappus 1.5—2.5 mm long, usually with brownish-gray tinge; scale- like inner pappus bristles 1/5—1/3 as long as largest outer pappus bristles, but almost indistinguishable in thickness. Flowering June to August. Steppes, old fields, up to lower montane zone.—European Part: Bessarabia, Black Sea Region, Crimea (doubtful record); Caucasus: Ciscaucasia (doubtful record). General distribution: Centrral Europe (southeastern part), Balkans (northern part). Described from Podolsk Uplands of Ukrainian SSR. Type in Leningrad. Note. This species, together with the similar Balkan species C. rumelica Boiss., occupies a very isolated position in the subgenus Lopholoma, and with respect to a series of distinguishing characters, approaches other subgenera. Quite possibly these species have origi- nated as a result of hybridization of C. orientalis with some species of the subgenus Acrolophus, which would seem to be confirmed by the presence of both pale pink and light yellow florets in plants within the limits of a single species. This is not found in other groups of the genus Centaurea L. The reports of C. stereophylla from Crimea and Ciscaucasia apparently are erroneous and relate to the widely distrib- uted hybrid C. orientalis L. x C. apiculata Ldb. Subgenus 15. Acrolophus (Cass.) Dobrocz. in Bot. Zhurn. Akad. Nauk URSR, VI, 2 (1949) 64, 69.—Acrolophus Cass. in Dict. Sc. nat. L. (1827) 253.—Capitula small- or medium-sized; appendages of in- volucral bracts decurrent on bract margin, scarious or semicoriaceous, at apex with soft or prickly cusp, pectinate-fimbriate, less often with small teeth; corolla purple or pink; achenes oblong, pappus double as long as or often shorter than achene, sometimes absent. Biennials, less often, perennials with arachnoid-hairs; stems strongly branched; basal and lower cauline leaves pinnately incised. Sil Ww Sid Type of subgenus: C. maculosa Lam. A well-separated group, which Soviet authors have recognized as a separate genus (S.A. Nevski) or subgenus (D.N. Dobroczaeva). It apparently is closest to the subgenus Phalolepis (Cass.) Dobrocz. (hy- brids between them are common). Subgenus Acrolophus (Cass.) Dobrocz. includes quite numerous montane-xerophytic, petrophytic, psammophytic, and steppe or forest- steppe species, widely distributed in the northern Mediterranean and in the forest-steppe, steppe, and semidesert zones of Eurasia. As also in the subgenus Phalolepis (Cass.) Dobrocz., there are no forest spe- cies among them. Both subgenera are clearly differentiated into geo- graphical races, which aggregate into series. The racial composition of Acrolophus (Cass.) Dobrocz. has been studied especially in the Ukrai- nian SSR (works of E.M. Lavrenko, Yu. D. Kleopov, D.N. Dobroczaeva, and the author), and in the Caucasus (investigated by T.A. Dumbadze). However, it is far from completely known for the whole territory of the Soviet Union. Considerable difficulty arises in the intensive inves- tigation of the systematics of the subgenus because of the profusion of interseries and partly intersectional hybrids, that can be distinguished readily in nature but not always in herbaria. We could not form a clear idea of C. pseudovina Illar. (in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVIII [1957] 308), described by N.B. Illarionova from this group of species. In her opinion, this species is closer to C. lJavrenkoana Klok., differing by having a shorter (1-1.5 mm) pappus, shorter fimbriae, and a less-developed cusp of the appendages of the involucral bracts. The reported differences do not transcend the limits of variation of the characters in C. lavrenkoana Klok. Authentic specimens of C. pseudovina Illar., except for a single shoot, could not be found in the Herbarium of the V.L. Komarov Botanical Institute in Leningrad. The species described from the Tarkhankut Peninsula in Crimea also requires collection of new mate- rial for definitive clarification. Without herbarium material, we could not include this species in the present treatment, more so because its available description is almost entirely contained in our description of the southern Ukrainian C. Javrenkoana Klok. 1. Involucre ovate or oblong-ovate, rarely almost cylindrical, often dichromatic, with more or less colored tips of bracts and append- ages; involucral bracts with 3—7 veins; appendages of involucral bracts, except central thick part, scarious with soft apical cusp and finibriaerate. week crete a, ey domed saath Niles, Piel meinem I ss 2s + Involucre cylindrical or ovoid-cylindrical, usually monochromatic, yellowish; involucral bracts with 1-3 veins; appendages of involu- 514 512 cral bracts coriaceous, with prickly cusp and stiff fimbriae ........ 18. Involucral bracts with 5—7 veins; appendages of involucral bracts pectinate-fimbriate, lacking distinct scarious auricles at base, black or brawi.eiSteppe plants 67% viet eals Ey ae eee se Meee 3. Involucral bracts with 3—5, rarely 7 veins; appendages of involucral bracts fimbriate, with more or less distinct scarious auricles at base, less often irregularly toothed, only in central thick part weakly colored. Plants on sands or stony OUtCTOPS .............:::cceeeeeeseeeeeeee 6. General inflorescence corymbose; appendages of outer and middle involucral bracts including fimbriae black. Western species not reach- Ine OMEpeN ey eR. ee ee 120. C. rhenana Boreau. General inflorescence paniculate; appendages of involucral bracts brownish, less often almost black, but with ochreous fimbriae (only inithe ‘Caucasian! C./kubanica Kiloks)va.3 a eee ee ee 4. Involucre oblong-ovate, 10-11 mm long and about 7 mm wide; appendages of outer and middle involucral bracts with 4—6 pairs of fimbriae, light brown. Terminal lobe of leaves narrowly linear, 1.0— Ssmmiwiderducwe) Audoosss. ah Ms. 123. C. micranthos Gmel. Involucre ovate or broadly ovate, much larger; appendages of outer and middle involucral bracts with 6—10 pairs of fimbriae dark brown to almost black. Terminal lobe of leaves linear or oblong, to 4—5 mat Wade) NT FEL OOP ee eee, Cae ee nee ee 3) Basal leaves bipinnate. General inflorescence broadly paniculate; appendage of outer and middle involucral bracts dark brown; pe- ripheral florets to 15 mm long; mature achenes brown,with pappus 1—1.5 mm long. Plants of plains and steppes ...............sceeeceeeeeeeeeeeee saree Se SH. PUREE aes OMEN hee. 121. C. pseudomaculosa Dobrocz. Basal leaves pinnate, with lobed segments. General inflorescence oblong-paniculate; appendages of outer and middle involucral bracts almost black, with ochreous fimbriae; peripheral florets 20—21 mm long; mature achenes blackish with pappus 2.3 mm long. Plants of montane’ steppes lacey. ieee... ena 122. C. kubanica Klok. Involucre oblong-ovate to ovate-conical; appendages of involucral bracts with soft fimbriae, usually with well-developed scarious au- ricles below or margin entirely irregularly toothed; pappus always well-developed, as long as achene or slightly shorter, rarely much shorter (in C. odessana Prod.). Plants exclusively on sands (obli- gateipsainimopliytes ies). Week ea Nee Sere: odie OES hae ie Involucre oblong-ovate to almost cylindrical; appendages of in- volucral bracts with few stiff fimbriae, with inconspicuous scarious auricles below; pappus always shorter than achene, often greatly underdeveloped. Lithophilous plants ..............cecceeeseeeeseeneeeees 12: Involucre 12-15 mm long; appendages of middle involucral bracts 515 123 13: 513 with 6-8 pairs of fimbriae, lacking distinct scarious auricles at base. Very rare plants; known only from sands of Savran River in OdessapResion tlw... Ok AB ee 129. C. savranica Klok. Involucre 9-12 mm long; appendages of middle involucral bracts with fewer fimbriae and with well developed scarious auricles at base or lacking distinct border, entirely irregularly toothed ......... 8. Appendages of outer and middle involucral bracts fimbriate, cen- tral thick part of appendage light reddish-brown or yellowish-brown with small dark purple spots at base ...........ecceeseeeseeeeesneeeseeeeeeneee 2. Appendages of outer and middle involucral bracts irregularly toothed, often divided; central thick part of appendage yellowish, lacking GSTAATea UNG Sf Sp. ak ese eds A ae Cede ape eaiha utes cat, aaa 10. Stems, especially in lower part, and leaves white-arachnoid-hairy- tomentose; pappus noticeably to much shorter than achene. Plants at coastal sands swe) 2)-.8..) eee: Bae 127. C. odessana Prod. Stems and leaves not densely woolly-arachnoid-hairy; pappus as long as achene or very slightly shorter. Plants of river sands .............. Weta peak Sh Beste oh ey al Sh aba 128. C. borysthenica Gruner. Involucre ovate-conical, 9-11 mm long, 6-8 mm in dia; stems sharply scabrous above, along ribs ............. 124. C. majorovii Dumb. Involucre oblong-ovate, narrower; stems entirely glabrous or less often weakly scabrous above, along ribs ..............ccccscceseseeeeneeeees I, Involucre 9-12 mm long, 5—6 mm in dia; stems rather densely arachnoid-hairy, somewhat thick, with short floriferous branches; leaves along margin scabrous from conical tubercles ................... pein, Lh A celled. Ata webs yeni coro samen A 125. C. sophiae Klok. Involucre 9-11 mm long, 3.5—5 mm in dia. Stems not densely arachnoid-hairy, glabrous above, somewhat thin, with long florifer- ous branches; leaves smooth on margin ....... 126. C. arenaria MB. Middle involucral bracts with 3 veins; pappus 1.5—2.5 mm long, much shorter than achene but well-developed. Plants of steppe re- Bronsvo tsp lang hia wel. Le.OA wide, Ve RE oe 13. Middle involucral bracts with 3—7 veins; pappus not longer than 1.5 mm, more or less reduced. Plants of Crimea-Caucasus ........ 14. Involucre 10—12 mm long, 5—6 mm in dia, ovate-cylindrical. Stems and leaves weakly arachnoid-hairy .......... 130. C. besseriana DC. Involucre 8—11 mm long, 3—5 mm in dia, almost cylindrical with conical base. Stems and leaves rather densely arachnoid-hairy ....... Uae te, Rhea eam selina lapenren colette ow entra bd 131. C. lavrenkoana Klok. ermal ivonh lier eaten, Avert ienndind sehied oy BS oes eh ak ieee ieee AR te IS: Pappus to 1.0—1.5 mm long. Plants of Caucasus ................ ee 16. Capitula all solitary at tips of branches 3—12 cm long; involucre 514 16. 20. Pile 10-12 mm long, 5—6 mm in dia ............. 132. C. steveniana Klok. Capitula partly sessile or subsessile, aggregated, partly solitary at tips of branches up to 4.5 cm long; involucre about 9 mm long, 345=4 Ooi dia i Sage. Aen 133. C. koktebelica Klok. Basal and lower cauline leaves bipinnate, with incised second-order segments. Involucre 11-13 mm long, 6-8 mm in dia; middle in- volucral bracts with 5—7 veins. Achenes 3.3—4.0 mm long ................. dabcansit hess Nd ARNE Rte RESET GI 136. C. gulissashvilii Dumb. Basal and lower cauline leaves with undivided second-order seg- ments. Involucre 10-11 mm long, 5—6 mm in dia; middle involu- cral bracts with 3-5 veins. Achenes not more than 3.2 mm long ... ERIN EEN et OES, BEDE, OR a, EER SO ane eam rs oe Stems with dense prickly bristles on ribs. Outer and middle involu- cral bracts with 5 veins; their appendages with cusp 0.9-1.5 mm long and 5-6 pairs of fimbriae to 1.5—2 mm long. Pappus about 1.5 aT OMG cA Be ee 135. C. ovina Pall. Stems scabrous on ribs due to scattered acute tubercles. Outer and middle involucral bracts with 5 inconspicuous veins; their append- ages with cusp about 0.5 mm long and 4—S pairs of fimbriae about Punmmilong, 084.2 Sea OPA: 134. C. vicina Lipsky Leaves with narrowly linear or linear, less often oblong-lanceolate or obovate, terminal lobe 1-4 mm wide. Capitula solitary at tips of short branches or partly aggregated; apical spine of appendages more or less elongate; pappus much shorter than achene or often not developed! Weedse. LRU ne TEER: BS ee ee 19. Leaves, at least cauline, with large terminal lobe or lyrate. All capitula aggregated in dense clusters of 5-10 at tips of main branches; apical spine of appendages very short; pappus not shorter thanvacheneMPlants of mGumtainsi es. Lee ee Ree, 22. Capitula often solitary, partly aggregated in small clusters; apical part of middle involucral bracts with appendage arcuately recurved outward and downward... 140. C. squarrosa Willd. Capitula solitary, less often in lax clusters; upper part of middle involicral bracts not recurved Yee CE kee 20. Involucre 10-15 mm long, 4-6 mm in dia; appendages of outer and middle involucral bracts with cusp 0.5—2.0 mm long and 3-9 pairs of fimbriae 0.4-1.7 mm long. Achenes about 3.5 mm long with pappus 2.5—3.0 mm long ........ eee eeeeeees 139. C. caspia Grossh. Involucre shorter and narrower; appendages of outer and middle involucral bracts with cusp to 4 mm long and 3-7 pairs of fimbriae, half as long as cusp. Achenes up to 3 mm long, with pappus incon- spictious or lacking) Tee RS oe ea eee onan oN" Stems and leaves scatteredly arachnoid-hairy. Capitula solitary at 517 515 tips of very short or somewhat longer branches. Involucre 8-10 mm long, 2.5—5.0 mm in dia; prickly cusp of appendages weakly recurved outward. Florets pale pink or whitish, rarely light purple CEN UD ALANS ACPI COPE EO EPEAT Ak i 137. C. diffusa Lam. + Stems and leaves rather densely arachnoid-hairy. Capitula usually more or less clustered at tips of second-order branches. Involucre 10-12 mm long, 3-3.5 mm in dia; prickly cusp of appendages strongly divaricate and in lower part of involucre more or less recurved! Blorets. purpley...Aeue2t. 2ane2 5. 138. C. aemulans Klok. 22. Stems entirely smooth but leaves white-arachnoid-tomentose. Ap- pendages of outer and middle involucral bracts with 24 pairs of HMITIAS ME WO LAS Bes, 141. C. albida C. Koch + Stems sharply scabrous above along ribs, leaves not densely arach- noid-floccose. Appendages of outer and middle involucral bracts with 4—7 pairs of fimbriae ....... 142. C. aggregata Fisch. and Mey. Section 1. Maculosae (Hayek) Dumb. in Dokl. Akad. Nauk Arm. SSR, V,2(1946) 48.—Sect. Acrolophus c) Maculosae Hayek, Centaur.- Art. Osterr.-Ung. (1901) 4, 81.—Plants of steppes and forest-steppes, weakly arachnoid-hairy. Involucral bracts with 5—7 veins; appendages of involucral bracts pectinately fimbriate, lacking scarious auricles, black or brown (thus, involucre bicolorous); pappus shorter than achenes. 120. C. rhenana Boreau, Flore du Centre de la France, ed. 3 (1857) 355; Hayek, Centaurea-Art. Osterr.-Ung. (1901) 85; Dobrocz. in Vizn. Rosl. URSR, 581.—C. maculosa ssp. rhenana Gugler in Ann. hist.-nat. Mus. Nat. Hung. VI (1908) 164, 265; Hegi, Ill. fl. VI, 2, 970.—Ic.: Hayek, loc. cit. Taf. VIII, fig. 4; Hegi, loc. cit., fig. 600 (a- b).—Exs.: Fl. exs. reip. Bohen.-Sloven. Nos. 638, 888. Biennial. Stem upright, 30-80 cm high, ribbed, scabrous, espe- cially above along ribs, including leaves weakly arachnoid-hairy, branched from middle; branches leafy to base of capitula, long, equal- ing stem. Leaves pinnately and bipinnately incised to linear or nar- rowly linear acuminate lobes 1.5—3 mm wide, scabrous, punctate-glan- dular; lower leaves petiolate, middle and upper ones sessile, apical leaves often undivided, linear, 1-1.5 cm long. General inflorescence corymbose; capitula solitary at tips of branches. Involucre ovate, about 14 mm long, 9-10 mm in dia; involucral bracts green, ochraceous in upper part, with 5 conspicuous veins; appendages of outer and middle involucral bracts black, triangular, bordering tip of bract, regularly fimbriate, with 6—8 fimbriae on each side, 1.0—1.5 mm long, black or dark brown, longer than tail-like tip of appendages; inner bracts 518 516 oblong-lanceolate with ovate or triangularly lanceolate, toothed, brown appendage surpassing middle bracts. Florets pink; peripheral florets sterile, to 15 mm long. Mature achenes black with white ribs, 2.5—3.0 mm long; pappus 1.5—2.0 mm long, white. Flowering July to September. Steppe slopes and stony outcrops, mostly limestones.—European Part: Upper Dnieper, Upper Dniester (south), Middle Dnieper (west, up to southern Bug in southeast and Pripyat River in northeast). Gen- eral distribution: Centrral Europe (up to Rhine in west and Danube in south). Described from France (Rhine Valley). Type in Paris(?). Note. To the east of the Rhine, this species replaces the westernmost race of C. maculosa Lam. s. str. According to the studies of Hayek (1901), Briquet (1902), Prodan (1930), D.N. Dobroczaeva (1949, 1950), and others, C. rhenana Boreau differs clearly from C. maculosa Lam. by having sparser arachnoid hairs on the stem and leaves, a larger involucre, a darker color of the appendages and fimbriae on the in- volucral bracts, and a much better developed pappus. In C. maculosa Lam., the fimbriae on the appendages of the involucral bracts are lighter in color and more numerous (to 10 on each side), and much lighter than the undivided blackish part of the appendage. 121. C. pseudomaculosa Dobrocz. in Bot. Zhurn. Akad. Nauk URSR, VI, 2 (1949) 73; in Vizn. Rosl. URSR, 582.—C. maculosa auct. fl. Ross. p. p. non Lam.—C. maculosa ssp. rhenana Ijin in FI. Yugo-Vost. VI (1936) 432. Biennial. Root string-like, strong. Stem, upright, 30—70 cm high, ribbed-sulcate, with leaves sharply scabrous and not densely arach- noid-hairy; lower branches not reaching stem tip, upper ones some- times even surpassing it. Basal and lowermost cauline leaves petiolate, bipinnately incised with 1-4 mm-wide, linear, acuminate, terminal pin- nules; others sessile, middle ones pinnately incised; apical leaves often undivided, oblong-lanceolate, 1.5—2.0 cm long; all leaves with profuse punctate glands on surface. General inflorescence broadly paniculate; capitula solitary at tips of densely leafy, relatively long or very short branches. Involucres ovate, 12-14 mm long, 10-13 mm in dia; involu- cral bracts green, with 5 prominent veins on outer side; outer bracts ovate,‘ middle ones oblong-ovate, inner bracts oblong-lanceolate; ap- pendages of outer and middle bracts dark brown, 2.5—3.0 mm long (including cusp), regularly fimbriate, short-decurrent on margin in upper part of bract, lacking distinct scarious auricles at base; fimbriae 6—9 on each side of appendage, somewhat lighter colored than its central part, 2.5—3.0 mm long; appendages of inner bracts ovate or oval, short- fimbriate or irregularly toothed. Florets pink, to 15 mm long. Achenes about 3 mm long, brown, with light colored ribs; pappus 1.0—1.5 mm long. Flowering July to August. 519 517 Forb-feather grass steppes, steppe slopes, stony, mostly limestone outcrops, edges and glades of steppe forests, sometimes as weed in fields —European Part: Middle Dnieper (east), Volga-Don, Trans- Volga, Volga-Kama (south), Black Sea Region (north), Lower Don (north), Crimea (?); Western Siberia: Upper Tobol (western part). Endemic. Described from vicinity of Makaevka in Donbass. Type in Kiev. 122. C. kubanica Klok. in Addenda XXVII, 615.—C. maculosa auct. fl. cauc. p. p. non Lam.—C. rhenana Dumb. in Dokl. Akad. Nauk Arm. SSR, V, 2 (1946) 48, non Boreau. Biennial. Stems solitary, upright, 10-110 cm high, ribbed-sulcate, along ribs especially above scabrous from conical tubercles, with very weakly arachnoid-hairy leaves in lower part, branched from below middle or above, with upward-spreading branches. Basal leaves with- ering by flowering time, petiolate, pinnately incised with more or less incised segments; cauline leaves short-petiolate (lowermost) to sessile, 1—8 cm long, pinnate with 1-11 pairs of lateral, undivided or incised segments in lower leaves, beneath and along margin sharply scabrous from spiniform bristles; terminal segment of leaves linear or oblong, 2-27 mm long, 0.3—5.0 mm wide, acute and somewhat prickly; cauline leaves sessile with undivided or divided auricles, apical leaves undi- vided, oblongly linear, acute, bent. General inflorescence in well de- veloped plants oblong-paniculate; floriferous shoots 0.5—14.0 cm long, not densely leafy. Capitula solitary at tips of these branches. Involucre ovate or broadly ovate, 12-15 mm long, 7-12 mm in dia, distinctly dichromatic; involucral bracts ovate to linear, 3-11 mm long, 1.5—3.2 mm wide, with 5—7 veins, greenish, with fine, sessile, yellow glands; appendages broadly ovate to elliptical (in lower bracts), 1.5—2.5 mm long, 1-4 mm wide; with triangular or ovate-triangular, black (except often greenish part at base), central part in outer and middle bracts, cusp 0.5—1.5 mm long; fimbriae blackish in lower part, ochreous above, filiform at tip, flexuous; appendages of inner bracts elliptical or almost ovate-lanceolate, toothed to entire. Corolla purple-pink, 20-21 mm long in peripheral florets, 14-15 mm in central florets. Achenes ob- long, somewhat laterally bent, 2.7—3.2 mm long, 1.2—1.5 mm wide, short and very thin hairy; mature achenes blackish, with yellow longi- tudinal striations; pappus 1.0—2.3 mm long. Flowering July to August. Mountain and foothill steppes, scrub.—European Part: Lower Volga (near Stavropol), Ciscaucasia, Dagestan. Endemic. Described from Teberda River. Type in Leningrad. Note. This species undoubtedly differs from the Centrral European C. rhenana Boreau in many characters, which was first noted by D.N. 520 518 Dobroczaeva. Apparently, it hybridizes in the contact zone with C. micranthos Gmel., producing forms with mixed characters. One such form was described by T.A. Dumbadze under the name C. micrantha Gmel. var. recurvata Dumb. (Dokl. Akad. Nauk Arm. SSR, V, 2 (1946) 47). 123. C. micranthos Gmel. Reise. I (1770) 135; Hayek, Centau- rea-Art. Osterr.-Ung. (1901) 92; Grossh. Fl. Kavk. IV, 221 and Opred. Rast. Kavk. 501; Dobrocz. in Vizn. Rosl. URSR, 582.—C. maculosa ssp. micranthos (Gmel.) Gugler in Ann. hist.-nat. Mus. Nat. Hung. VI (1908) 164, 179, 265; Ijin in Fl. Yugo-Vost. VI, 432.—C. biebersteinii DC. Prodr. VI (1837) 583; Ldb. Fl. Ross. II, 2, 703.—C. maculosa auct. fl. Ross. p. p. non Lam.—Exs.: GRF No. 1115 (sub C. maculosa Lam.); Fl. exs. reip. Bohem.-Sloven. No. 890; Hayek, Cent. exs. crit. No. 69 (sub C. maculosa ssp. micranthos). Biennial (perennial). Stems upright, 20—75 cm high, ribbed-sul- cate, branched from base or middle, with leaves scabrous and dis- tinctly arachnoid-hairy, leafy to top. Leaves pinnately and bipinnately incised, with narrowly linear, acuminate terminal lobe, 1.0—1.5 mm wide, pale green from less dense arachnoid-hairy tomentum; lower leaves petiolate, others sessile; apical leaves sometimes undivided, oblong-lanceolate, about 2 cm long. Capitula solitary at tips of stem and numerous branches, forming paniculate inflorescence. Involucre oblong-ovate, 10-11 mm long, about 7 mm wide, somewhat arach- noid-hairy; involucral bracts pale green, outer bracts ovate, middle ones oblong-ovate, with 5 prominent veins on outer side, with brown, triangular, regularly fimbriate apical appendage; fimbriae 4—6 on each side, about 2 mm long; inner bracts oblong-lanceolate, with scarious border, with oval or ovate, short-fimbriate or irregularly toothed apical appendage 1.0-1.5 mm long. Florets pink, peripheral about 15 mm long. Achenes about 3 mm long, blackish, with whitish ribs; pappus about 1 mm long, white. Flowering July to August. Dry steppes, chalky and clayey outcrops.—European Part: Middle Dnieper (south), Bessarabia, Black Sea Region, Lower Don, Volga- Don, Trans-Volga, Crimea; Western Siberia: Upper Tobol (west); Caucasus: Ciscaucasia. General distribution: Centrral Europe (south- east), Balkans (north). Described from Don. Type missing. Note. D.N. Dobroczaeva described a form in this species, f. campestris Dobrocz., which differs from the type by having subsessile capitula aggregated in groups of 3-5 at the tips of the branches. The significance of this form is not clear. Series 2. Arenariae (Hayek) Dumb. in Dokl. Akad. Nauk Arm. SSR, V, 2 (1946) 47.—Sect. Acrolophus d) Arenariae Hayek, Centau- 52 — keg rea-Art. Osterr.-Ung. (1901) 4, 94 p. p.—Plants of river and marine sands. Involucral bracts with 3—5 veins; appendages of involucral bracts whitish-scarious except colored central part, with irregular fine teeth or fimbriae but with distinct fine-toothed auricles at base. Pappus al- most as long as achene or slightly shorter, less often much shorter. 124. C. majorovii Dumb. in Dokl. Akad. Nauk Arm. SSR, V, 2 (1946) 47.—C. arenaria auct. fl. Ross. p. max. p. non MB. Biennial. Stems 1—3, upright, 25-60 cm high, branched from base or middle, sharply scabrous above along ribs, with leaves rather densely arachnoid-hairy. Leaves pinnately or bipinnately incised with oblong- lanceolate or narrowly linear, entire, acuminate lobes, 1-4 mm wide, sharply scabrous, punctate-glandular; basal and lower cauline leaves more or less long-petiolate, middle and upper ones sessile, apical leaves undivided, narrowly lanceolate. Capitula solitary at tips of stems and its short branches. Involucre ovate-conical, 9-11 mm long, 6—8 mm in dia; outer bracts ovate, about 4 mm long, with appendage 1.0—1.5 mm long, 1.5—1.75 mm wide; middle bracts oblong-ovate, 7-8 mm long, 2.5—3 mm wide, with 5 veins on outer side, with broadly ovate apical appendage 2—3 mm long and 2—3.5 mm wide; appendages finely toothed with small broad notch and short cusp in it; central part of appendage narrowly triangular, yellowish, lacking conspicuous spot at base; inner bracts narrow, linear, 9-11 mm long and 1-2 mm wide, with 3 veins on outer side, and oblong-elliptical apical appendage about 1 mm long. Florets pink, peripheral 15—16 mm long, central ones 12—13 mm long. Achenes 3.0—3.5 mm long, blackish with whitish ribs; pappus 2-3 mm long. Flowering July to August. River, more rarely marine and dry valley sands.—European Part: Middle Dnieper (east), Volga-Don (south), Trans-Volga, Black Sea Region (northeast), Lower Don, Lower Volga; Caucasus: Dagestan, Eastern Transcaucasia. Endemic. Described from Dagestan sands near village of Kumtorkal. Type in Yerevan. Note. I compared plants from the sands of the Caspian Coast, particularly from Kumtorkal, with those from the Urals, Volga and Donetsk. However, I could not find appreciable difference among them. Moreover, this species is fairly variable, and I am not fully convinced of its racial uniformity. 125. C. sophiae Klok. in Addenda XXVII, 615. Biennial. Stems about 50-60 cm high, somewhat cylindrical be- low, blunt-ribbed above, smooth along ribs up to top, rather densely arachnoid-hairy, strongly branched, with upward-spreading branches. Basal leaves few at flowering time; lower cauline leaves pinnately 522 520 incised with linear or narrowly linear segments about 1—2 mm wide, along margin scabrous from conical tubercles; apical leaves linear or narrowly linear, 4-15 mm long, 0.5—1.0 mm wide. Capitula solitary at tips of 2.5—9.0 mm long branches. Involucre oblong-ovate, 9-12 mm long, 5—6 mm wide; involucral bracts ovate to linear, 2-8 mm long, 1.2-2.5 mm wide, with 5 veins on outer side, brownish-green like central part of appendage; appendages scarious except in their central part whitish, transversely oval in outer and middle, 1.5—2.0 mm long, 2.5—3.5 mm wide, toothed and finely incised in upper part; appendages oblong-ovate-lanceolate in inner bracts, about 1.5—2.0 mm long, al- most entire. Corolla pink, in peripheral florets about 15 mm and in central florets about 13 mm long. Achenes oblong, 3.2—3.6 mm long, 1.2-1.6 mm wide, mature ones olive-dark-brown; pappus 2—3 mm long. Flowering July to August. River sands.—European Part: Lower Don. Endemic. Described from riverbank sands of the Don in vicinity of village of Tsimlyanskaya. Type in Leningrad. Note. This plant was collected only once by S.G. Gorshkova in whose honor I name this species. In its characters it occupies an inter- mediate position between C. arenaria MB. and C. majorovii Dumb., clearly differing from both of them. 126. C. arenaria MB. in Willd. Sp. pl. III, 3 (1803) 2278 p. p. (non alior.); Fl. taur.-cauc. II, 347; III, 590; DC. Prodr. VI, 581; Ldb. Fl. Ross. II, 2, 702; Schmalh. Fl. II, 128 p. p—C. wolgensis DC. Prodr. VI (1837) 581; Ldb. Fl. Ross. II, 2, 702. Biennial. Stems usually few, upright or ascending, 25—80 cm high, relatively thin, angular, smooth on faces, less often noticeably sca- brous above on branches, less densely arachnoid-hairy below, glabrous above, branched from below middle, with upward-spreading branches. Basal and lower cauline leaves petiolate, bipinnately incised, weakly arachnoid-hairy, smooth; terminal lobe of leaves mostly linear to nar- rowly linear, sometimes oblong-elliptical, 1-15 mm long, 0.5—1.0 mm wide; apical leaves undivided, narrowly linear. General inflorescence rather lax paniculate-corymbose; capitula solitary at tips of relatively long (2—13 cm) and thin, weakly leafy floriferous branches. Involucre oblong-ovate, 9-11 mm long, 3.5—5.0 mm in dia; involucral bracts ovate to linear, 2-10 mm long, 0.8—2.0 mm wide with 5 veins; append- ages round-ovate to oblong-elliptical (in inner bracts), 1.2—2.0 mm long, 0.7—2.0 mm wide, scarious except central yellowish part, whitish with inconspicuous cusp 0.1—0.3 mm long, toothed or irregularly fim- briate, fimbriae up to 0.6 mm long, with entire, decurrent auricles below. Corolla pink, about 15 mm long in peripheral florets. Achenes 523 521 oblong-linear, 2.8—3.7 mm long, 1.2—1.4 mm wide, finely hairy; pap- pus 2—4.2 mm long, pappus hairs often yellowish. Flowering July to August. Sands, often coastal—European Part: Lower Volga; Caucasus: Dagestan, Eastern Transcaucasia (Apsheron Peninsula); Soviet Central Asia: Aralo-Caspian. Endemic. Described from lower reaches of Kuma River. Type in Leningrad. Note. Marschall-Bieberstein referred to this species also Ukrai- nian plants from Dnieper, reporting them in his third volume of the Flora as a separate, unnamed variety, later described by L. Gruner as a separate species, C. borysthenica Gruner. According to Bieberstein, it differs from typical C. arenaria MB. by having much larger, ovate involucres and finely divided leaves. The type of this species in Bieberstein’s herbarium has the label: “Ex deserto cumano a. 1803,” and agrees fully with the description of the typical form of the species. The characters of the species indicated by Bieberstein and the type specimen do not come at all close to those of the southeastern plant, for which the name C. arenaria MB. was established. Appar- ently, the latter must be named C. majorovii Dumb. at least if the Caspian race, to which this name directly belongs, and the Donetsk race do not differ. However, Ledebour and De Candolle distinguished our plant from C. arenaria MB and even referred it to a separate section, which was treated by Cassini as the separate genus Spilacron (S. crupinoides Cass. in Dict. Sc. nat. L. 238). The reason for such a separation lies in that in some of the specimens of this species examined by Cassini and Ledebour the achenes were entirely lacking the inner pappus, which consists of short, apically connate scales. In the other specimens col- lected from the same locality, which did not differ in any way from the former, the pappus was normal and double. Plants from the more southern localities (for example, the Apsheron plants) differ somewhat from the Volga-Dagestan plants by having branches that are scabrous on the sides and probably also other char- acters. 127. C. odessana Prod. in Contrib. bot. Cluj. I, 17 (1928) 11; Klok. in Nauk. Zap. KDU, VII, 6, 69; Dobrocz. in Vizn. Rosl. URSR, 582.—C. arenaria auct. fl. ucr. p. min. p. non MB. Biennial. Stems upright, 30—70(100) cm high, 3—5 (7) less often solitary, rather thick (2.0—5.5 mm thick) at base, branched from base or middle, with leaves arachnoid-white-tomentose especially below, like leaves beneath. Basal and lower cauline leaves more or less long- petiolate, bipinnately incised, with narrowly linear, acuminate, lobes 524 522 1.5-4.5 mm wide; apical leaves undivided, sessile like middle. Capitula solitary at tips of stem and its rather long branches, with 1-2 apical leaves. Involucre oblong-ovate, 10-12 mm long, 4.5—6.5 mm in dia, arachnoid-hairy; outer involucral bracts ovate, 3.0—4.5 mm long, about 2.5 mm wide, with appendages up to 2 mm long, having short cusp and 3—4 fimbriae on each side; middle bracts oblong-ovate, 5.5-8.0 mm long, 2.0-3.5 mm wide, with appendages 1.5—2.5 mm long and as wide, having short cusp and 4—5(6) regular fimbriae on each side; inner bracts linear, 9.0-10.5 mm long, 1.5—2.0 mm wide, with appendages about 1.5 mm long, undivided part of appendages grayish-reddish brown, fimbriae of lighter color than central part and longer than its width. Florets pink. Mature achenes 3—4 mm long, blackish; pappus 1.0—2.5(3.0) mm long, shorter than achenes. In other characters, it resembles C. borysthenica Gruner. Flowering July to September. Coastal sands.—European Part: Black Sea Region (only along coast of Black and Azov seas, from Vilkov at estuary of Danube to vicinity of Zhdanov). Endemic. Described from vicinity of Odessa. Type in Cluj. 128. C. borysthenica Gruner in Bull. Soc. Nat. Mosc. XLI (1866) 426; Kleopov in Vizn. Kiivsk Bot. Sadu, IX, 74, rest.; Klok. in Nauk. Zap. KDU, VII, 6, 62; Dobrocz. in Vizn. Rosl. URSR, 582. Biennial. Stems upright, 25-120 cm high, solitary or 2—5, branched from base or middle, cylindrical below, ribbed-sulcate above, scabrous along ribs, not entirely densely woolly-arachnoid. Basal and lower cauline leaves petiolate, pinnately or bipinnately incised with oblong or oblong-ovate terminal lobe, terminating short cusp; middle and upper leaves sessile, pinnately incised, with linear, acuminate lobes, some- times with 1-2 teeth at base; apical leaves undivided, linear or nar- rowly linear; all leaves scabrous from small, sharp bristles and with dense punctate glands. Capitula solitary at tips of stems and branches, subtended by 1-2 leaves shorter than involucre. Involucre oblong- ovate, 9-12 mm long, 5—6 mm in dia, weakly arachnoid-hairy; outer involucral bracts ovate, 3-6 mm long, 2.5—3.0 mm wide, with apical appendage bearing short cusp and 3 short fimbriae on each side; middle bracts oblong-ovate, 6-9 mm long, about 3 mm wide, with (3) 5 promi- nent purple veins on outer side; appendages of middle bracts about 2 mm long and 2.5-3.5 mm wide, with cusp up to 1 mm long and 4-5 parallel fimbriae on each side, rather broad at base, with irregularly small toothed, scarious border; central part of appendages of middle and outer bracts triangular or ovate-triangular, yellowish-brown at base with more or less distinctly dark purple spots; inner bracts linear or 525 5238 narrowly linear, 10-11 mm long, 1.0—2.5 mm wide, with 3 conspicu- ous veins on outer side, with scarious border and apical appendage, bifid, 1.5—2 mm long and about 1 mm wide, with short cusp in its deep notch, both appendage lobes toothed or with 2-3 short fimbriae. Flo- rets light purple; corolla of peripheral florets about 16 mm long, of central florets about 12 mm long. Achenes 2.5—3.5 mm long, blackish, with whitish ribs; pappus as long as achene or slightly shorter. Flow- ering June to September. River sands, rarely on granite outcrops (distinct form).—Euro- pean Part: Upper Dnieper (south), Middle Dnieper (west), Black Sea Region (west). This species is endemic to sands of Dnieper Basin and Southern Bug. Described from vicinity of Aleksandrovsk (now Zaporozhy). Type in Leningrad. Note. The unique form reported from the granite out crops along the Southern Bug deserves attention; it has distinct spots on the ap- pendages of the involucral bracts. 129. C. savranica Klok. in Nauk. Zap. KDU, VII, 6 (1948) 67 (ucrainice), 80 (latine); Dobrocz. in Vizn. Rosl. URSR, 582. Biennial. Stems upright or ascending at base, 30-60 cm high, soli- tary branched from below or above middle; branches few, ribbed- sulcate from base, distinctly tetraquetrous above, sharply scabrous along ribs, all whitish-arachnoid-hairy throughout. Basal leaves to 12 cm long, with petiole 4—6 cm long, bipinnately incised with linear or nar- rowly linear, acuminate, terminal lobe; lower cauline leaves also bi- pinnately incised, short-petiolate, with linear auricles at base of peti- oles, middle sessile, pinnately incised, auriculate; apical leaves undivided, linear, 5-10 mm long; terminal lobe of pinnately incised leaves oblong-obovate, oblong or linear, 0.8—4.5 mm wide, with ob- tuse, triangular cusp; all leaves grayish from arachnoid pubescence and sharply scabrous from thin bristles. Capitula solitary, at tips of stem and short branches 1-3 cm long or subsessile, subtended by 1- 2 apical leaves. Involucre oblong-ovate, 12-15 mm long, 5—6 mm in dia, characteristically dichromatic from dark purple involucral bracts and dull colored appendages; outer involucral bracts ovate, about 3 mm long, their appendages cuspidate, longer than 3—4 short fimbriae on each side; middle bracts with 5 veins on outer side, oblong-ovate, 7-10 mm long, 2.5—3.0 mm wide, their appendages to 2 mm long and 2.7 mm wide, decurrent on margin, short-cuspidate, with parallel arcs of very regular, 6-8 fimbriae on each side, appressed to bract; scari- ous auricles below last row of fimbriae not distinct; inner involucral bracts with 3 veins on outer side, narrowly linear, 9-12 mm long, 0.75—2.0 mm wide, with bifid appendage 1.0—1.5 mm long and as wide having short cusp in notch between lobes. Florets pink. Achenes 2.75— 526 524 3.5 mm long, olive-brown with whitish rib; pappus 2.0—2.75 mm long, shorter than achenes. Flowering July to August. River sands.—European Part: Middle Dnieper (Odessa Region, village of Puzhaikovo on Savran River). Endemic. Type in Kiev. Note. So far plants are known only from the reported locality. Series 3. Ovinae Dumb. in Dokl. Akad. Nauk Arm. SSR, V, 2 (1946) 48, ampl. Klok.—Sect. Acrolophus d) Arenariae Hayek, Cen- taurea-Art. Osterr.-Ung. (1901) 4, 94 p. p.—Lithophilous xerophytes. Involucre oblong-ovate to almost cylindrical, 3—6(8) mm in dia, usu- ally considerably narrower and longer than in plants of preceding se- ries. Involucral bracts with 3, less often 5 veins; bract appendages fimbriate with: indistinctly toothed, scarious auricles below crest of fimbriae; pappus considerably shorter than achenes, more or less re- duced. Apparently, a composite series subject to further division. Its Transcaucasian species are distinguished by more ovate involucres and stiff fimbriae on the appendages. 130. C. besseriana DC. Prodr. VI (1837) 585; Ldb. Fl. Ross. II, 2, 705; Klok. in Nauk. Zap. KDU, VII, 6, 72; Dobrocz. in Vizn. Rosl. URSR, 582.—C. arenaria var. besseriana Schmalh. FI. II (1897) 129. Biennial. Stems 1-3, upright or often ascending, 20-45 cr high, branched from base or middle, ribbed-sulcate, tetraquetrous above, along ribs sharply scabrous, with leaves weakly arachnoid-hairy. Basal leaves rather long-petiolate, bipinnately incised with narrowly linear to almost filiform terminal lobe; lower cauline leaves short-petiolate, pinnately or bipinnately incised with lobes to 2 mm wide; apical leaves undivided, narrowly linear. Capitula solitary at tips of stems and short branches, with 1—2 subtending leaves. Involucre ovate cylindrical, 10— 12 mm long and 5—6 mm in dia, weakly arachnoid-hairy; outer involu- cral bracts ovate, 3.0-3.5 mm long, their appendages about 2 mm long, terminating in cusp and on each side bearing upto 6 fimbriae shorter than acute; middle bracts oblong-ovate, 6—8 mm long with 3 less prominent veins on outer side, their appendages 2.0—2.5 mm long, with short cusp and below it with 4—6 regular fimbriae on each side, scarious below fimbriae, with finely toothed auricles; central part of appendages triangular, brownish-yellowish, with dark purple spots at base along veins; inner bracts narrowly linear or linear, to 10 mm long and 2 mm wide, with 1-3 veins on outer side, their appendages ob- long, 2.0—2.5 mm long, narrowed at apex and terminating in very short cusp in shallow notch between two lobes. Florets light purple; corolla of peripheral florets 15-17 mm long, of central florets 10-12 mm 52 — 525 long. Achenes (2.5)2.75—3.25 mm long; pappus 2.0—2.5 mm long, shorter than achenes. Flowering June to August. Limestones and loessic slopes, also on granite outcrops.—Euro- pean Part: Middle Dnieper (southwest), Bessarabia, Black Sea Region (west). Endemic. Described from Podolye. Type (cotype) in Kiev. 131. C. lavrenkoana Klok. in Nauk. Zap. KDU, VII, 6 (1948) 73 (ucrainice), 81 (latine); Dobrocz. in Vizn. Rosl. URSR, 582.—C. ovina auct. fl. ucr. non Pall. Biennial. Stems 1-3, upright, 25—65 cm high, branched above or below middle, sharply scabrous above along ribs, with leaves rather densely arachnoid-hairy. Basal leaves petiolate, 8-14 cm long, bipin- nately incised; lower cauline leaves short-petiolate, pinnately or bipin- nately incised, others sessile, pinnately incised; apical leaves undi- vided; terminal lobe of pinnately incised leaves linear or narrowly linear, acuminate; all leaves acutely scabrous and punctate-alveolate. Capitula solitary at tips of stems and very many branches, 2-5 mm long or to 2 cm long, with 1-2 linear, subtending leaves. Involucre oblong (almost cylindrical), conical at base, 8-11 mm long, 3—5 mm in dia, arachnoid-hairy, often dark purple in upper part, with regularly fimbriate apical appendages; central part of appendage narrowly trian- gular, blackish or dark brown, narrowed into terminal more or less recurved, short cusp; lateral fimbriae lighter in color brownish-yellow- ish; middle bracts oblong-ovate or linear, 4.0—7.5 mm long, 1-2 mm wide, with 1.5—3.0 mm long and as wide appendages terminating in short, weakly recurved cusp, with regular, 4—7 long fimbriae on each side and in addition 2—3 short fimbriae below; inner bracts narrowly linear or linear, 7-10 mm long, 0.75—1.25 mm wide, with apical ap- pendage 1.5—2.0 mm long and 0.5—1.0 mm wide, terminating in cusp to 1 mm long; all bracts three-veined. Florets pink. Achenes 2.5—3.0 mm long; pappus 1.5—2.0 mm long. Flowering June to August. Chalk, limestone and shale outcrops.—European Part: Black Sea Region (east), Lower Don (west). Endemic. Type in Kiev. Note. This species was first noticed by V.M. Czernyaev, who des- ignated it in the herbarium as a separate chalk variety of C. ovina Pall. or C. tenuiflora DC. 132. C. steveniana Klok. in Addenda XXVII, 616.—C. caprina auct. fl. taur. non Stev. Biennial. Stems to 80 cm high, more or less cylindrical and gla- brous below, ribbed, and scabrous above along ribs from fine spinous bristles, with leaves arachnoid hairy more or less strongly branched with divaricate but mostly upward-spreading branches. Basal and lower 528 526 cauline leaves petiolate, bipinnately incised, densely setose beneath spinous bristles with 0.2—0.3 mm long; terminal leaf lobe oblong-lin- ear, oblong-lanceolate or obovate, to 20 mm long and 3—4 mm wide; middle leaves sessile, pinnately incised, with 1—3 pairs of undivided or lobed (submedian leaves) segments. Capitula solitary at tips of branches 3-12 cm long. Involucre oblong-ovate, (8) 10-12 mm long, 5—6 mm in dia; involucral bracts ovate to linear, 3.0—9.5 mm long with 1.0—2.5 mm wide appendages; with 3—5 veins; appendages round to elliptical (in inner bracts), 1.2—2.7 mm wide, with rather soft cusp 0.2—1.2 mm long; appendages of inner bracts with very short cusp not exserted from apical notch, outer and middle bracts with 2—5 pairs of regular whitish fimbriae, about 0.5—2 mm long and wide, more or less toothed or irregularly incised scarious auricles below; inner bracts except cen- tral part with scarious finely toothed auricles separated by notch, cen- tral part of appendage ovate-lanceolate, reddish-brown or dull purple. Florets pinkish-purple; corolla of peripheral florets about 14 mm long, of central florets 11-12 mm long. Achenes 2.5—3.0 mm long and 1.0— 1.4 mm wide, whitish, without or with strongly reduced pappus about 0.5 mm long. Flowering June to August. Coastal shell sands and rubble stopes.—European Part: Crimea (southern coast). Endemic. Described from Eupatoria. Type in Kiev. Note. This plant is known by the name C. caprina Stev. However, the authentic specimens of the latter species, collected by Christian Steven and preserved in the herbarium of the V.L. Komarov Botanical Institute of the Academy of Sciences of the USSR, markedly differ by having a much longer and stiffer, recurved cusp of the appendage and in the over-all habit are more similar to C. diffusa Lam. These, evi- dently, are a hybrid form of C. steveniana x C. diffusa with a predomi- nance of characters of C. diffusa Lam. It is not possible to identify it with our species, which is quite widespread on the southern coast. 133. C. koktebelica Klok. in Addenda XXVII, 617. Biennial. Stems few, upright, 25—70 cm high, almost cylindrical at base, ribbed-sulcate above, acutely scabrous along ribs from conical tubercles, rather densely floccose-arachnoid-hairy when young, later glabrescent, divaricately branched from base, branches numerous. Basal and lower cauline leaves petiolate, pinnately incised to parted, arach- noid-hairy to villous, sharply scabrous beneath and along margin from spinous bristles; terminal lobes (segments) of leaves 2-15 mm long, 1-4 mm wide, acuminate or tiny-cuspidate; middle leaves pinnately divided with linear or narrowly linear, acuminate or cuspidate lobes; apical leaves undivided, linear or narrowly linear, cuspidate, recurved like lobes of middle leaves. Capitula some sessile, others subsessile, Se LEA Plate XXVI. 529 Centaurea polypodiifolia Boiss. 530 528 aggregated, often solitary at tips of branches to 4.5 mm long. Involu- cre oblong ovate, about 9 mm long, 3.5—4 mm in dia; involucral bracts oblong-ovate to narrowly linear, 2.5—-8.5 mm long, with appendages 0.8—1.7 mm wide and 1-5, mostly 3, veins, green at flowering; middle and inner bracts usually purple, their appendages round, 1.5—2.5 mm long, 1.5—3 mm wide, cuspidate; cusp, 0.5—1.3 mm long and (in middle involucral bracts) with 4—6 pairs of regular fimbriae 1—1.5 mm long, with weakly developed toothed scarious auricles at base below fim- briae; central undivided part of appendage triangular, yellowish, often with dark, reddish-brown-purple spot at base; at fruiting entire involu- cre brownish-yellow. Florets purple; corolla of peripheral florets 12— 13 mm long, of central florets, 9-10 mm long. Achenes 3.1—3.2 mm long, about 1.5 mm wide, olive-reddish-brown, with strongly reduced pappus 0.1—0.2 mm long. Flowering August to October. Wormwood steppes.—European Part: Crimea (east). Endemic. Described from vicinity of village of Planersky. Type in Leningrad. 134. C. vicina Lipsky. in Tr. Peterb. Bot. Sada, XIII (1894) 312. Biennial. Stems 35—60 cm high, ribbed, scabrous along ribs from scattered sharp tubercles, densely arachnoid-pubescent, white-tomen- tose below, divaricately branched, with long upward-spreading branches. Basal and lowermost cauline leaves petiolate, bipinnately incised, with linear terminal lobes; middle cauline leaves sessile, pinnately parted, with 3—5 pairs of lateral segments; segments narrowly linear, revolute, sharply scabrous beneath and along margin; floriferous branches 2.5— 11.0 cm long, 5—25, weakly leafy. Involucre oblong-ovate, about 10 mm long and 6 mm in dia, whitish-yellowish, arachnoid-hairy; outer and middle involucral bracts oblong-ovate, 3-7 mm long including appendage, 1.5-3 mm wide, and with 3 inconspicuous veins; append- age broadly ovate, 1.5—2.5 mm long and 2—3.5 mm wide, with cusp to 0.5 mm long, with toothed, scarious auricles at base, fimbriate above with 4—5 fimbriae on each side to 1 mm long; central part of append- age triangular, yellowish, with inconspicuous dull reddish-brown spots at base; inner bracts linear, about 8 mm long, 1—-1.5 mm wide, with oblong-lanceolate appendages 1.2—1.5 mm long. Florets purple pink; corolla to 15 mm long. Achenes oblong-cuneate, 2.3—2.6 mm long, 1- 1.2 mm wide, blackish, with yellowish ribs; pappus 0.3—1.0 mm long, undeveloped. Flowering June to August. Stony slopes.—Caucasus: Western Transcaucasia (in Novorossiisk region). Endemic. Described from Novorossiisk. Type missing (?). 135. C. ovina Pall. in Willd. Sp. pl. I, 3 (1803) 2292; DC. Prodr. VI, 585; Schmalh. FI. II, 122 p. p.; Grossh. Opred. Rast. Kavk. 531 529 501.—C. intacta Ldb. Fl. Ross. II, 2 (1845) 705; Grossh. Fl. Kavk. IV, 219. Biennial. Stems 1—3, upright, about 50-75 cm high, sharply ribbed, deeply sulcate, sharply scabrous along ribs from spinous bristles, more densely arachnoid-hairy, branched from about middle with upward- spreading branches and divaricate branchlets of second and third or- der. Basal and lowermost cauline leaves petiolate, bipinnately incised, grayish from villous arachnoid pubescence, sharply scabrous beneath; terminal lobes oblong-linear to oblong-obovate, to 15 mm long and 0.8—2.0 mm wide, acuminate; middle cauline leaves sessile, pinnately parted with 2—6 segments or divided with small, shortly linear lateral lobes and oblong; terminal lobe to 3—4 mm wide; apical leaves undi- vided, linear, to 20 mm long and about 1 mm wide. Floriferous branches 2-14 cm long; capitula solitary at tips of branches. Involucre oblong- ovate, 10-11 mm long, 5—6 mm wide; outer and middle involucral bracts ovate or oblong, 3—8 mm long including appendage, 1.5—2.5 mm wide, greenish-yellowish, with 3—5 veins, their appendages broadly ovate to triangular, 2-3 mm long, 2.5—4 mm wide, cuspidate, with spinescent cusp 0.9—1.5 mm long, and thin-scarious toothed auricles at base, fimbriate above, with regular 5—6 fimbria on each side, to 1.5— 2 cm long; central part of appendage ovate-triangular, light reddish- brown; inner bracts linear or narrowly linear, 7.5—9.0 mm long, 0.7— 1.7 mm wide, with ovate-lanceolate, scarious appendage 1.5—2.0 mm long, 1.0—1.5 mm wide, lacking cusp and fimbria. Florets light purple; corolla of peripheral florets about 15 mm long, of central florets about 12 mm long. Achenes oblong, about 3.0-3.2 mm long and 1.3 mm wide, pappus about 1.5 mm long. Flowering June to August. Stony montane slopes.—Caucasus: Dagestan, Eastern Trans- caucasia. Endemic. Described from eastern Georgia. Type lost. Note. Apparently an aggregate species, requiring further study. Plants included in this species exhibit a fairly variegated picture of variation in all characters, only partly explained by hybridization. 136. C. gulissashvilii Dumb. in Dokl. Akad. Nauk Arm SSR, V, 2 (1946) 48. Biennial or short-lived perennials. Root string-like, woody, 7—15 mm in dia. Stems 1—5, upright, 25-80 cm high, obtuse-ribbed and sulcate, scabrous along ribs from obtuse tubercles, arachnoid-tomen- tose below, sparsely arachnoid-hairy above, branched almost from base, with long spreading branches. Basal and lowermost cauline leaves 10-— 20 cm long, petiolate, bipinnately incised with incised second order segments, arachnoid-tomentose, especially on petiole or sparsely arach- noid-hairy, sharply scabrous beneath and along margin of lobes from a2 530 spinous bristles; terminal lobes of leaves ovate-triangular to oblong- elliptical or lanceolate, to 7 mm long and 2.0—3.5 mm wide, obtuse or sub-obtusely acuminate; middle cauline leaves sessile, pinnately in- cised with lobed auricle at base and oblong-linear or lanceolate, acumi- nate segments; apical leaves linear or linearly lanceolate, acute; all leaves punctate-glandular. General inflorescence obpyramidal; capitula solitary at tips of less densely leafy branches 2—17 cm long, about 17— 22 mm long including florets. Involucre oblong-ovate, 11—13 mm long, 6-8 mm in dia, dichromatic; involucral bracts yellowish-brownish, outer ones with appendages, about 3—4 mm long; appendage weakly demar- cated from bract, lanceolate, yellowish-brown with 2-3 pairs of stiff, whitish fimbriae and larger cusp; middle bracts ovate, 4-7 mm long, 3—3.5 mm wide, with 5—7 veins, and scarious border, their appendages broadly ovate, about 3—4 mm long, 4—5 mm wide with about 0.7—2 mm long cusp, 4—7 pairs of regular fimbriae, 1.5—2.5 mm long, and with scarious, toothed or irregularly incised auricles, at base; tips of ap- pendages cuspidate, recurved; central part of appendage dark reddish- brown. Florets about 15 mm long. Achenes oblong, 3—3.4 mm long, 1.2—1.5 mm wide, olive-brown (mature), with pappus1.1—1.5 mm long. Flowering July to September. Stony slopes and sands.—Caucasus: Southern Transcaucasia. Gen- eral distribution: Probably in Armenia and Kurdistan. Described from Stepanavan (gorge of Dzoraget River). Type in Yerevan. Note. C. demetrii Dumb. (op. cit.), according to the authentic description and the key in Opredelitel Rastenii Kavkaza [Keys to the Plants of the Caucasus] by A.A. Grossheim, differs from C. gulissashvilii Dumb. by having much larger capitula. We did not see the specimens of this species and could not form a definite opinion about it. This species has been described from the Dagestan plains. Series 4. Cylindraceae (Hayek) Dumb. in Dokl. Akad. Nauk Arm. SSR, V, 2 (1946) 50.—Sect. Acrolophus 1) Cylindraceae Hayek, Centaurea-Art. Osterr.-Ung. (1901) 4, 106.—Involucre cylindrical, 2.5— 3.5 mm in dia; appendages of involucral bracts lanceolate, pectinate- fimbriate, terminating in stiff spine, often more or less recurved; pappus usually very short or undeveloped. 137. C. diffusa Lam. Encycl. meth. I (1783) 675; DC. Prodr. VI, 586; Ldh. Fl. Ross. II, 2, 706; Boiss. Fl. or. III, 650; Schmalh. FI. II, 29; Grossh. Fl. Kavk. IV, 220; Iljin in Fl. Yugo-Vost. VI, 433; Dobrocz. in Vizn. Rosl. URSR, 582.—C. parviflora Bess. Enum. pl. Volhyn. (1822) 35, non Desf.—Ic.: Hayek, Centaurea.-Art. Osterr.-Ung. Taf. VIII, fig. 7—Exs.: GRF No. 421; Fl. exs. reip. Bohem.-Sloven. Nos. 637, 889; Fl. exs. austro-hung. No. 234. 33)3) 531 Biennial. Stems upright or ascending at base, 15-50 cm high, ribbed-sulcate, strongly branched from base with divaricate branches, with leaves acutely scabrous and weakly arachnoid-hairy. Basal and lower cauline leaves petiolate, bipinnately incised into narrowly linear, acuminate lobes, other leaves sessile; middle cauline leaves pinnately incised; apical leaves undivided, linearly lanceolate, 1-3 mm wide. .Capitula solitary at tips of very many branches, 0.5—1.0 or 2.0—3.5 cm long leafy to capitula. General inflorescence paniculate. Involucre ovate- cylindrical 8-10 mm long, 2.5—5 mm in dia; outer and middle involu- cral bracts greenish or yellowish, with distinct keel in middle and 2 lateral veins on outside; appendages of involucral bracts decurrent on scarious margin, in form of hard brownish triangles with 4—6 stiff fimbriae on each side 1-2 mm long, cuspidate (spiny); spine 3—4 mm long, somewhat recurved; fimbriae and spine yellowish or reddish- brown; inner bracts dull green, linear, with scarious border, and ob- long, obtuse or acuminate, entire or irregularly toothed scarious ap- pendage. Florets dull pink or whitish, less often light purple. Achenes 2.5—3.0 mm long, brownish, glabrous; pappus scarcely noticeable, consisting of fewer hairs, or absent. Flowering July to September. Open places, on stony slopes and coastal sands, as weed.—Euro- pean Part: Middle Dnieper, Bessarabia, Black Sea Region, Volga-Don (south), Lower Don, Crimea; Caucasus: all regions? Often introduced in north and other regions not mentioned above. General distribution: Balkans-Asia Minor. As introduced plant in many regions of southern and Centrral Europe. Native of Asia Minor. Described from western Europe. Type in Paris. Note. Very readily hybridizes with many species. Hybrids are known with C. micranthos Gmel., C. majorovii Dumb., C. lavrenkoana Klok., -C. sterilis Stev., C. margarita-alba Klok., C. protomargaritacea Klok., C. paczoskyi Kotov ex Klok., and others. Some of these hybrids have been described by me as separate species, which is indicated in the corresponding places in the treatment of this subgenus and the subgenus Phalolepis (Cass.) Dobrocz. 138. C. aemulans Klok. in Addenda, XXVII, 617.—C. squarrosa auct. fl. taur. non Willd. Biennial. Root string-like, vertical, strong. Stem solitary, upright, 30-60 cm high, cylindrical in lower part with thin ribs and furrows, ribbed and deeply sulcate above especially on branches, more or less densely setose along ribs with spinous bristles, rather densely arach- noid-hairy, with leaves arachnoid-tomentose below, branched from base, with numerous strongly divaricate branches. Basal and lowermost cauline leaves petiolate, with pinnately parted or partly undivided 534 532 segments; terminal lobes oblong, oblong-lanceolate, or obovate, 3—15 mm long, 1-4 mm wide, narrowed at apex into short fine cusp, densely setose along margin and beneath with spinous bristles 0.2-0.3 mm long; middle and upper leaves sessile, pinnately parted, with oblong or linearly lanceolate lobes, attenuate into prickly cusp, their terminal lobe larger than lateral ones, sometimes oblong-obovate; apical leaves at tips of branches oblong-linear or narrowly linear, 1.5—2 mm long and 0.3—0.4 mm wide (below capitulum). Capitula more or less clus- tered at tips of second-order branches, subsessile or peduncles not surpassing involucres, less often to 2 cm long. Involucre almost cylin- drical, 10-12 mm long, 3—3.5 mm in dia, stramineous, weakly arach- noid-hairy; involucral bracts oblong to narrowly linear, with append- age 2.5-10 mm long, 0.5-—1.5 mm wide, and 1-3 inconspicuous veins; appendages 2—6 mm long, with apical spine 0.7-4.0 mm long and 3- 7 pairs of stiff fimbria 0.3—1.7 mm long; prickly cusp of appendages deflexed and in outer bracts more or less recurved. Corolla purple, pale on withering, 10-12 mm long. Achenes 2.3—2.5 mm long, 0.9-1.1 mm wide, glabrous, lacking pappus. Flowering July to October. Wormwood steppes and as weed in open places.—European Part: Black Sea Region (Melitopol), Crimea (eastern part). General distri- bution: Not determined. Possibly it is endemic to Crimea. Described from vicinity of village of Planersky (Koktebel). Type in Leningrad. Note. This plant differs quite clearly from C. diffusa Lam. by having strongly crowded capitula, longer involucres, and longer and stiffer, mostly recurved apical spines on the appendages, as well as persistently purple florets. Because of these differences, Crimean flora specialists have considered it to be C. squarrosa Willd. The latter species was reported for Crimea by S.S. Stankov in the second edition of Opredelitel [The Keys] (1957) 419, apparently in place of our spe- cies, since C. squarrosa does not grow in Crimea. 139. C. caspia Grossh. Fl. Kavk. IV (1934) 219; Opred. Rast. Kavk. 500; Dumbadze in Dokl. Akad. Nauk Arm. SSR, V, 2, 50 (latine). Biennial. Stem 20—40 cm high, ribbed above, scabrous along ribs from conical tubercles, entirely villous-arachnoid-hairy, divaricately branched. Basal and lower cauline leaves petiolate, bipinnately parted, with linear or oblong to oblong-lanceolate, obtuse, or subobtuse, acumi- nate, terminal lobe 1.5—7.5 mm long, 0.4—2 mm wide, arachnoid-hairy, sharply scabrous beneath and along margin; cauline leaves sessile, auriculate, bipinnately incised and (in upper part) pinnate, lobes linearly lanceolate to narrowly linear, 1.5—9.0 mm long and 0.5-1.5 mm wide, acuminate or subobtusely cuspidate. Capitula solitary at tips of rather short (to 5 cm long) branches. Involucre ovate-cylindrical, 353 535 10-15 mm long, 4—6 mm in dia, almost monochromatic, villous-arach- noid; involucral bracts oblong-ovate to narrowly linear appendages 2.5-13.0 mm long, with 1.0-1.7 mm wide with 3 veins, greenish; appendages of outer and middle bracts ovate to round, 1.5—-4.0 mm long, 1-4 mm wide, with 0.5—2.0 mm long cusp, fimbriate, with regu- lar, like cusp yellowish fimbriae 0.4-1.7 mm long, 3—9 on each side, more or less recurved; central thick part of appendage triangular, ochre- ous; appendage of inner bracts ovate-lanceolate, 1.5—3.0 mm long, 0.4—2.5 mm wide, toothed. Corolla purple-pink, of peripheral florets 18-19 mm long, of central florets 15-16 mm long. Achenes oblong, about 3.5 mm long, 1.2 mm wide, brown; pappus 2.5—3.0 mm long. Flowering June to July. Coastal sands.—Caucasus: Dagestan, Eastern Transcaucasia, Talysh (all along Caspian Coast). General distribution: Possibly found in northern part of Iran. Described from Dagestan. Type in Baku (?). Note. A unique species, maybe not even belonging to this series. Series 5. Squarrosae Klok.—Sect. Acrolophus m) Squarrosae Hayek, Centaurea-Art. Osterr. Ung. (1901) 5 p. p. nomen.—Weakly differing from the preceding series. Capitula partly aggregated in small clusters at tips of branches; appendages of involucral bracts much stiffer, arcuate at tip. 140. C. squarrosa Willd. Sp. pl. II, 3 (1803) 239; Grossh. FI. Kavk. IV, 220 and Opred. Rast. Kavk. 500; Dumbadze in Dokl. Akad. Nauk Arm. SSR, V, 2, 50.—C. virgata var. Squarrosa Boiss. FI. or. III (1875) 651.—Acrolophus Squarrosus (Willd.) Nevski in Tr. Bot. Inst. Akad. Nauk SSSR, Ser. 1, 4 (1937) 291. Biennial. Stems solitary or few, upright, about 35-70 cm high, ribbed-sulcate, scabrous along ribs, quite densely arachnoid-hairy, divaricately branched almost from base or middle. Basal and lower cauline leaves petiolate, bipinnately incised, not persisting at flower- ing time; middle leaves sessile, auriculate, pinnately incised with 1—5 pairs of lateral segments, arachnoid-hairy, very weakly or strongly scabrous beneath and along margin from undeveloped bristles, leaf lobes divergent, linear to oblong-lanceolate 2-18 mm long, 0.5—2.2 mm wide, obtusely acuminate or cuspidate; apical leaves undivided, oblong-linear to obovate, 1-22 mm long, 0.4-3.7 mm wide. Capitula numerous, at tips of short branches to 1.5—4 cm long or partly often sessile and crowded at tips of branches. Involucre oblong-ovate to almost cylindrical, 9-11 mm long, 2.5—3.5 mm in dia, usually dichro- matic, mottley, arachnoid-hairy; involucral bracts oblong-ovate to lin- ear, 2-10 mm long, 0.8—2 mm wide, yellowish or pale purple, with 3 536 534 often inconspicuous veins; appendages of involucral bracts broadly ovate to oblong-elliptical (in inner bracts), 1.5—3.5 mm long, 1-4 mm wide, at apex (in outer and middle) with yellow, prickly cusp 0.5—2 mm long, and on sides with stiff, yellowish fimbriae to 1.2-1.7 mm long, 4-6 on each side; central part of appendage lanceolate, very thick, yellowish or often dull to dark purple; cusp on middle append- ages recurved; appendages of inner bracts semi-membranous, finely toothed. Florets pink or pinkish-purple; corolla 9-14 mm long. Achenes 2.4—4.2 mm long, 1.3—1.7 mm wide, olive-dark-brown with yellowish stripes on ribs; pappus 1.0—-2.7 mm long. Flowering June to August. Stony slopes and weedy places.—Caucasus: Dagestan, Eastern and Southern Transcaucasia, Talysh; Soviet Central Asia: mountainous Turkmenia, Pamiro-Alai, Tien Shan. General distribution: Asia Mi- nor, Armenia and Kurdistan, Iran, Afghanistan. Described from “the East”. Type in Berlin. Note. This species is highly variable and probably is an aggregate. T.A. Dumbadze described four varieties from the Caucasus, which, however, do not have their own ranges. Series 6. Aggregatae Dumb. in Dokl. Akad. Nauk Arm. SSR, V, 2 (1946) 50.—Sect. Acrolophus m) Squarrosae Hayek, Centaurea-Art. Osterr.-Ung. (1901) 5 p. p. nomen.—Montane xerophytes; leaves lyrate or pinnately incised, with large terminal lobe; capitula in dense clus- ters of 5—10 at tips of primary branches; involucre almost cylindrical; pappus not shorter than achenes. The Iranian species differ sharply from the other members of the subgenus. 141. C. albida C. Koch. in Linnaea, XXIV (1851) 433; Grossh. Fl. Kavk. IV, 219 and Opred. Rast. Kavk. 500; Dumbadze in Dokl. Akad. Nauk Arm. SSR, V, 2, 50.—C. aggregata var. albida Boiss. F1. or. III (1875) 653.—Exs.: Fl. cauc. exs. No. 223. Biennial, perennial. Stems few or solitary, upright, 20-70 cm high, ribbed, smooth, with leaves entirely white-arachnoid-tomentose, branched above middle, with upward-spreading branches. Basal and lower cauline leaves petiolate, bipinnately incised or sometimes pin- nately parted; terminal lobe of leaves oblong or oblong-obovate, 3-20 mm long, 1.5—5 mm wide, attenuate into short, triangular cusp; middle cauline leaves pinnately parted to almost lyrate, with large, oblong- elliptical terminal lobe, like lower cauline leaves scabrous or setose beneath and along margin; terminal lobe smooth, oblong, 2-5 mm long, 0.8—1.5 mm wide, obtusely cuspidate, like lobes of middle leaves. Capitula usually all sessile, aggregated in dense clusters of 5—10 at 535 537 tips of branches. Involucre ovate, cylindrical, weakly narrowed toward apex, 11-12 mm long, 3—3.5 mm in dia, monochromatic, reddish-brown- yellowish; involucral bracts ovate to linear, 2—9 mm long, 1.0—1.5 mm wide; with 3 prominent veins, greenish or yellowish, weakly arach- noid-hairy; appendages 1.0—2.5 mm long, 1—4 mm wide, outer and middle ones with acute angled, reddish-brown, triangular, central part and short (0.4-0.7 mm long) but rather thick cusp and 2-4 pairs of regular yellow fimbriae. Corolla light-purple, 12-13 mm long. Achenes oblong, 2.5—3.0 mm long, 1.2—1.5 mm wide, olive-brown; pappus 2.5— 3.5 mm long. Flowering June to August. Stony mountain slopes.—Caucasus: Western Transcaucasia (south). Specific localities from beyond USSR unknown. General distribution: Armenia and Kurdistan. Described from former Artvin District. Type in Leningrad. 142. C. aggregata Fisch. and Mey. in DC. Prodr. VI (1837) 585; Grossh. Fl. Kavk. IV, 219 and Opred. Rast. Kavk. 500; Dumbadze in Dokl. Akad. Nauk Arm. SSR, V, 2, 50. Biennial, perennial. Stems solitary or few (2—8), upright or almost ascending, 40-100 cm high, ribbed-sulcate, sharply scabrous along ribs in upper part, less densely floccose-arachnoid-hairy, branched mostly above middle but often also from below. Basal and lower cauline leaves petiolate, pinnately incised, with large, oblong elliptical, termi- nal lobe and similar or oblong, much smaller lateral lobes, white- tomentose beneath when young, later glabrescent, green; cauline leaves mostly lyrate with very large terminal lobe and small oblong to linear lateral lobes, scabrous-setose on both sides and with traces of arach- noid-tomentum disappearing by flowering time; apical leaves smooth, oblong, at apex attenuate into very short, triangular cusp, as also many lobes of lower leaves. Capitula sometimes solitary on short branches, mostly aggregated at tips of primary branches, in dense clusters of 3— 10, and then sessile. Involucre oblong-ovate or cylindrical, 10-13 mm long, 3—4 mm in dia, monochromatic, reddish-brown-yellow or mottley; involucral bracts ovate to linear, 2.7—10.5 mm long (including append- age), 1-2 mm wide, with 3 prominent veins, greenish, yellowish, or inner ones often pale purple; appendages broadly ovate (in outer bracts) to round and oblong-lanceolate (in inner bracts), 1.5—3.0 mm long, 0.7-5 mm wide, light yellow or often reddish brown in outer and middle bracts, with short, somewhat thick cusp 0.4—0.7 mm long and pectinate, 4—7 pairs of regular fimbriae along margin 0.5—2 mm long; appendage of inner bracts scarious, toothed. Corolla light-purple, of peripheral florets 14-16 mm long, of central florets 11-12 mm long. 538 539 536 Achenes 2.5—2.9 mm long, 1.3-—1.4 mm wide, light reddish-brown, with whitish ribs; pappus 2.5—2.9 mm long. Flowering July to August. Stony slopes and meadows; in mid-montane zone—Caucasus: Southern Transcaucasia. General distribution: Iran. Described from bordering regions of Iran. Type in Leningrad. Subgenus 16. Phalolepis (Cass.) Dobrocz. in Bot. Zhurn. Akad. Nauk URSR, VI, 2 (1949) 63.—Phalolepis Cass. in Dict. Sc. nat. L (1827) 248.—Centaurea sect. Phalolepis (Cass.) DC. Prodr. VI (1837) 568. p. p.—Capitula mostly quite large; involucral bracts coriaceous with large, round, entire or weakly toothed appendages; corolla pink, purple, light yellow, or white; pappus as long as achene, slightly longer, or much shorter. Biennials. rarely perennials; leaves arachnoid-hairy, simple, pinnately or bipinnately incised. Type of subgenus: C. alba L. -1.Appendages [of involucral bracts] not sharply differentiated into central thick part and scarious border (former gradually merging into latter); pappus more or less as long as achene, much longer, or slightly shorter. Obligate psammophytes of steppe zone of Euro- pean part of USSR (one species in this zone found on granite outcrops):is fees A onoladomam 2h ede eas ae ee Dy + Appendages [of involucral bracts] more or less sharply differenti- ated into central thick coriaceous part with all veins passing through it and thin scarious, entirely transparent border; pappus always much shorter than achene (usually half or less). Lithophilous plants of mountains of Crimea and! Caucasus’. ..22822.22R0 RS... 14. 2. Involucre 8-10 mm long; appendages of middle involucral bracts cuspidate. Achenes 2.0—2.5 mm long with distinctly shorter pappus. Growing on granite outcrops .......... 143. C. pseudoleucolepis Kleop. + Involucre much larger (if not so large, then its middle bracts not cuspidate); achenes also with larger pappus. Growing on sands... 3 3. Appendages of involucral bracts silky-lustrous, almost entirely opaque, and translucent only along margin .............. cc eeeeeeeeeeees 4. + Appendages of involucral bracts with oily or micaceous luster, mostly transparentiandsthinyithickionlyabbase. cunt 6. 4. Corolla creamish, almost white, never pink; achenes lenticular, whitish, not blackening at maturity ...... 145. C. margaritacea Ten. + Corolla purple or pink, sometimes very pale but only in individual specimens; achenes almost tetraquetrous, blackening at maturity .... 5. Involucre globose, 15—20 mm in dia, appendages of involucral — bright white; corolla;purpleybmehtiwd 25 .2eeek. SA ee 10. 540 11. 53) MITE DROITS, OF Cs ees 146. C. margarita-alba Klok. Involucre somewhat oblong, 11—16 mm long, to 15 mm in dia. appendages of involucral bracts yellowish to light orange; corolla Pinks Pale’. IG 144. C. proto-margaritacea Klok. Appendages large, to 10 mm long and wide, very thin, scarious, dorsally carinate and cuspidate; narrow band of opaque tissue of appendage extending along keel up to very tips; all or at least some appendages incurved at apex, forming unique, divergent valves, but HAtENSA WE LO WL EE, HAR nae I wt 2 SNe 53, 8)5 a. Appendages small, less thin, keel and cusp not developed or incon- spicuous; opaque tissue of appendage triangular, basal but not reach- ing up to apex; appendages more or less converging and not incurved APABORS TOPMUN S Val VES ecvcasscsnscins EN I le RS cc sade 8. Corolla creamish, almost white; involucre large, upto 25 mm in dia, with spreading appendages; stem sharply scabrous above .............. RAEI, Sta horde th EUR, GSS, me gad, Mas 148. C. appendicata Klok. Corolla pink; involucre smaller, up to 16-17 mm in dia, mostly with somewhat flat appendages; stem smooth above ..............::ccee8 ok SAORI So R a et anne San) Cte 147. C. konkae Klok. Corolla pink or pale purple; appendages, at least in inner involucral bracts, dichromatic, with dark purple basal spot; stem more or less Scabrous*above Glens TrOSeWA Ne Se es cos RP i 9. Corolla creamish, almost white or (in one species) light yellow; appendages usually monochromatic, yellowish or light yellow; stem Sninotlirabovel wawwlinioot. jm. atealh, dell ce ee ee, Ss 12. Appendages yellowish, stiff, strongly convex, almost hemispheri- cal, with dark inconspicuous spot at base or entirely monochro- matic, corolla pale pink to almost White .0..........ccceeccsseeeeeeceeeneeneeeeeees ES Se an oy Se eas 149. C. proto-gerberi Klok. Appendages whitish or greenish softer and less convex, almost all with well-developed dark basal spot; corolla brighter ................ OF Involucre about 10-13 mm long and 6-10 mm in dia; appendages of middle involucral bracts elliptical, greenish, plants tall (up to 100 cm or more) with slender, virgate stems and long lower leaves divided into very long, narrow segments ............::ccscccssccsseeesseeeeeees RIO Sih ROSEN. teawesdaa rea viandnasee i eiadssaainaseannaeatnns 152. C. breviceps Iljin Involucre longer, appendages of middle involucral bracts more or less round, not greenish. Plants shorter with much thicker branches and smaller leaves with shorter, equally broad lobes or segments .... Involucre subglobose, 12-15 mm long, 10-16 mm in dia; achenes 4-5 mm long, light brown at maturity, not blackening ............. ed ac ee Ree COMP te eRe MP ye 6a 150. C. donetzica Klok. 538 12. 13% 14. Ls? 16. Involucre ovate or oblong-ovate, 15-18 mm long, 10-14 mm in dia; achenes 3.5-3.7 mm long, blackening at maturity EERE TIE TE BOSC, BUA STN. ARE REeN Oe 115. C. pineticola Ijin. Involucre 15-27 mm long, 15—25 mm in dia, often compressed globose; involucral appendages weakly convex and entirely cover- ing bracts, large, in central florets about 6-7 mm long and 8-9 mm wide Tkiiee eee ER, cele Sad eet ee eh 155. C. dubjanskii [jin Involucre smaller and not compressed above; involucral append- ages strongly convex and not entirely covering bracts, smaller and notawider thamlonpry. ik ted ye. ghee ee Bee ee 13. Corolla creamish; appendages of involucral bracts yellowish, all monochromatic. Mature achenes blackish; pappus somewhat shorter than achene, 3=4°mm long <..icc..0...-.ese.ee reese 154.C. gerberi Stev. Corolla light yellow; appendages of involucral bracts darkish yel- low, middle and inner often with purple spot at base. Mature achenes whitish; pappus much longer than achene, 4.5—7.0 mm long ....... ODL RELL. Nod seGa 8. Hat ROE SO 153. C. paezoskyi Kotov. ex Klok. All appendages of involucral bracts with more or less developed cusp; capitula fewer (to 10 per stem). Plants of Crimea ............... Middle and inner involucral bracts with appendages entirely lack- ing cusp; capitula more numerous. Plants of Caucasus (and one eastern CrintedmiSpecies)iz..Auiiys acne. eee alee eee | Wie Involucre ovate to oblong-ovate, 10-14 mm long, 6—10 mm in dia; stem to 100 cm high. Plants of foothills or southern coast but not alpinepusually biennial re oe. ee 158. C. sterilis Stev. Involucre ovate or broadly ovate to globose, much larger; stem to 35cm igh. ‘Perennialvalpinesplants vert: alee etek eee 16. Involucre ovate or broadly ovate, 14-16 mm long, 10-14 mm in dia; appendages of middle involucral bracts about 3-4 mm long, with large ovate-triangular dark reddish-brown basal spot ............. NOUS CLES. S55, EE UT ee ee dre oes: 157. C. vankovii Klok. Involucre broadly ovate or subglobose, 16—22 mm long, 12—21 mm in dia; appendages of middle involucral bracts about 10 mm long, with oblong-lanceolate light reddish-brown dorsal spot ................. ci Me A on se tn Maman ferent het net 156. C. semijusta Juz. Involucre 9-10 mm long and 4—6 mm in dia; appendages of middle involucral bracts 2.5—3.0 mm long ............. 164. C. alexandri Bordz. Involucre and appendages much larger ...............ceceeeseeseeeneeseees 18. Appendages distinctly toothed to almost fimbriate; achenes 3.5—4.5 mm long. Plants of eastern Crimea and Western Transcaucasia ....... eee eee meee cere teres eee eee eee eae ee eee EEE OSES EEE REET EH OES EES OEE EES ESEES TEESE OSES ESEEESESSSESE ESTEE EEE 541 539 + Appendages entire or at places indistinctly toothed; achenes 2.5— 3.0 mm long. Plants of Eastern and Southern Transcaucasia ...... 2M. 19. Segments or terminal lobes of basal and divided cauline leaves to 6 mm wide, linear- or oblong-lanceolate. Involuere 10-16 mm in dia, yellowish, with monochromatic (lacking dark basal spots) ap- pendages; appendages of middle involucral bracts 5-6 mm long. Plants of eastern’‘@rimeae2 eae 159. C. sarandinakiae []lar. + Segments or terminal lobes of divided leaves 0.5—-3.0 mm wide, narrowly linear to linear-lanceolate. Involucre 8-12 mm in dia, whitish or yellowish-light reddish-brown (predominant color); ap- pendages of involucral bracts with dark (reddish-brown) basal spots; appendages of middle bracts 3-4 mm long. Plants of Western ‘PrahnsCaucasias..2 5. Se A ee BANAL 20. DOM Univoliicre whitish!.2..2.220 Jee 160. C. novorossica Klok. + Involucre yellowish-light-reddish-brown or reddish-brown-light By SLLCIOVAR: Se ERE Sorel LE 161. C. georgica Klok. 21. Segments or lobes of basal and lower cauline leaves linear-lan- ceolate, about 1.5—2.0 mm wide; segments of middle cauline leaves 1-3 mm wide. Involucre compressed, ovate, 12-13 mm long, 11- Domina ee. Ae 163. C. transcaucasica Sosn. and Grossh. + Segments of basal and lower cauline leaves pinnatifid, ovate in outline with oblong-ovate (and obovate) lobes, 1.5—5.0 mm wide; segments of middle cauline leaves oblong, 1-5 mm wide. Involucre broadly ovate to subglobose, 11-14 mm long, 10—15 mm in dia, MOt COMIPRESSEMe AN tek. SVE lect rite. dente 162. C. latiloba Klok. Section 1. Pseudophalolepis Klok. in Addenda XXVII, 618; Dobrocz. in Bot. Zhurn. Akad. Nauk URSR, VI, 2 (1949) 63, 65 (ucrainice).—Exclusively biennials. Appendages of involucral bracts almost entirely membranous or scarious with small, triangular thicker part at base, less often with narrow band in middle, lacking distinct dark spot, less often with spots, thicker part of appendage indistinctly separated from scarious part; pappus more or less as long as achene, less often slightly longer or much shorter; stems more or less strongly branched; corolla pink, pale purple, creamish or sometimes light yel- low. Type of section: C. gerberi Stev. A palaeopontic relict section, native exclusively to the steppe zone of eastern Europe. Except for one, all the known species of this sec- tion are obligate psammophytes; one species grows on granite out- crops. All species appear to be rare and more or less narrowly endemic plants. 542 543 540 Series 1. Pseudoalbae Dobrocz. in Ukr. Bot. Zhurn. XIX, 1 (1962) 43, descr.; in Bot. Zhurn. Akad. Nauk URSR, VI, 2 (1949) 63, nomen.— Capitula smaller; appendages of involucral bracts transparent, toothed- fimbriate, dorsally carinate, and with small, soft cusp; pappus shorter than in members of other series. Only one species belongs to this series. It differs sharply from all other members of the section and approaches section Phalolepis in some characters. 143. C. pseudoleucolepis Kleop. in Izv. Kievsk. Bot. Sada, IV (1926) 26; Dobrocz. in Vizn. Rosl. URSR, 578. Biennial. Stem upright, about 50 cm high, strongly branched above middle, with leaves weakly arachnoid-hairy. Leaves pinnately and bi- pinnately incised with narrowly linear, ‘acute, terminal lobe; lower leaves long-petiolate, middle and upper ones sessile, uppermost leaves with larger, narrowly lanceolate, central lobe and 2 small lateral lobes at base. Capitula small, solitary at tips of quite long, leafy branches, forming lax paniculate inflorescence. Involucre ovate, 8-10 mm long, 7-8 mm in dia with conical base [obconical]; involucral bracts yellow- ish-green, with scarious border; appendages of outer and middle bracts round, convex, transparent, thicker in middle up to tip, opaque, very short-cuspidate; inner bracts with ovate, scarious, apical appendages. Corolla pale pink. Achenes 2.0—2.5 mm long, dark brown, with whitish ribs; pappus shorter than achene, 1.0—2.5 mm long. Flowering July to August. Granite outcrops.—European Part: Black Sea Region (only in Azov Region, in “Kamennye mogily” reserve of Ukrainian Academy of Sciences, former Rozov District, Zaporozhye Region). Endemic. De- scribed from locality mentioned above. Type in Kiev. Note. This species and the unique yarrow, Achillea glaberrima Klok, growing together with it, are strictly endemic to the reserve, covering an area of about 4 km’. Both probably were distributed widely at sometime in the Ukrainian low hills area of the Azov uplands but persisted only in the indicated locality, thanks to the more favorable moisture conditions. Series 2. Margaritaceae Klok.—Appendages. of involucral bracts yellowish or lustrous white, silky, almost entirely opaque, transparent only along margin, and with irregular small teeth, convex, with promi- nent dorsal keel, incurved at apex. Stem tall, very strongly branched. All three species of this series are known only from sands of the lower reaches of the Southern Bug River. The series is undoubtedly relictual, becoming extinct. 544 541 144. C. proto-margaritacea Klok. in Tr. Inst. Bot. KhDU, I (1936) 81, 102; Dobrocz. in Vizn. Rosl. URSR, 578.—C. klokovi Oppermann ex Klok. op. cit. Biennial. Stem upright, 45-80 cm high, strongly branched, with leaves rather weakly arachnoid-hairy, smooth. Leaves pinnately and bipinnately incised (lower), with narrowly linear to almost filiform lobes, uppermost leaves narrowly linear, undivided, subtending ca- pitula. Capitula solitary, at tips of leafy branches. Involucre short- elliptical or subglobose, 11-16 mm long, 11-15 mm in dia; appendage of involucral bracts yellowish or light orange, membranous, almost entirely opaque, transparent only along margin at base, round or ellip- tical, to 5-6 mm long and 4-5 mm wide; appendages of middle and inner bracts with dark spot at base, usually convex at flowering time, incurved. Corolla pink to almost white (creamish). Achenes 2.8—4.0 mm long, usually about 3.5 mm, blackish with whitish ribs; pappus to 3 mm long, always distinctly shorter than achenes. Flowering July to September. Sands.—European Part: Black Sea Region (only on sands of Bug estuary below Nikolaev). Endemic. Described from vicinity of Nikolaev. Type preserved in Kharkov; cotype in Leningrad. Note. We have already expressed the opinion (1936) that this species is a progenitor of both of the following members of this series. 145. C. margaritacea Ten. Syll. Fl. Napol. Add. 3 (1831) 628; Klok. in Tr. Inst. Bot. KhDU, I, 84, 100; Dobrocz. in Vizn. Rosl. URSR, 578.—C. splendens Lang. and Szovitz in sched, non L.—Exs.: Lang and Szovitz, Herb. ruth. No. 115 (sub C. splendente L.). Biennial. Stems upright, 50-120 cm high, strongly branched, with leaves densely arachnoid-hairy, ribbed above and sharply scabrous on ribs. Leaves mostly pinnately incised with narrowly linear lobes 1—2 mm wide; lower leaves bipinnately incised with oblongly linear termi- nal lobe to 3.5 mm wide; uppermost leaves undivided, linear, subtend- ing capitulum, usually shorter than involucre. Capitula solitary at tips of leafy branches. Involucre compressed-globose, 15-18 mm long and 20-22 mm in dia, yellowish; appendages of involucral bracts round, 6—7 mm long, 7-9 mm wide, silky, yellowish, almost opaque, only at base along margin transparent, translucent with irregular small teeth, dorsally distinctly carinate, terminating in soft, very short cusp in shallow notch at tip of appendage; appendages more or less convex at flowering time, incurved, forming acute head; appendages of middle bracts more or less lacerate. Corolla light creamish, almost white. Achenes 3.5—4.25 mm long, lenticular, whitish or light brown, mono- 542 chromatic, not blackening at maturity; pappus to 4.5 mm long, some- what longer than achene or as long. Flowering July to August. Sands.—European Part: Black Sea Region (only on right bank sands of Southern Bug River above Nikolaev). Endemic. Described from vicinity of Nikolaev, evidently from specimens distributed by Lang and Szovitz. Type probably in Naples; cotypes in various places. Note. This very rare species has not been found by anyone since P.A. Oppermann, who collected it in 1928 and 1931 between the vil- lages of Solonikh and Pristen. In 1955, D.N. Dobroczaeva conducted an extensive search at this locality but was unable to find it and con- sidered that possibly it was already extinct. 146. C. margarita-alba Klok. in Tr. Inst. Bot. KhDU, I (1936)81, 100; Dobrocz. in Vizn. Rosl. URSR, 578.—C. margaritacea auct. non Ten.: Schmalh. FI. II (1897) 121 p. min. p.; Iljin in Izv. Glavn. Bot. Sada SSSR, XXVI, 1 (1927) 4. Biennial. Root string-like, rather strong. Stem upright, 45—85 cm high, strongly branched above middle, with leaves floccose-arachnoid- hairy, smooth, scabrous only below capitula. Leaves mostly pinnately incised with filiform or narrowly linear lobes to 1 mm wide; lower leaves bipinnately incised with wider lobes, uppermost ones narrowly linear, undivided or with 1—2 teeth at base, subtending capitulum. Capitula solitary at tips of branches. Involucre globose, 15—20 mm wide; involucral bracts coriaceous, yellowish, entirely covered by appendage; appendages larger, to 6-8 mm long, and 8-10 mm wide, membranous, opaque and transparent only along margin at base, pearl- white, silky-lustrous, lacking dorsal keel but with distinct longitudinal vein in middle and often with small triangular ochreous basal spot (more prominent in inner bracts), with fine teeth, with less conspicu- ous notch at apex, and short soft cusp; most appendages at flowering time involute at apex resembling acute head. Corolla purple. Achenes 3.5-4.0 mm long, blackish with whitish ribs and base; pappus 3-4 mm long, as long as achene or slightly shorter. Flowering July to September. Sands.—European Part: Black Sea Region (only on left bank sands of Southern Bug River in immediate environs of Nikolaev). Endemic. Described from vicinity of Nikalaev (nursery). Type preserved in Kharkov; cotypes in Kiev and Leningrad. Note. A very rare plant with unusually ornamental involucres. It is in extreme need of in situ protection (almost extinct!) and introduc- tion into cultivation. It hybridizes with C. diffusa Lam.; one such hybrid form is described by I.K. Paczoski as a separate species, C. hypanica Pacz. 545 546 543 Series 3. Appendicatae Klok.—Appendages of involucral bracts large, scarious, with oily luster, mostly transparent and thin, thick only at base but with narrow band of opaque tissue, reaching apex, with small ochreous or dark purple spot at base, dorsally with inconspicu- ous keel and with short cusp, not convex in involucre, somewhat flat or incurved at apex as acute valve. Two endemic Dnieper species belong to this also very distinctive and relictual series. 147. C. konkae Klok. in Tr. Inst. Bot. KhDU, I (1936) 93, 100; Dobrocz. in Vizn. Rosl. URSR, 579. - Biennial. Stem about 50 cm high, branched above, with leaves quite weakly arachnoid-hairy, smooth. Leaves pinnately incised with linear lobes to 3.5 mm wide; lobe smooth, along margin sharply sca- brous or scabrously serrate from short, acute, upright bristles. Capitula solitary at tips of leafy branches. Involucre subglobose, 15-16 mm long, and to 17 mm in dia; appendages of involucral bracts round (inner) to obreniform (middle and outer); outer appendages large, 6—7 mm long, 8—10 mm wide, with or lacking indistinct spot on dorsal side, spot well-developed in middle and inner appendages, oblong- triangular, ochreous or dark purple, from base extending upward along midrib, except middle thick narrow part; all appendages parchment- like scarious, thin, transparent, dorsally carinate, terminating in dis- tinct soft cusp in apical notch, with irregular small teeth; in involucre at flowering time appendages more or less appressed to bracts and entirely covering them, some outer appendages incurved. Corolla pale purple, 14-16 mm long in central bisexual florets. Achenes 3.5—5.0 mm long, dark brown, with whitish ribs; pappus 4.25—5.5 mm long, often distinctly longer than achenes, but sometimes even shorter. Flow- ering July to September. Sands.—European Part: Black Sea Region (on left bank of Dnieper from Dnepropetrovsk to Zaporozhye). Endemic. Described from Velikie Kuchugury in floodplain of Konka River. Type preserved in Kharkov; cotype available in Leningrad. Note. For the most part the range of this species is inundated. The species is preserved, however, on the island that has been formed and, possibly, is still above Dnepropetrovsk (in Dnepropetrovsk, there are collections from the vicinity of the villages of Kurilovka. 148. C. appendicata Klok. in Tr. Inst. Bot. KhDU, I (1936) 94, 101; Dobrocz. in Vizn. Rosl. URSR, 579. Biennial. Stems upright, 60-90 cm high, branched above middle with upward-spreading branches, with leaves distinctly arachnoid-hairy, 54 — 544 ribbed above, sharply scabrous along ribs from short bristles and tiny tubercles. Leaves mostly pinnately incised, with narrowly linear lobes 1—3 mm wide, sharply scabrous, especially along margin; lower leaves bipinnately incised, with somewhat broader terminal lobe; upper leaves linear, undivided, with white-scarious, aristate tip, subtending capitu- lum. Capitula solitary at tips of branches. Involucre globose, 22—27 mm in dia; involucral bracts yellowish, not entirely covered by ap- pendages; appendages more or less round, outer and middle ones round- rhombic, 10—12 mm long, 6.5—10.5 mm wide; inner.and adjacent middle ones dorsally with oblong-triangular, ochreous-purple or less often purple spot, transparent above spot, thin-scarious, yellowish with green- ish or ochreous tinge; outer appendages lacking distinct spot; all ap- pendages with irregular small teeth, dorsally with indistinct keel, ter- minating in soft short cusp in apical notch; appendages more or less conically convolute, divergent with acute tips. Corolla pale yellowish or creamish, larger than other species, to 20 mm in central bisexual florets and to 25 mm long in peripheral sterile florets. Achenes 4.5— 5.0 mm long, olive-brown with whitish ribs; pappus 5—7 mm long, distinctly surpassing achenes, white. Flowering July to August. Sands.—European Part: Black Sea Region (only on right bank sands of Dnieper, below Zaporozhye). Endemic. Described from “Lysaya Gora” [Bald Mountain] 7 km below Belenkoe wharf. Type preserved in Kharkov; cotypes in Leningrad and Kiev. Note. A very rare and unique species needing protection as a monument of nature. Series 4. Gerberianae Klok.—Appendages of involucral bracts smaller, with triangular, thicker, opaque part only at base, lacking dorsal keel and cusp; convex, margins not convolute at apex, mono- chromatic, lacking distinct dark basal spots. A very widely distributed series. 149. C. proto-gerberi Klok. in Tr. Inst. Bot. KhDU I (1936) 89, 102; Dobrocz. in Vizn. Rosl. URSR, 578. Biennial. Stem 30-90 cm high, branched from below middle or less often above middle, with upward-spreading branches, with leaves more or less densely arachnoid-hairy, sharply scabrous below capitula from conical tubercles like apical leaves. Cauline leaves pinnately or (lowermost) bipinnately incised, their terminal lobe narrowly linear or linear but somewhat broadened at apex, to 2 mm wide; apical leaves undivided, subtending, not surpassing capitulum. Involucre broadly ovate to subglobose, about 14-15 mm long; appendages of involucral bracts strongly convex, stiff, yellowish, oblong-elliptical to round- 545 triangular, with almost flat margins; largest appendages (in middle bracts) about 5 mm long and 5—6 mm wide, lacking conspicuous dark spots at base in upper third, indistinct spot sometimes only on append- age of inner bracts. Corolla pale pink to almost white or light creamish. Achenes 4—5 mm long, at maturity brownish or blackish, with pappus about 4—5 mm long. Flowering July to August. Open (not pine forest) sands.—European Part: Lower Don (along left bank of lower course of Donets River). Endemic. Described from Lugansk Region (village of Petropavlovka). Type in Kiev. Note. In all characters, this species is intermediate between C. donetzica Klok. and C. gerberi Stev., and can be distinguished with difficulty from them, although they differ quite sharply from each other. We have expressed the opinion (1936) that C. proto-gerberi Klok. is a progenitor species from which both C. donetzica Klok. and C. gerberi Stev. have originated. 150. C. donetzica Klok. in Tr. Inst. Bot. KhDU, I (1936) 86, 101; Dobrocz. in Vizn. Rosl. URSR, 578.—C. margaritacea auct. nonnull. fl. ucr. p. p. non Ten. Biennial. Stems solitary or many, 30-50 cm high, branched from below middle, with leaves more or less arachnoid-hairy above, espe- cially below capitula, scabrous, on ribs covered with acute tubercles. Leaves (except uppermost) pinnately incised with narrowly linear to almost filiform lobes 0.25—2.0 mm wide; basal and lower cauline leaves bipinnately incised, with lobes to 4 mm wide; all leaves stiff and acutely scabrous along margin, especially upper leaves; apical leaves subtending capitulum, undivided, narrowly linear, usually with white scarious tips. Capitula solitary at tips of branches; involucre at flow- ering time globose or somewhat oblong, 12—15 mm long, 10-16 mm in dia; appendages of involucral bracts elliptical, middle ones 5-6 mm long, 4-5 mm wide, inner ones much smaller; all basally with reddish- ochreous triangular spot along midrib, more conspicuous in inner bracts, surrounded by stiffer and thicker, less transparent, middle part in form of white triangle, very distinctly separated from transparent scarious border, whitish like central part, with characteristic micaceous luster; all appendages weakly convex, spatulate, not entirely covering bracts. Corolla pink, often pale, 14-19 mm long in central bisexual florets. Achenes 4—5 mm long, at maturity light brown, with pappus 4.0—5.5 mm long; pappus bristles thicker. Flowering June to August. Sands.—European Part: Lower Don (along Donets, from mouth of Oskol River to Lisichansk District on pine sands). Endemic. De- scribed from vicinity of village of Kremennoe at mouth of Krasnaya River. Type preserved in Kharkov; cotypes in Kiev and Leningrad. 549 546 151. C. pineticola [jin in Izv. Glavn. Bot. Sada SSSR, XXVI, I (1927) 34, 35; in Majevski, Fl. ed. 8, 617. Biennial. Stems solitary or 2, upright or basally assurgent, 25—65 cm high, branched from about middle, with upward-spreading branches, with leaves not densely arachnoid-hairy, smooth below, scabrous above along ribs from tiny conical tubercles. Basal and lowermost cauline leaves petiolate, about 5-10 cm long, pinnately and bipinnately in- cised, with linear or oblong, obtuse or subobtusely acuminate seg- ments or conical lobes, to 2—2.3 mm long and 0.7—4 mm wide, rather densely setose beneath and along margin; middle cauline leaves 3-9 cm long, sessile, pinnately incised with 3-8 pairs of lateral segments, scabrous-setulose beneath and along margin, segments linear and subobtusely acuminate to oblong-spatulate, 3-24 mm long, 1.0—3.5 mm wide; apical leaves subtending capitulum, undivided, linear or narrowly linear, about 10-15 mm long, cuspidate. Capitula solitary at tips of variously long branches, usually numerous. Involucre ovate or oblong ovate, 15-18 mm long, 10-14 mm in dia; appendages of in- volucral bracts weakly convex, more or less round to oblong-ovate (inner), lustrous, whitish, translucent with more or less distinct (espe- cially in appendages of outer bracts) ovate-triangular, dark purple, basal spot, somewhat toothed; lacking cusp, appendages of middle bracts about 4-6 mm long. Corolla 14-18 mm long, light purple. Achenes 3.5—4.7 mm long, whitish, more or less blackening at matu- rity with pappus 3.2—-5.0 mm long. Flowering June to August. Pine sands.—European Part: Volga-Don (known only from Khrenovsk pine forest in former Bobrovsk District of Voronezh Re- gion). Endemic. Described from above locality. Type in Leningrad. 152. C. breviceps Iljin in Izv. Glavn. Bot. Sada SSSR, XXVI, 1 (1927) 35; Klok. in Tr. Inst. Bot. KhDU, I, 90; Dobrocz. in Vizn. Rosl. URSR, 578.—C. leucolepis Ldb. Fl. Ross. II, 2 (1845) 689 p. min. p. non DC.—C. margaritacea auct. fl. ucr. p. p. non Ten. Biennial. Stem usually solitary, upright, 50-100 cm (and more) high, branched from middle or below, less often from above with more or less numerous divaricate branches, arachnoid-hairy with leaves, smooth below, more or less scabrous above along ribs. Basal and lower cauline leaves petiolate, to 15—20 cm (and more) long, pinnately and bipinnately incised with filiform narrowly linear or less often lin- ear lanceolate, obtuse segments; terminal lobe 5—8 mm long, 0.5—3.0 mm wide, smooth along margin and beneath like upper leaves (very rarely leaves rather densely setulose and with sharp tubercles.—f. asperula Klok. op. cit.); cauline leaves sessile, pinnately incised, mostly with almost filiform segments 0.5—1.0 mm wide; apical leaves sub- 550 547 tending capitula, filiform or narrowly linear, about 5-10 mm long, acuminate. Capitula solitary, at tips of more or less elongated branches. Involucre ovate, about 10-13 mm long and 6-10 mm in dia; append- ages of involucral bracts convex, elliptical, weakly lustrous, greenish- whitish or yellowish, transluscent, with small oblong triangular red- dish-brown-purple basal spot, scarcely toothed or entire lacking cusp; appendages of middle bracts 4-5 mm long and 3.5 mm wide. Corolla 10-14 mm long, pale purple. Achenes 3—4 mm long, ochreous or dark brown, with ribs whitish at maturity; pappus 2.5—-3.8 mm long, slightly shorter than achene. Flowering June to August. Open sands, sandy steppes.—European Part: Black Sea Region (only along left bank of Dnieper in lower-most course, on famous lower Dnieper sands). Endemic. Described from Aleshka (now Tsyurupinsk). Type in Leningrad. Note. It is the most characteristic representative of the endemic Lower Dnieper psammophilous flora. 153. C. paczoskii Kotov ex Klok. in Tr. Inst. Bot. KhDU, I (1936) 92, 101, descr.; Kotov in Dnevn. Vsesoyuzn. Sezda Bot. (1926) 101, 102, nomen seminudum; Dobrocz. in Vizn. Rosl. URSR, 578.—C. gerberi auct. non Stev.: Ijin in Izv. Glavn. Bot. Sada SSSR, XXVI, 1 (1927) 36, 39, p. min. p. quoad forman chersonicam. Biennial. Stems solitary or 2, upright or ascending at base, 30—50 cm high, with leaves more or less arachnoid-hairy, branched from above or below middle; branches upward spreading, more or less distinctly scabrous below capitula, along ribs from conical subobtuse or acute tubercles. Leaves (except uppermost) pinnately incised or (basal and lowermost cauline) bipinnately incised with narrowly linear to almost filiform, often involute lateral segments or terminal lobes, 0.3—1.5 mm wide; terminal lobe noticeably or considerably larger than lateral ones; sometimes to 4 mm wide; apical leaves subtending capitulum, undi- vided; narrowly linear, to 35 mm long, awned. Capitula solitary at tips of mostly short secondary branches. Involucre subglobose at flowering time, 15-18 mm long, 15-19 mm in dia; appendages of involucral bracts strongly convex, not covering bracts entirely, darkish yellow, oily lustrous, round, except inner elliptical, to 6 mm long, weakly toothed, lacking cusp; middle and inner appendages often (but not always) dichromatic, with oblong triangular, dark purple basal spot. Corolla about 16-17 mm long, light yellow. Achenes 3.5—5.0 mm long, whitish, not blackening at maturity; pappus 4.5—7.0 mm long, always considerably surpassing achene. Flowering June to August. Open sands, sandy steppes.—European Part: Black Sea Region (only on left bank sands of Ingulets River—a right tributary of Dnieper). 551 548 Endemic. Described from Ingulets River (vicinity of village of Novogrednevka). Type in Leningrad. 154. C. gerberi Stev. in Bull. Soc. Nat. Mosc. XXIX, 2 (1856) 391, in textu; Iljin in Izv. Glavn. Bot. Sada SSSR, XXVI, 1, 36, No. 5, excl. specimina chersonica; Klok. in Tr. Inst. Bot. KhDU, I, 90; Iljin in Majevski, Fl. ed. 8, 617. Biennial. Stems 1—3, upright or basally somewhat ascending, 30- 75 cm high, branched from above or below middle, with upward spread- ing branches, with leaves not densely arachnoid-hairy, smooth or al- most smooth above along ribs. Leaves (excluding uppermost) 2—11 cm long; basal (usually absent at flowering time) and lowermost cauline leaves petiolate, bipinnately incised or with incised segments, others sessile, pinnately incised; lateral segments-or terminal lobes narrowly linear to linear lanceclate, 3.5—30.0 mm long, 0.5—2.5 mm wide, acumi- nate; terminal lobe somewhat larger than laterals; apical leaves sub- tending capitulum, undivided, narrowly linear, about 10—15 mm long. Capitula solitary at tips of short or reduced branches, often partly aggregated. Involucre ovate or broadly ovoid to subglobose, 11-16 mm long, 9-14 mm in dia; appendages of involucral bracts strongly convex, not entirely covering bracts, more or less round or ovate (in- ner), with oily lustre, light yellow, lacking distinct, dark, basal, spots, weakly toothed, outer appendages indistinctly cuspidate, in middle bracts about 3—4 mm long. Corolla yellowish (creamish), about 12—15 mm long. Achenes 3.5—4.5 mm long, whitish, at maturity turning black; pappus 3-4 mm long. Flowering June to September. Open hummocky sands and sandy steppes.—European Part: Lower Don, Lower Volga, Trans-Volga (south); Volga-Kama (south). En- demic. Described from Volga. Type in Helsinki. Note. It is the most widespread species of the series and the entire section Pseudophalolepis Klok. 155. C. dubjanskii Iljin in Izv. Glavn. Bot. Sada SSSR, XXVI, I (1927) 36; in Majevski, Fl. ed. 8, 617; Klok. in Tr. Inst. Bot. KhDU, I, 86. Biennial. Stems solitary or few, upright or weakly ascending from base, with leaves not densely and [or] sparsely arachnoid-hairy, smooth along ribs, even below capitula. Leaves (except uppermost) 3.5—16.0 cm long; basal and lowermost cauline leaves petiolate, bipinnately incised, their lateral segments or terminal lobe mostly narrowly linear to linear-lanceolate, 5-25 mm long and 0.5—1.5 (3.0) mm wide, acumi- nate; terminal lobe slightly larger than laterals, to 40 mm long and 3 mm wide; apical leaves subtending capitulum, narrowly linear, about Spy 549 20-30 mm long and 1—1.5 mm wide, with whitish scarious tips. Ca- pitula solitary at tips of short branches. Involucre subglobose, often compressed above, wider than long, 15—27 mm long, 15-25 mm in dia; appendages of involucral bracts not strongly convex, entirely covering bracts, all more or less round, light yellow, with oily lustre, lacking dark basal spots, weakly toothed, lacking cusp; appendages of middle bracts about 6—7 mm long and 8—9 mm wide, of inner ones to 5 mm long and as wide. Corolla about 16-18 mm long, yellowish. Achenes 4.5—5.2 mm long, blackening at maturity; pappus 3.5—5.0 mm long. Flowering June to August. Hummocky sands.—European Part: Volga-Don (known from vi- cinity of villages of Bereznyak and Kazansaya, from sands along Khopyor River near Borisoglebsk). Endemic. Described from vicinity of village of Bereznyak of former Boguchar District. Type in Leningrad. Note. A very rare and relict species requiring special protection as a monument of nature. Section 2. Phalolepis.—Sect. Euphalolepis Dobrocz. in Bot. Zhurn. Akad. Nauk URSR, VI, 2 (1949) 64, 65, nomen.—Biennial or, less often, perennial plant; appendages of involucral bracts sharply differ- entiated into thick, completely opaque, central part with 5 veins tra- versing through it and completely transparent, scarious border; pappus always much shorter (most often half as long) than achene; stems more or less branched (not strongly in all species); corolla purple. Type of section: type of subgenus. An eastern Mediterranean section, represented in the USSR only in Crimea and the Caucasus by two endemic series. Series 1. Steriles Klok.—Relatively weakly branched perennials, less often biennials with fewer (to 10) capitula; all appendages of involucral bracts with distinct cusp. Endemic. Crimean mountain se- ries. 156. C. semijusta Juz. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIV (1951) 43. Perennial. Plants positively perennial with quite thick woody root. Stems to 10, ascending from prostrate base, 25-35 cm high, weakly branched, with 1-3 branches, 9-24 cm long, more or less densely arachnoid-tomentose and weakly scabrous or almost smooth below, sparsely arachnoid-hairy and scabrous-setulose above along ribs. Basal and lower cauline leaves 4-12 cm long, petiolate, bipinnately incised, arachnoid-hairy, like all other, leaves densely setulose, their terminal lobe oblong- or linear lanceolate, 7-15 mm long, 1.5—4.0 mm wide; 353) 550 middle and upper cauline leaves pinnately incised to undivided, with large, to 5-7 mm wide, terminal lobe, often almost lyrate. Capitula solitary at tips of long branches, but numerous on entire plant, larger than all other Crimean species of the section. Involucre broadly ovate to subglobose, 16-22 mm long and 12—21 mm in dia, lustrous from large scabrous appendages entirely covering involucral bracts; involu- cral bracts with small teeth, and soft cusp about 1-3 mm long; middle bracts with round appendages about 10 mm long; inner bracts with broadly ovate, smaller appendages; all appendages dorsally often with distinct oblong-lanceolate, light reddish-brown spot, central part of appendages membranous, weakly thickened but with 5 distinct veins. Corolla purple. Achenes oblong, narrowed toward base, 4—5 mm long, thinly hairy, blackening at maturity; pappus 1.5—2.0 mm long. Flower- ing July to August. Stony mountain slopes and on screes.—European Part: Crimea (known only from Chatyrdag Mountain). Endemic. Described from indicated locality. Type in Leningrad. 157. C. vankovii Klok. in Addenda, XXVII, 618.—C. nikitensis Illar. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVIII (1957) 305 p. p. quoad pl. jailicam. Perennial. Stems 2—10, ascending, about 30—35 cm high, more or less branched, more densely arachnoid-hairy, and smooth below, sharply scabrous above along ribs. Basal and lower cauline leaves 4-10 cm long, petiolate, pinnately incised, with undivided or partly 2—3-fid seg- ments, arachnoid-hairy, setulose beneath, like others scabrous-finely serrate; lateral lobes of leaves mostly linearly lanceolate, 9-25 mm long, 1.0-2.5 mm wide, cuspidate; terminal lobe somewhat larger than lateral ones; upper leaves at base 3-parted with shortened lateral lobes and large terminal lobe to 3 mm wide; apical leaves below capitulum smooth, linear, 5-15 mm long, 1 mm wide, almost smooth above, sharply scabrous beneath. Capitula solitary at tips of short branches. Involucre ovate or broadly ovate, 14-16 mm long, 10-14 mm in dia; appendages of involucral bracts round or broadly ovate with ovate triangular, dark reddish-brown central part and yellowish, scarious margin; margin toothed or almost fimbriate; appendages of middle bracts about 3—4 mm long, with shallow notch at apex and soft cusp 0.5—0.7 mm long, almost not exserted from it, with large dark reddish- brown spot below, extending beyond thick central part of appendage; appendage of outer bracts with more distinct cusp but in inner bracts lacking it; corolla light purple; achene oblong, 4.0—4.2 mm long, thinly hairy, with pappus 1.5—1.8 mm long, turning black at maturity. Flow- ering June to July. 554 551 Stony mountain slopes.—European Part: Crimea (mountains; known from Ai-Petri Mountain and “yaila” above it, Stavri-Kai, Uch- Kosh Gorge). Endemic. Described from Ai-Petri “yaila.” Type in Leningrad. Note. As indicated above, N.B. Illarionova referred some plants of this species to C. nikitensis Illar. Unfortunately, the type specimen of the latter is missing in Leningrad. There are topotypes from the vicinity of the Uchan-su-Waterfall, which we refer to C. sterilis Stev. They conform to the characters given in the authentic description, according to which the involucre in C. nikitensis Illar. is ovate, 12—13 mm long and 9-12 mm in dia, while the achenes are 3—3.5 mm long. Such measurements are entirely characteristic of C. sterilis Stev., but the alpine race replacing the latter is characterized by larger measure- ments. Moreover, C. vankovii Klok. is a perennial plant, whereas C. nikitensis Illar. has been described as biennial. 158. C. sterilis Stev. in Bull. Soc. Nat. Mosc. XXIX 2 (1856) 390; Boiss. Fl. or. III, 622; Schmalh. FI. II, 121; Fedtsch. and Fler. FI. Evrop. Ross. 1019.—C. alba MB. FI. taur.-cauc. II (1808) 353, non L.—C. leucolepis Ldb. FI. Ross. II, 2 (1845) 689, p. p. non DC.—C. stankovii Ullar. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVIII (1957) 305.—C. nikitensis Illar. ibid. 304, p. max. p.—Exs.: Dorfler. Herb. norm. No. 4152. Biennial, perennial. Plant biennial or short-lived (flowering in sec- ond year but also in third year). Stems upright, ascending, branched from base or sometime (under unfavorable conditions) procumbent, solitary or few (2-3), 10-95 cm high, more or less branched, angular, with leaves more or less distinctly arachnoid-hairy, acutely scabrous along ribs from erect, small, thickish bristles and only at base almost smooth. Basal and lower cauline leaves about 6-12 mm long, with distinctly developed or relatively long petiole and pinnately incised lamina, individual (middle) segments 2—3-fid; segments of terminal lobes of leaves linear-lanceolate, 5-32 mm long, 1—3 mm wide, cus- pidate; petiole and leaf lobes arachnoid pubescent and rather densely covered with small, thickish bristles about 0.3—1.1 mm long; middle and upper leaves sessile, weakly lobed to undivided, scabrous, finely setose along margin and beneath; uppermost leaves narrowly linear, about 7-13 mm long and 1.0—1.5 mm wide, terminating in short, thin cusp. Capitula few (2-10 per stem, less often more), at tips of long primary or short second order branches, with florets about 16-18 mm long. Involucre ovate to oblong-ovate, 10-14 mm long, 6-10 mm in dia, weakly arachnoid-hairy; involucral bracts coriaceous, with 5 veins; usually greenish; appendages of involucral bracts with thick, coria- 552 ceous, central part and sharply demarcated, thin-scarious, and trans- parent border, more or less toothed, dorsally carinate, continued into thin, 0.4-2.0 mm long cusp; appendages of middle bracts round, 3—6 mm long, with more or less narrow, yellowish or often reddish-brown, triangular central part and whitish or often yellowish border; append- ages of inner bracts ovate, smaller, usually with reddish-brown spot in central part. Corolla light purple. Achenes roundish-cuneate, 2.5—3.5 mm long, whitish or light reddish-brown, scatteredly thin-hairy; pap- pus 0.4—2.0 mm long, consisting of unequal bristles, sometimes almost undeveloped. Flowering June to August. Stony slopes.—European Part: Crimea (common in foothill part and on southern coast). Endemic. Described from Crimea (probably from vicinity of Simferopol). Type in Helsinki. Note. A highly polymorphic species, it hybridizes readily with C. diffusa Lam., C. ovina Pall. s. 1. and other species of the subgenus Acrolophus (Cass.) Dobrocz. These hybrids include the ones described by N.B. Illarionova as separate species—C. Jongiaristata Illar. and C. iljiniana Il\lar. (in the article “Novye vidy roda vasilek flory Kryma” [New species of centaureas in the flora of Crimea], included in the cited volume of Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, 306, 307. It is well known that centaurea hybrids break down and do not produce stable, widely distributed forms; hence these should not be described as hybridogenic species. The difference between the southern coastal populations of this species and the ones from the foothills, to the extent that we could ascertain, is not expressed in systematic charac- ters. Hence, treating these populations as two different species—C. stankovii Illar. (described from the Nikitsky Botanical Garden) and C. nikitensis Ilar. (described from the vicinity of Yalta, near the Uchan- Su Waterfalls)—is useless. All of them, to all appearances, form a single geographic race. The plants collected by S. Grigorev along the Shtangev track near Yalta differ more sharply from the type specimen of C. sterilis Stev. by having the leaves strongly divided and the pappus of the achenes up to 4 mm long. The plants collected by S. Dzevanovsky from the Esk stone quarry which are distinguished by completely black achenes also are distinctive. However, the material on these forms is too limited for any final judgment. Plants from the Ai-Petri “Yaila” clearly differ from C. sterilis Stev.; they were partly included by N.B. Illarionova along with Steven’s species in the composition of her C. nikitensis Illar. They can be considered as an alpine race, altitudinally replacing C. sterilis Stev., and are described above as a separate species. The unique form described by S.V. Juzepczuk from the Baider “yaila,” under the name C. ninae Juz. in Bot. Mat. Gerb. Bot. Inst. 555 553 Akad. Nauk SSSR, XIV (1951) 42, apparently is of hybrid origin (C. sterilis Stev. x C. maculosa s. 1.). Series 2. Transcaucasicae Klok.—Biennial or less often perennial plants with strongly branched stem and numerous capitula; middle and inner involucral bracts lacking cusp. A Caucasian (Transcaucasian) series extending into the eastern part of Crimea. Herbarium material of this series is still very insufficient, and the species composition of the series is far from being worked out. 159. C. sarandinakiae Illar. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVIII (1957) 303. Perennial. Root woody, to 3 cm thick. Stems usually many, as- cending, 30-50 cm high, branched with quite many, somewhat proc- umbent branches, ribbed, with leaves not densely pubescent, smooth below, scabrous or almost smooth above along ribs. Basal and lower cauline leaves about 6-13 cm long, petiolate, pinnately incised, with linearly or oblong-lanceolate segments to 40 cm long and 6 mm wide, finely setose; middle and upper leaves pinnatifid, with smaller lateral lobes and larger terminal lobe, sessile, sharply scabrous or almost smooth above, scabrous-finely serrate along margin and beneath along midrib; apical leaves below capitulum shorter than involucre, undi- vided, oblong-lanceolate, cuspidate. Capitula solitary at tips of short, second order branches, about 16-20 mm long with florets. Involucre broadly ovate, 12-17 mm long, 10-16 mm in dia, yellowish and lus- trous from appendages of involucral bracts entirely covering them; appendages of outer bracts ovate lanceolate, about 4 mm long, with soft cusp to 2 mm long; appendages of middle bracts round, about 5— 6 mm long, with small, light yellow triangular, thick part at base and below middle, to 2 mm long, lacking keel and cusp above, with tiny teeth like outer; appendages of inner bracts almost identical. Corolla pale purple. Achenes oblong, narrowed at base, 3.5—4.2 mm long, finely hairy, blackening at maturity; pappus 1.2—2.0 mm long. Flower- ing July to August. Stony talus and rocks.—European Part: Crimea (known only from Karadag in eastern Crimea). Endemic. Described from Karadag, from Karagach Mountain. Type in Leningrad. Note. The lone specimen collected by I.I. Sprygin at the Ai-Petri foothills near Miskhor is very distinctive and similar to this species in the structure of the appendages. Most of its leaves are bipinnately incised with long narrow lobes; the appendages of its involucral bracts have distinct light reddish-brown spots; the achenes are blackish and 556 554 4.5—-4.7 mm long, with a pappus about 2.5 mm long. Apparently it flowers earlier than other species. Possibly it is an introduced plant. C. sarandinakiae Il\lar. shows affinity not to the Crimean but to the Caucasian members of the section. 160. C. novorossica Klok. in Addenda XXVII, 619.—C. sterilis auct. non Stev.: Grossh. Fl. Kavk. IV (1934) 211; Opred. Rast. Kavk. 496 and auct. al. fl. cauc. Biennial. Stems 1—3, upright or ascending from base, 20—70 cm (and more) high, branched, angular, with leaves relatively densely floc- cose-arachnoid-hairy, almost smooth at base, acutely scabrous above along ribs. Basal and lower cauline leaves petiolate, bipinnately in- cised, with linear or narrowly linear terminal lobe, often withering before flowering time; middle cauline leaves about 2—6 mm long, sessile, pinnately incised, like other leaves scabrous-finely setose be- neath and along margin, weakly scabrous to almost smooth above, with narrowly linear or narrowly linear-lanceolate lateral lobes up to 5 pairs, 5-25 mm long and 0.5—1.5 mm wide; terminal lobe larger, to 32 mm long and 2.5—-3.0 mm wide; upper leaves smaller, pinnatifid, apical leaves below capitulum undivided or with basal tooth, linear, 7— 12 mm long, 1.2—1.5 mm wide, cuspidate. Capitula solitary at tips of branches, (1—-3)7—18 per stem, 17-20 mm long. Involucre ovate or broadly ovate, 12-15 mm long, 8-12 mm in dia, whitish, rather dull, weakly lustrous; appendages of outer involucral bracts round-rhombic, like others finely toothed, with soft cusp; to 1.5 mm long; appendages of middle bracts round, about 3.5—-4.0 mm long, somewhat larger than others, with fimbriate margin (even in budding stage), lacking cusp, with reddish-brown or light reddish-brown triangular spot at base of central part not reaching apex; appendages of inner bracts oval, also lacking cusp, with spots. Corolla light-purple. Achenes oblong-cuneate, about 4—4.5 mm long, finely hairy, blackening at maturity; pappus 1.5-3.3 mm long. Flowering June to August. Stony slopes, as roadside weed.—Caucasus: Western Transcaucasia (north, known from vicinity of Taman, Novorossiisk, Gelendzhik and Markotkh Range). Endemic. Described from vicinity of Novorossiisk. Type in Leningrad. Note. This Novorossiisk species clearly differs from C. sterilis Stev. by the absence of the cusp on the appendages of the middle and upper involucral bracts and by the longer, blackened achenes with a well-developed pappus. It apparently approaches the eastern Crimean species C. sarandinakiae Ilar., from which it also is well distinguished. The specimens collected by T. Pojarkova from the Markotkh Range differ from all the others by having a lower stature (20-22 cm), fewer Si 3150) (1-3) capitula on the stem, and somewhat more divided leaves. In these specimens, the involucre is more oblong. 161. C. georgica Klok. in Addenda XXVII, 619.—C. pseudodeusta auct. fl. cauc. p. p. non Hayek. Biennial. Stems solitary or few, upright, 40—70 cm high, branched from middle or less often above, with more or less procumbent, vir- gate branches, with leaves not densely arachnoid-hairy, smooth below, sharply scabrous above along ribs. Basal and lower cauline leaves petiolate, bipinnately incised, with linear or linearly lanceolate, nar- row, terminal lobe, mostly withering by flowering time; middle cauline leaves 1.5—4.0 cm long, sessile, pinnately incised or some with pin- natifid segments, sharply scabrous along margin and beneath from dense spinules, like upper leaves weakly scabrous above; segments or termi- nal lobes linear or linear-lanceolate, to + 10 mm long, 0.5—1.5 mm wide, terminal lobe larger, up to 2-3 mm wide; apical leaves undi- vided, linear or narrowly linear, cuspidate. Capitula solitary at tips of long primary branches or short second-order branches, (3)12—30 per stem, 17-19 mm long. Involucre ovate to broadly ovate, about 12—13 mm long and 8-9 mm in dia; yellowish to light reddish-brown or reddish brown to light yellow, lustrous; appendages with yellowish- reddish-brown thick central part and yellowish scarious border, finely toothed to almost fimbriate (lower); appendages of outer bracts broadly ovate, with distinct apical notch having cusp to 1 mm long; append- ages of middle bracts round, about 3—4 mm long lacking notch and cusp, of inner bracts oblong-elliptical, smaller, lacking cusp. Corolla light purple. Achenes about 3.7 mm long, brownish-olive when ma- ture; pappus about 1.5 mm long. Flowering June to July. Stony slopes, in low-montane zone.—Caucasus: Western Trans- caucasia (known from vicinity of Borzhomi and Gori). Endemic. Described from Borzhomi. Type in Leningrad. 162. C. latiloba Klok. in Addenda, XXVII, 620. Biennial. Stems solitary or few, upright, 20-70 cm high, divaricately branched, with leaves somewhat densely or sparsely arachnoid-hairy, smooth below, sharply scabrous above along ribs from spinules. Basal and lower cauline leaves petiolate, to 8-12 cm long, pinnately incised with ovate, pinnatifid segments, finely setose beneath; terminal lobe oblong-ovate or less often oblong-obovate, 2-15 mm long, 1.5—5.0 mm wide; middle cauline leaves sessile, pinnately incised, 2—4 cm long, sharply setose beneath like upper leaves, finely setose-serrate on margin, with 1-3 pairs of lateral segments oblong, 3—20 mm long and 1-4 mm wide, toward apex usually somewhat broadened and abruptly 558 556 attenuate into subobtuse tip, middle lobe noticeably distinctly larger than laterals, to 5 mm wide; apical leaves undivided, oblong, 8-25 mm long, 2-6 mm wide, abruptly attenuate into subobtuse tip. Capitula solitary at tips of branches, 7-40 per stem, 16-21 mm long. Involucre broadly ovate to subglobose, 11-14 mm long, 10-15 mm in dia, yel- lowish or reddish-brown-yellow, lustrous; appendages with light yel- low or light reddish-brown, thick, central part and yellowish or yel- lowish-whitish, scarious border, weakly toothed in apical part or subentire; appendages of outer bracts broadly ovate, emarginate with small inconspicuous cusp in notch; appendages of middle bracts about 5—6 mm long, inner ones elliptical, smaller. Corolla light purple. Achenes about 2.5 mm long, olive-brown; pappus about 1 mm long. Flowering June to July. Mountain slopes, meadows, as weed in fields.—Caucasus: Eastern and Southern Transcaucasia. Endemic. Described from Transcaucasia. Type in Leningrad. Note. We could not identify this species with C. schemachensis Sosn. ex Grossh. to which it approaches by its wide leaf lobes C. schemachensis Sosn. ex Grossh. has been mentioned only in Opredelitel Rastenii Kavkaza [Manual of the Plants of the Caucasus] by A.A. Grossheim, and, according to him, differs from other similar Cauca- sian species by having very large involucres of the capitula, reaching about 20 mm in diameter, which does not agree with our specimens. Specimens identified by D.I. Sosnowsky or A.A. Grossheim could not be found in Leningrad. In the size of the involucres, our specimens do not differ from C. transcaucasia Sosn. ex Grossh. and C. sterilis auct. fl. cauc. (C. novorossika Klok.) referred by A.A. Grossheim to the group of species with smaller, 10-12 mm-wide, involucres. For similar reasons, we also still do not understand C. daghestanica Sosn. ex Grossh. (Opred. Rast. Kavk. (1949) 496, absque diagn.). 163. C. transcaucasica Sosn. ex Grossh. Fl. Kavk. IV (1934) 212; Opred. Rast. Kavk. 496. Biennial. Stems 1—4, upright or ascending at base, 40-70 cm (and more) high, branched from middle or slightly below, with upward- spreading branches, less often divaricate, with leaves densely or sparsely arachnoid-hairy, smooth below, sharply scabrous above along ribs from spinules or tubercles. Basal and lower cauline leaves petiolate, about 4-7 cm long, pinnately incised, partly bifid segments, spinulose be- neath and along margin; leaf segments or lobes linear-lanceolate, about 1.5-2.0 mm wide; middle cauline leaves sessile, pinnately incised, about 3—5 cm long, mostly with 3 pairs of lateral lobes, sharply sca- brous or scabrous-serrate on margin, not setose beneath, quite weakly 557 559 scabrous, almost smooth above; their lateral lobes linearly lanceolate, about 7-15 mm long, 1.0—2.5 mm wide, middle (terminal) lobe to 22— 32 mm long and 2.5—3.0 mm wide; apical leaves undivided, linearly lanceolate, 7-22 mm long, 1.2—3.2 mm wide, cuspidate. Capitula solli- tary at tips of branches, 10-20 on stem, 17-19 mm long. Involucre short-ovate, more or less compressed, 12—13 mm long and 11-13 mm in dia, yellowish, lustrous; appendages of outer involucral bracts round- rhombic, emarginate, with or lacking small inconspicuous cusp in notch; appendages of middle bracts broadly obovate, 4-6 mm long, entire or weakly toothed, lacking cusp; central thick part of appendage ovately triangular, yellowish or brownish-light-yellow, with inconspicuous, light reddish-brown basal spot or lacking it; appendages of inner bracts oblong-ovate, quite small, lacking cusp. Corolla light purple. Achenes 2.53.1 mm long, brownish-olive at maturity, with pappus 0.7—-1.0 mm long. Flowering June to July. Mountain slopes.—Caucasus: Eastern Transcaucasia. Endemic. Described from Azerbaidzhan. Type in Leningrad. Note. The incomplete specimens collected by F. Alekseenko in the vicinity of the village of Khazra on the Samur River are distin- guished by a longer pappus, as long as (3 mm) the achenes. 164. C. alexandri Bordz. in Fedde, Repert. XXXVI (1934) 306; in Zhurn. Inst. Bot. VUAN, II (1934) 80; Grossh Opred. Rast. Kavk. 495. Biennial, less often perennial. Stems 1—3, upright, 60-90 cm high, branched from middle or below with upward-spreading branches, more or less densely or sparsely arachnoid-hairy, with leaves smooth below, more or less scabrous above from conical tubercles (and not spines). Basal and lowermost cauline leaves petiolate, about 10-15 cm long, bipinnately incised or with more or less lobed segments, lacking spines or bristles beneath, almost smooth or weakly scabrous, noticeably scabrous along margin from conical tubercles; their terminal lobes linear to linearly- or oblong-lanceolate, cuspidate, 0.5—2.5 mm wide; middle cauline leaves sessile, 1.5-7.5 cm long, bipinnately to pin- nately incised, more or less scabrous along margin and beneath; their lobes narrowly linear to linear-lanceolate, lateral lobes to 22 mm long and 1.5 mm wide, middle (terminal) lobes to 32 mm long and 2 mm wide; apical leaves linearly lanceolate, smaller, cuspidate. Capitula solitary at tips of variously long branches, sometimes on very short branches, more or less numerous, 12-14 mm long. Involucre ovate to oblong-ovate, 9-10 mm long and 4—6 mm in dia, yellowish or whitish- yellowish; appendages of outer involucral bracts broadly ovate, with indistinct notch at apex and very small cusp, more or less toothed; 560 561 558 appendages of middle bracts round, about 2.5—3.0 mm long, weakly toothed to entire, lacking cusp, central thicker part of appendage ovately triangular, light-yellow or light reddish-brown, with scarcely visible darkening at base (as in outer appendages), of inner bracts ovate, smaller, lacking cusp. Corolla light-purple. Achenes oblong, basally narrowed to oblong-obovate, 3.0—3.3 mm long, brownish-olive at ma- turity, weakly hairy with pappus 1.5—2.5 mm long. Flowering June to July. Slopes and rocks, in lower montane zone.—Caucasus: Eastern and Southern Transcaucasia. Endemic. Described from Armenia (Megri- Artatsur in gorge of Araks River). Type in Yerevan. Note. Simultaneously with the species, E.I. Bordzilowsky described C. alexandri var. rhodolepis Bordz., differing from the type by the dirty-ochreous margins of the appendages of the involucral bracts. Subgenus 17. Microlophus (Cass.) Hayek, Centaurea-Art. Osterr. Ung. (1901) 2.—Microlophus Cass. in Dict. Sc. nat. XLIV (1826) 37 and LIV (1829) 490.—Piptoceras Cass. ibid. L. (1827) 469 and LIV (1829) 487.—Centaurea sect. Microlophus (Cass.) DC. Prodr. VI (1837) 567.—Centaurium sect. Microlophus (Cass.) C. Koch in Linnaea, XXIV (1851) 418.—Capitula medium-sized, with many flo- rets, rather numerous, solitary at tips of main stem and its lateral branches, often aggregated in lax corymbose or paniculate inflores- cence. Involucre ovate, glabrous or subglabrous (with thin glabrescent arachnoid tomentum), 8—20 mm in dia; involucral bracts coriaceous, brownish-greenish lacking or with inconspicuous veins, gradually en- larged from broadly triangular-ovate outer to oblong-lanceolate inner- most, acuminate or with short spine, lacking appendages (in species of our flora), less often with small incurved, lanceolate appendage, not decurrent on bract, pectinate-ciliate and terminating in spine or prickle. All florets tubular, heterogeneous; central florets bisexual, peripheral ones not large (usually even smaller), sterile, lacking or with 1—4 staminodes, included in corolla tube; stamen filaments flattened on sides and mostly covered on two sides with crisped papilliform hairs; stigma short-bifid. Achenes about 4—5 mm long, initially sparsely short- hairy; mature achenes glabrous or subglabrous; pappus as long as achene or longer, persistent, double, outer pappus consisting of several irregu- lar rows of bristles scabrous on two sides, gradually elongated from very short outer to middle and inner, 1/6—1/3 as long as outer, consist- ing of single row of slightly broader, erose-ciliate bristles scarious in upper part. Perennial or biennial herbs lacking underground shoots, with string-like root and upright, strongly branched (often almost from base) stems. Basal rosette and lower cauline leaves usu- 359 ally more or less pinnately lobed, others entire or more or less toothed, usually decurrent on stem, short-scabrous with or lacking inconspicu- ous punctate glands. Type of subgenus: C. behen L. Of the 8-10 species of this subgenus, ‘distributed mostly in the countries of West Asia and in Iran, three are found in the USSR. Many authors of the Caucasus flora recognize this subgenus as the separate genus Microlophus Cass. In fact, the species belonging here occupy a very distinctive position among the other species of Centau- rea L. in the USSR flora. However, in West Asia there is a whole range of species occupying a more or less intermediate position be- tween the subgenus Microlophus and the other subgenera of this ex- tensive polytypic genus. 1. Capitula aggregated in corymbose inflorescence; apical leaves close to base of capitula and often enveloping them; leaves of.basal rosette pinnately lobed, with numerous lateral lobes and terminal lobe as, wide as lateraltones! Mey Awe RB, SR Dt + Capitula usually not aggregated in corymbose inflorescence; apical leaves not close to base of capitula; leaves of basal rosettes undi- vided or pinnately lobed, but then with strongly enlarged terminal lobe and few lateral lobes ..............:ceccesceesseeseeee 163. C. behen L. 2. Involucres 13-17 mm in dia; subtending apical leaves quite large and wide, often entirely enclosing capitula ..............cecceeseesceeteeseeeeeees ART RE VINA) SPRY HN RY oe 166. C. szovitsiana Boiss. + Involucres 9-13 mm in dia; apical leaves smaller and narrower, usually somewhat removed from base of capitula ................... Mei hb) yee WRAL, OTS Te ARN 167. C. polypodiifolia Boiss. Section 1. Microlophus.—Centaurea sect. Microlophus (Cass.) DC. Prodr. VI (1837) 567 p. p.—Capitula not in corymbose inflores- cence; apical leaves not close to base of capitula. Perennial polycarpic plants, covered with very short papilliform hairs; basal and lower cauline leaves undivided or pinnately lobed but then terminal lobe larger than lateral lobes. Type of section: type of subgenus. 165. C. behen L. Sp. pl. (1753) 914; Boiss. Fl. or. III, 682 p. p.; Grossh. Fl. Kavk. IV, 210.—C. alata Lam. Encycl. meth. I (1783) 665; MB. FI. taur.-cauc. II, 342 and III, 586; DC. Prodr. VI, 567; Ldb. Fl. Ross. II, 2, 687.—C. acuta Vahl. ex MB. Beschreib. Land. Casp. Meer. (1800) 82 and App. 211, non Lam. (1789).—C. babylonica auct. non L.: MB. op. cit. (1800) 82 and App. 211.—C. bordzilowskii 562 560 Lonacz. in Tr. Yurev. Bot. Sada, IX (1908) 162.—Piptoceras behen (L.) Cass. in Dict. Sc. nat. LIV (1829) 488.—Microlophus alatus (Lam.) Cass. ibid. 491.—M. behen (L.) Takht. in Takht. and Fedtsch. FI. Erevana (1945) 326; Grossh. Opred. Rast. Kavk. 494.—Centaurium behen (L.) C. Koch in Linnaea, XXIV (1851) 418.—Ic.: Lonacz. op. cit. 163; Takht. and Fedtsch. Fl. Yerevana, Atlas (1946) Plate 117. Perennial. Plants to 1.5 m high, covered with very short, papilli- form and spiniform hairs mixed with weakly floccose arachnoid-hairs, weakly scabrous; sometimes subglabrous. Stems upright, branched above middle, with numerous capitula usually at different height. Basal and lower cauline leaves very large, with quite long petioles, oblong, undivided or lyrate, with terminal lobe much larger than fewer lateral lobes, irregularly toothed; middle and upper cauline leaves smaller, entire or very weakly toothed, sessile, base of lamina decurrent on stem as broad wings; apical leaves not close to base of capitulum. Involucre 13-17 mm in dia, 23—27 mm long; involucral bracts acute, lacking appendages. Florets yellow. Achenes about 5 mm long, with pappus 6—7 mm long. Flowering July to August. Stony slopes, meadows, gravel beds, roadsides, sometimes in fields; up to mid-montane zone.—Caucasus: Eastern and Southern Trans- caucasia; Soviet Central Asia: mountainous Turkmenia. General dis- tribution: West Asia, Armenia and Kurdistan, Iran. Described from West Asia (Lebanon?). Type in London. Note. Not having the type material for C. behen L. and C. alata Lam., I am not at all sure about its complete identify as well as its affinity with the species described by me. Possibly, one of these names refers to one of the two following species. Section 2. Polypodiifoliae Tzvel. in addenda XXVII, 620.—Ca- pitula aggregated in more or less corymbose inflorescence; apical leaves usually more or less close to base of capitula. Biennial or short-lived(?) monocarpic plant, covered with short papilliform hairs and longer crisped hairs; basal and lower cauline leaves pinnately lobed, their terminal lobe almost of same size as lateral lobes. Type of section: C. polypodiifolia Boiss. 166. C. szovitsiana Boiss. Fl. or. III (1875) 683; Grossh. Fl. Kavk. IV, 210 p. p.; Tzvelev in Bot. Mat. Gerb. Bot. Inst. Akad.. Nauk SSSR, XX (1960) 29.—C. behen auct. non L.: DC. Prodr. VI (1837) 567; Ldb. FI. Ross. II, 2, 687.—Microlophus szovitsianus (Boiss.) Takht. in Takht. and Fedtsch. Fl. Yerevana (1945) 326 p. p.; Grossh. Opred. Rast. Kavk. 495 p. p. 563 561 Biennial or perennial. Plants to 1 m high, covered with short spiniform and papilliform hairs (mostly along leaf margin) and longer but sparse crisped hairs mixed with some arachnoid hairs, scabrous. Stem upright, strongly branched above middle, with numerous capitula in corymbose inflorescence. Basal and lower cauline leaves quite large, petiolate, pinnately lobed (almost to midrib), with 6-10 oblong, ob- tuse, lateral lobes and terminal lobe almost as large as lateral lobes; middle and upper cauline leaves numerous, entire or weakly toothed, sessile, base of lamina decurrent on stem as broad wings; apical leaves enveloping capitulum rather large; all leaves distinctly punctately glandular with glands lighter than in preceding species, often with yellowish tinge. Involucre 13-17 mm in dia, 20—25 mm long; involu- cral bracts acute, lacking appendages. Florets yellow. Achenes un- known. Flowering July to August. Stony slopes, mountain meadows, often in fields, as weed; mostly in mid-montane zone.—Caucasus: Southern Transcaucasia (northeast- ern part of Nakhichevan ASSR). General distribution: Armenia and Kurdistan(?). Described from Nakhichevan ASSR (Kashadara Moun- tain). Type in Geneva; isotype in Leningrad. 167. C. polypodiifolia Boiss. Diagn. pl. or. ser. I, 6 (1845) 126 and FI. or. III, 683; Tzvelev in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XX (1960) 29.—C. szovitsiana auct. non Boiss.: Grossh. FI. Kavk. IV (1934) 210 p. p.—Microlophus szovitsianus (Boiss.) Takht. in Takht. and Fedtsch. Fl. Yerevana (1945) 326 p. p.; Grossh. Opred. Rast. Kavk. 495 p. p.—M. polypodiifolius (Boiss.) Agadsh. in Fl. Azerb. VIII (1961) 443.—Exs.: Fl. or. exs. No. 24 (sub C. szovitsiana). Biennial or perennial. Plants to 70—80 cm high, short- and sparsely crisped hairy-mixed with some arachnoid hairs, scabrous from spiniform hairs mostly along leaf margin. Stem upright, strongly branched above middle or almost from base, with numerous capitula in corymbose inflorescence. Leaves as in preceding species, somewhat attenuate at apex; apical leaves smaller, at flowering time usually somewhat distant from base of capitulum. Involucre 9-13 mm in dia and 16-20 mm long; involucral bracts acute, lacking appendages. Florets yellow. Achenes about 4 mm long, with pappus about 5 mm long. Flowering July to August. (Plate XXVI). Stony slopes, mountain meadows and meadow-steppe areas, road- sides; mostly in mid-montane zone.—Caucasus: Southern Transcaucasia (western part including area of Lake Sevan). General distribution: Asia Minor (eastern part), Armenia and Kurdistan, Iran (western part), Iraq. Described from Iraq. Type in Vienna. 564 562 Note. In relation to the preceding species it is a more western ecogeographical race, widely distributed in Armenia. A hybrid of this species with C. stevenii MB. is known from the environs of Leninakan (collections of N.N. Tzvelev and S.K. Czerepanov, 1960). Subgenus 18. Seridia (Juss.) Czer. comb. nova.—Seridia Juss. Gen. pl. (1789) 173; Cass. in Dict. Sc. nat. XLVIII (1827) 498.— Capitula medium-sized, with many florets, solitary at tips of stems and their lateral branches. Involucre ovate, 10—15 mm in dia, weakly arach- noid-hairy; involucral bracts coriaceous, greenish (yellowing on dry- ing), with inconspicuous veins in upper part, gradually elongated from broadly ovate outer to oblongly lanceolate inner ones; appendages of outer and middle bracts small, not decurrent, coriaceous, pale reddish- brown, with palmately divergent short spines, of which middle spine usually slightly longer and very slightly thicker than others; append- ages ef innermost bracts very small, membranous, unevenly toothed. All florets tubular, yellow, 1.5 times as long as involucre, heteroge- neous; central florets bisexual, peripheral sterile, lacking staminodes and rudiments of style, not enlarged, with 4(5)-fid corolla; stamen filaments flattened, covered with papilliform hairs; style shortly bifid. Achenes oblong or oblong ellipsoidal, 4.5-5.0 mm long, scatteredly hairy; pappus 1/2—2/3 as long as achenes, whitish or weakly ochreous, persistent, double; outer pappus consisting of several irregular [rows of] weakly serrate, scabrous bristles, gradually elongating from outer to inner; inner pappus of single row of bristles, similar to outer pappus bristles but slightly shorter, sometimes pappus almost simple. Peren- nial herbs with long, thick root, fibrous, branched above, with branched rhizome, and prostrate, at ends of ascending stems; leaves entire, lyrately lobed or divided, petiolate. (Description of the subgenus has been compiled, as is conventional in our Flora, only with reference to our species.) Type of subgenus: C. seridia L. Pending clarification of the true systematic position of C. stevenii MB. in the genus Centaurea L., I am referring it provisionally to the subgenus Seridia (Juss.) Czer. Probably the yellow-flowered perennial species C. cheirolopha (Fenzl) Wagenitz, C. lancifolia Sieb., C. hololeuca Boiss., and C. stevenii MB. should be split off in a separate section or even subgenus. 168. C. stevenii MB. FI. taur.-cauc. H (1808) 356; Ldb. FI. Ross. II, 2, 710; Grossh. Opred. Rast. Kavk. 495.—Phaeopappus stevenii (MB.) Boiss. Fl. or. III (1875) 603; Grossh. Fl. Kavk. IV, 206.—Exs:.: PiSvorexst) Now323; 565 563 Perennial. Root long, thick, vertical or oblique, fibrous, branched above, usually with short-branched rhizome. Stems usually prostrate, at end ascending, 10—40 cm long, with few short branches, sometimes simple, grayish-green or more or less grayish from fine lax arachnoid- hairs often mixed with articulate, somewhat stiff, short hairs. Leaves lanceolate or oblanceolate, acute, subacute, less often subobtuse, peti- olate (upper cauline leaves subsessile), entire, lyrately lobed or parted with oblong, obtuse, entire, small lateral lobes and oblong-elliptical or ovately rhombic larger terminal lobe, with articulate, stiff hairs mixed with few arachnoid hairs, covered with pale, sessile, punctate glands. Capitula solitary, at tips of stems and lateral branches. Involucre ovate, 15-20 mm long and 10-15 mm in dia; involucral bracts from broadly ovate outer to oblong-lanceolate inner; appendages of outer and middle bracts coriaceous, pale reddish-brown, with (3)5—7 palmate, almost equal, small spines; appendages of innermost bracts membranous, unevenly toothed. Florets yellow. Achenes 4.5—5.0 mm long and 2.0— 2.5 mm wide; pappus 2.5—4.0 mm long. Flowering June to July. Herb slopes, mid-montane zone.—Caucasus: Eastern and South- ern Transcaucasia. General distribution: Armenia and Kurdistan (former Kara Region). Described from Georgia. Type in Leningrad. Note. One specimen of this species is preserved in the Herbarium of the Botanical Institute of the Academy of Sciences of the USSR in Leningrad with the label “Nowotscherkask Pabo.” The lack of new collections from the area of Novocherkassk and generally from Ciscaucasia leads me to assume that in the present case a label mixup has occurred. The report of the occurrence of C. stevenii MB. in Dagestan is based on Lagov’s specimens from Derbent. It also is not confirmed by new collections. The hybrid C. polypodiifolia Boiss. x C. stevenii MB. is known from the vicinity of Leninakan (Armenian SSR, Achinsk District between Ani and Bagravan stations, on the border of a field, 2-3 km from Bagravan Station, 5.VII.1960, No. 847, N. Tzvelev and S. Czerepanoy). Subgenus 19. Rhizocalathium (Tzvel.) Tzvel. comb. nova.—Cen- taurea auct. Rhizocalathium Tzvel. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX (1959) 438.—Centaurea sect. Acrocentron*** Rhizanthae Boiss. Fl. or. III (1875) 618 and 674 p. p.—Capitula medium-sized sometimes rather large, with many florets, one each in center of rosette of basal leaves on strongly reduced peduncles (often subsessile). Involucre 10-25 mm in dia, broadly ovate, to subglobose, more or less thinly tomentose and also with very short, erect hairs to subglabrous; involucral bracts coriaceous, greenish, with 3—7 quite ‘inconspicuous veins, gradually elongated from broadly ovate outer to 566 567 564 lanceolate innermost; appendages of involucral bracts quite large, thin- coriaceous, triangular-lanceolate, less often ovate to suborbicular, of- ten recurved, usually not decurrent on bracts and not surpassing them in width, brownish or of various shades, gradually acuminate-spinescent, pectinately ciliate, with quite long, stiff cilia, with cilia many to 1-2 on each side, sometimes appendages entirely lacking, replaced by short deflexed spine at leaf tip. All florets tubular, various shades of yellow, 1.52.0 times as long as involucre, heterogeneous, peripheral florets sterile, not large (usually even smaller than bisexual central florets) with staminodes not exserted from corolla tube; stamen filaments weakly flattened, covered wtih very short, papilliform hairs; stigma shortly bifid. Achenes,4—6 mm long, sparsely pubescent, mature achenes subglabrous, somewhat lustrous; pappus 1/5—1/3 as long as achenes, persistent, double: outer pappus consisting of many irregular rows of scarious bristles, weakly scabrous on two sides, gradually elongated toward middle and inner; inner pappus of single row of scarious bristles, similar to outer but usually slightly thicker and shorter. Perennial herbs with long, strongly branched rhizome, producing vegetative shoots with rosette of basal leaves and reproductive shoots; leaves with quite long petioles, with leaf glades oblong, more or less divided or lobed, less often entire or subentire, covered with short crisped hairs and incon- spicuous punctate, glands. Type of subgenus: C. rhizantha C.A.M. The seven species of the subgenus, which, except for C. oltensis, are very closely related, are distributed in Turkey, Iran, and Transcaucasia, extending also to the montane regions of Turkmenia. Five species are found in the USSR. In the original diagnosis of the section Rhizocalathium, based exclusively on herbarium material, the species belonging here were described as not producing rhizomes. Later observations in nature have shown that this is incorrect; long, strongly branched rhizomes are, on the contrary, a very typical characteristic of the subgenus Rhizocalathium, clearly distinguishing it from the subgenus Lopholoma in whose species stolons are always absent. 1. ‘Lateral segments of pinnately incised leaves many (8-16 on each side), sinuate-toothed or lobed, almost always narrowed toward base, terminal lobe almost equaling lateral ones; appendages of involucral bracts deflected from involucre, but not recurved, long andswide wie ei tr bate cethseh erie 2 SA. 171. C. oltensis Sosn. + Lateral segments of lyrate or pinnately lobed leaves fewer (up to 6-8 on each side), entire, basally usually broadened, terminal lobe almost always larger than lateral ones; appendages of involucral 568 bracts recurved, rarely partly ascending «00.0.0... eee eeeeeeeeeeeeees 2. 2! Florets light-yellow (pale); leaf blades relatively weakly incised, with few (1-3 on each side) lateral lobes only near base, often partly undivided and entire. Southern Georgia and northern Arme- mii Heydar oh one) es. Rela Ls es 171. C. rhizanthoides Tzvel. + Florets yellow; leaf blades relatively more incised, usually lobed or divided up to half or more than half the length, with 1-8 lobes on Sriclipsidenseht hate ence tie he eA A, LA ee 3. | Appendages of involucral bracts very large, partly ascending, al- most covering involucre, to 12 mm long and 2.5—3.6 mm wide (excluding lateral cilia), cilia to 6 mm long, 5-10 on each side; involucre 20-25 mm in dia... eee 172. C. armena Boiss. + Appendages of involucral bracts smaller, recurved, lateral cilia isuallyishortenand fewer. 2k 2208 Re ee 4. 4. Leaf blades rather narrow, relatively more incised, with 3—8 lobes on each side; terminal lobe not much longer than lateral ones, usu- ally much shorter than half the blade length. Nakhichevan ASSR .... Rowiiie soe otel lantoriat..cbie. fans it 170. C. grossheimii Sosn. + Leaf blades less incised, with 1—5 lobes on each side; terminal lobe much longer than lateral ones; often nearly half as long as blade ..... upeneniins: |. oA aentre tl catmbergel Eure uate Cen e 169. C. rhizantha C.A.M. Series 1. Rhizanthae Tzvel.—Involucral bracts with more or less recurved appendage; lateral lobes or leaf segments usually entire. 169. C. rhizantha C.A.M. Verzeichn. (1831) 64; DC. Prodr. VI, 594; Ldb. Fl. Ross. II, 2, 710; Sosn. in Fl. Gruzii, VIII, 564; Tzvelev in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX. 439.—C. sessilis auct. non Willd. Boiss. Fl. or. III (1875) 676; Grossh. Fl. Kavk. IV, 213 and Opred. Rast. Kavk. 497.—Exs.: GRF No. 4398. Perennial. Plants stemless or with strongly reduced (to 5 cm long) stem, green, more or less crisped-hairy. Leaves lyrate or pinnately lobed, rather long-petiolate, with 1-5 lobes on each side, terminal lobe much larger than lateral ones, often hastate or sagittate. Capitula usu- ally solitary, in center of leaf rosette. Involucre 15—25 mm in dia; appendages of involucral bracts lanceolate, recurved, ciliate, usually with 1—6 cilia on each side. Florets yellow. Achenes 4—6 mm long, with whitish pappus 1—2 mm long. Flowering June to July (Plate XXVII, Fig. 2). Stony slopes, meadows, more moist places of mountain steppes, mostly in mid-montane zone.—Caucasus: Eastern Transcaucasia (south- western part), Southern Transcaucasia (southeast of Aragats Moun- tain), Talysh; Soviet Central Asia: mountainous Turkmenia. General 569 566 distribution: Armenia and Kurdistan, Iran (northern part). Described from Talysh. Type in Leningrad; isotype in Tbilisi. Note. Apparently, even in the more restricted sense considered by me, this species splits up into many weakly differentiated ecogeographical races, differing mostly by the size and shape of the appendages of the involucral bracts. From Nakhichevan ASSR, a sterile intergeneric hybrid is known: C. rhizantha x Tomanthea phaeopappa (Shakhbuz District, stony steppe, 2-3 km northwest of the village of Buzgov, 26.VI.1957, N. Tzvelev and S. Czerepanov). 170. C. grossheimii Sosn. in Bot. Zhurn. SSSR, XXXIV, 3 (1949) 288; Grossh. Opred. Rast. Kavk. 497.—C. rhizantha auct. non C.A.M.; Tzvelev in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XIX (1959) 439 p. p.; Karjagin in Fl. Azerb. VIII, 474 p. p. Perennial. Plants stemless or with strongly reduced stem, green or grayish-green, quite densely crisped-hairy. Leaves pinnately lobed or parted, with 3—8 lobes on each side, terminal lobe not much longer than lateral ones, usually much shorter than half length of leaf blade. Capitula usually solitary, incenter or leaf rosettes. Involucre 15—22 mm in dia; appendages of involucral bracts lanceolate, recurved, cili- ate, usually with 3-7 cilia on each side. Florets yellow. Achenes 4—6 mm long, with pappus 1-2 mm long. Flowering June to July. Stony and rubbly limestone slopes, at 1,000—2,000 m.—Caucasus: Southern Transcaucasia (Nakhichevan ASSR). Endemic(?). Described from Nakhichevan ASSR (Karagut Massif). Type and isotype in Leningrad; isotype in Tbilisi. Note. A very weakly separated. more xerophilous and lower mon- tane ecogeographical race associated with limestone massifs of Nakhichevan ASSR. 171. C. rhizanthoides Tzvel. in Addenda XXVII, 620.—C. sessilis auct. fl. cauc. p. p. quoad pl. Georgiae et Arm. bot.—C. sessilis var. integrifolia Bordz. ex Grossh. Fl. Kavk. IV (1934) 213. Perennial. Plants stemless or with strongly reduced stem, green, more or less crisped-hairy. Leaves lyrate, with very long petioles, few (1-3 on each side) lateral lobes only closer to base, often partly un- divided and entire. Capitula usually solitary in center of leaf rosette. Involucre 12—20 mm in dia; appendages of involucral bracts lanceolate, recurved, ciliate, usually with 1-5 cilia on each side. Florets light yellow (pale). Achenes 4—5 mm long, with pappus 1—2 mm long. Flow- ering June to July. 570 567 Stony slopes, meadows, pastures; mostly in middle mountain zone.—Caucasus: Western Transcaucasia (southern part), Southern Transcaucasia (north of Aragats Mountain and Lake Sevan). General distribution: Asia Minor (eastern part), Armenia and Kurdistan. De- scribed from Armenia (Sevan District, meadow-steppe stony slope toward Lake Sevan, 3—4 km northeast of city of Sevan, 13 July 1957, No. 875, N. Tzvelev and S. Czerepanov). Type in Leningrad. Note. Mass herbarium material, collected by S.K. Czerepanov and me in the summer of 1960 in the region of the Dzhavakhet and Pambak ranges, revealed considerable stability of the morphological characters of this northernmost of ecogeographical races of C. rhizantha s. 1. C. rhizantha s. str. is found also on the slopes of Aragats Mountain, but in the area of Lake Sevan the geographical boundary between the ranges of these two species is still not altogether clear. Near the state border with Turkey, it is possible to find specimens of another similar species—C. rhizocalathium (C. Koch) Boiss.—in which the append- ages of the involucral bracts are reduced to a single spine without lateral cilia. 172. C. armena Boiss. Fl. or. III (1875) 676. Perennial. Plants stemless or with strongly reduced stem, green, more or less crisped-hairy. Leaves lyrate or pinnately lobed, with very long petioles and 1—5 lobes on each side, terminal lobe much longer than lateral ones. Capitula usually solitary, in center of leaf rosette. Involucre 20—25 mm in dia; appendages of involucral bracts lanceolate, very large, often ascending, almost covering bracts, to 12 mm long and 2.5-3.5 mm wide (excluding lateral cilia), cilia 5-10 on each side. Florets yellow. Achenes 5—6 mm long, with pappus 1-2 mm long. Flowering June to July. Stony slopes.—Caucasus: Southern Transcaucasia (Daralgez). General distribution: Armenia and Kurdistan. Described from north- eastern Turkey. Type in Geneva; isotype in Leningrad. Note. The specimens available from Daralgez (Ekhegnadzor Dis- trict, Armenian SSR, 3—4 km northwest of the village of Ortakend, 8 July 1957; Nos. 781 and 785, N. Tzvelev, and S. Czerepanov) are not entirely consistent with the typical plants of this species, differing primarily by the recurved appendages of the involucral bracts, and, possibly, belong to a separate Daralgez ecogeographical race. Series 2. Oltenses Tzvel.—Involucral bracts usually with ascend- ing appendages; lateral lobes or segments of leaf blades numerous, more or less sinuately toothed or lobed. 571 568 173. C. oltensis Sosn. in Zam. po Sist. i Georg. Rast. Tbil. XXI (1959) 60.—C. ustulata auct. non DC.: Grossh. Opred. Rast. Kavk. (1949) 497. Perennial. Plants stemless or with strongly reduced stem, crisped- hairy, mixed with some arachnoid-hairs, green. Leaves pinnately in- cised, with very long petioles, and 8—16 lobes on each side, unequal, mostly sinuate-toothed or lobate, usually narrowed toward base, termi- nal lobe almost as long as laterals. Capitula solitary, in centre of leaf rosettes. Involucre 14—20 mm in dia; appendages of involucral bracts lanceolate or broadly lanceolate, divergent from involucre but ascend- ing, with 3-6 stiff cilia on each side. Florets yellow. Achenes 4—6 mm long; pappus about 1.5 mm long, whitish; inner pappus bristles much thicker than outer pappus bristles. Flowering June to July. Stony limestone slopes of middle mountain zone.—Caucasus: Southern Transcaucasia (Karagut Massif in Nakhichevan ASSR). Gen- eral distribution: Armenia and Kurdistan (northwestern part). Described from northwestern Turkey (near Oltu). Type in Tbilisi. Note. Specimens of this species from the Karagut Massif (from the Herbarium of the Botanical Institute of the Academy of Sciences of Azerbaidzhan SSR) were identified by A.A. Grossheim as “C. ustulata DC.” However, this latter species (there are isotypes of it in the Herbarium of the Botanical Institute of the Academy of Sciences of the USSR) differs from C. oltensis by a whole series of significant characters (structure of the involucre, color of the florets, and others). The great similarity in the structure of the involucre of C. oltensis with the hybrid, C. rhizantha x Tomanthea phaeopappa, mentioned above, prompts me to suggest the possibility of a hybrid origin of the plants of C. oltensis, as a result of the hybridization of C. rhizantha with some species of the genus Tomanthea. Subgenus 20. Solstitiaria (Hill.) Dobrocz. in Bot. Zhurn. Akad. Nauk URSR, VI, 2 (1949) 64, 69.—Solstitiaria Hill, Veg. Syst. IV (1762) 21.—Leucantha S.F. Gray, Nat. Arrang. Brit. Pl. II (1821) 443.—Mesocentron and Triplocentron Cass. in Dict. Sc. nat. XLIV (1826) 38.—Verutina Cass. op. cit. XLIV (1826) 38, LVIII (1829) 8.—Centaurea sect. Mesocentron (Cass.) DC. Prodr. VI (1837) 592 p. p.; Boiss. Fl. or. Ill (1875) 619; O. Hoffm. in Pflanzenfam. IV, 5, (1893) 331.—Centaurea subgen. Centaurea Schmalh. Fl. Yugo-Zap. Ross. (1866) 329 p. p.; Fl. II (1897) 120 p. p.—Centaurea subgen. Calcitrapa sect. Mesocentron (Cass.) Hayek, Centaurea-Art. Osterr.- Ung. (1901) 3, 15.—Calcitrapa auct. p. p. non Adans.: Lam. FI. Fr. II (1778) 29; Dostal, Kli¢. (1958) 734.—Capitula medium-sized, with many florets, solitary at tips of stem and lateral branches, aggregated 569 in racemose or paniculate-racemose inflorescence. Involucre ovate- conical, 7—12(15) mm in dia; involucral bracts coriaceous, greenish (yellowing on drying), smooth, from ovate outer to oblongly lanceolate and oblong inner; outer and middle bracts with 5(7) palmately divaricate spines, of which central spine usually much longer (up to 30 mm) and thicker than others (2.5—4.0 mm long); inner bracts with small mem- branous, unevenly toothed appendage. All florets tubular, yellow, het- erogeneous; central florets bisexual; peripheral ones sterile lacking staminodes, very slightly shorter than central florets, with 3-5 fid corolla; stamen filaments flattened, covered with papilliform hairs; style short-bifid or not (its branches connate to tip). Achenes oblong- ellipsoidal or oblong obovate, 2.5 mm long, initially scatteredly hairy, mature ones glabrous; pappus as long as achenes, persistent, double outer pappus consisting of many rows of flat, serrate-scabrous bristles, elongated from outer to inner; inner pappus much shorter than outer, of single row of bristles. Biennial herbs; stem upright, branched, nar- rowly-winged, leafy to tip; cauline leaves sessile, decurrent, with stem arachnoid-tomentose, mostly entire, lower leaves short-petiolate, lyrately pinnately incised, to undivided and even entire. Type of subgenus: C. solstitialis L. Of the 17 species of this subgenus growing in southern Europe, northern Africa, and southwestern Asia, only two are found in the USSR. 1. Involucre 13—15(18) mm long and 7—12(15) mm in dia, outer and middle involucral bracts with 5(7) yellow spines, of which central spine is much thicker and longer than others ..............::.cssccceseeeees Shaklee cttatan Lec emted. aed, eel Raut, 2opaiaets aus ble 174. C. solstitialis L. + Involucre 12—13(15) mm long and 7—8(10) mm in dia, outer and middle involucral bracts with 5(7) usually brownish spines, of which central spine is thin and short, slightly longer than others or almost Asrlonppedt.. cece deinen tt ai. meee ented ey: 175. C. adamii Willd. 174. C. solstitialis L. Sp. pl. (1753) 917; DC. Prodr. VI. 594 p. p.; Ldb. Fl. Ross. II, 2, 710 p. p.; Boiss. FI. or. III, 685 p. p.; Schmalh. Fl. II, 121; Hayek, Centaurea-Art. Osterr.-Ung. (1901) 18; Briquet, Monogr. Centaurees Alpes Maritimes, 158; Fedtsch. and Fler. Fl. Evrop. Ross. 1032 p. p.; O. and B. Fedtsch. Perech. Rast. Turk. IV, 272, excl. syn. C. adamii Willd.; Grossh. Fl. Kavk. IV, 210; Opred. Rast. Kavk. 495; Dobrocz. in Vizn. Rosl. URSR, 582; Sosn. in Fl. Gruzii, VIII, 562; Stank. and Tal. Opred. Vyssh. Rast. ed. 2, 414; Czer. in FI. Turkm. VII, 274.—C. sicula auct. non L.: Guldest. Reise, I (1787) 572 196, nomen.—C. adamii auct. non Willd.: MB. FI. taur.-cauc. II, 355, 570 III, 593.—?C. cyanifolia Poir. in Lam. Encycl. meth. Suppl. I (1811) 158.—C. solstitialis ssp. eu-solstitialis var. typica Gugler in Ann. hist.- nat. Mus. Nat. Hung. VI (1908) 203.—Solstitiaria solstitialis (L.) Hill. Veg. Syst. IV (1762) 21.—S. flava Hill. Herb. Brit. I (1769) 77.— Calcitrapa solstitialis (L.) Lam. Fl. Fr. I (1778) 34.—C. solstitialis ssp. solstitialis: Dostal, Klic. (1958) 734.—Cyanus solstitalis (L.) Baumb. Enum. strip. Transsilv. HII (1816) 79.—Leucantha cyanifolia S.F. Gray, Nat. Arrang. Brit. Pl. II (1821) 444.—L. solstitialis (L.) A. and D. Love in Bot. Notis. CXIV, 1 (1961) 44.—Seridia solstitialis (L.) Sweet, Hort. Brit. ed. 1 (1827) 218.—Triplocentron solstitiale (L.) Fourr. in Ann. Soc. Linn. Lyon, nov. ser. XVII (1867) 97.—Ic.: Hill, Herb. Brit. I (1769) tab. 54, fig. 2; Rchb. Ic. Fl. Germ. et Helv. XV (1852) tab. 795, I-1; Hayek, op. cit. Taf. Il, fig. 3; Hegi, Ill. Fl. VI, 2 (1929) fig. 668a; Javorka and Csapody, Iconogr. Fl. Hung. (1933) 545, fig. 3895.—Exs.: GRF No. 1879; Fl. cauc. exs. No. 224; Sintenis. It. trans-casp.-pers. 1900-1901, No. 388; Bornm. Pl. exs. Anat. or. 1889, No. 1502; Kotschy, It. cilic.-kurd. 1859, No. 314; FI. exs. austro- hung. No. 3435; Schultz, Herb. norm. No. 2215; Billot. Fl. Gall. et Germ. exs. No. 266; FI. exs. reip. Bohem.-Sloven. No. 1181; Rchb. FI. Germ. exs. No. 316. Biennial. Whole plant grayish-arachnoid-tomentose. Stem (15)30— 100 cm high, erect, ribbed, narrowly-winged, mostly branched from middle and below, with few upward-spreading branches, or simple. All leaves scabrous along margin from small acute spines; lower leaves lyrately pinnately incised to undivided or even entire, short-petiolate, withering early, their lateral segments triangular, oblong-lanceolate or oblong with remote small teeth or entire, terminating in small cusp, 2— 6 on each side; terminal segment rather large, lanceolate, elliptical or almost rhombic, with occasional small teeth or entire; cauline leaves lanceolately linear, usually entire, awned, sessile, long-decurrent on stem. Capitula solitary at tips of stem and lateral branches, aggregated in racemose or paniculate-racemose inflorescence. Involucre ovate- conical 13—15(18) mm long and 7—12(15) mm in dia, weakly arach- noid-hairy; outer and middle involucral bracts with 5(7) palmately spreading yellow spines, of which central spine in middle bracts is stiff, long (12-30 mm), more or less strongly recurved, others thin, short (2.5—4.0 mm long) at base of central spine, 2(3) on each side; middle spine of outer bracts mostly thinner and shorter (4—7 mm long); inner bracts with small membranous, unevenly toothed appendages. Florets yellow. Achenes 2.5 mm long and about 1 mm wide, outermost pappus, others with 5 mm long pappus. Flowering May to September (October). SS 571 Dry rubbly and clayey slopes, foothills and lower mountains; weedy places.—European Part: Bessarabia, Black Sea Region (western half of region), Middle Dnieper (extreme southwest), Volga-Don (vicinity of Kharkov, introduced). Crimea (south); Caucasus: entire region; Soviet Central Asia: mountainous Turkmenia, Pamiro-Alai (western part). General distribution: Iran, Armenia and Kurdistan, Balkans-Asia Mi- nor, Mediterranean, Central Europe (southern part), Atlantic Europe (introduced), North America (introduced). Described from western Europe. Type in London. Note. The absence of true C. adamii Willd. in Bieberstein’s herbarium forces me to consider C. adamii MB. as a synonym. 175. C. adamii Willd. Sp. pl. III, 3 (1803) 2310; DC. Prodr. VI, 594; Hayek, Centaurea-Art. Osterr.-Ung. (1901) 16 p. p.; Grossh. FI. Kavk. IV, 210; Opred. Rast. Kavk. 495; Sosn. in Fl. Gruzii, VIII, 563; Stank. and Tal. Opred. Vyssh. Rast. ed. 2, 414.—C. solstitialis B. adamii (Willd.) Heuff. in Verh. Zool. Bot. Ges. Wien. VII (1858) 144; Boiss. FI. or. III, 685, excl. syn. C. lappacea Ten.—C. solstitialis ssp. eu-solstitialis var. adamii (Willd.) Gugler in Ann. hist.-nat. Mus. Nat. Hung. VI (1908) 203.—Seridia adamii (Willd.) Sweet, Hort. Brit. ed. 1 (1827) 218.—Calcitrapa adamii (Willd.) Schur. Enum. pl. Transsilv. (1866) 409.—C. solstitialis ssp. adamii (Willd.) Dostal, Kli¢ (1958) 734.—Ic.: Rchb. Ic. Fl. Germ. et Helv. XV (1852) tab. 795, II- 2; Hayek, op. cit. Taf. II, fig. 2. Biennial. Differs from preceding species in insignificant features: involucre much smaller, 12—13(15) mm long and 7—8(10) mm in dia; spines of outer and middle involucral bracts usually brownish and more or less equal; central spine in some middle involucral bracts often slightly thicker and longer (4—7, less often to 10 mm long) than others. Flowering June to August. Dry slopes, steppes, and weedy places.—European Part: Bessarabia, Black Sea Region (environs of Odessa; settlement of Sositskaya), Middle dnieper (southern Podolia: former Balta District), Crimea (Lake Saks; Kazantin Peninsula); Caucasus: Ciscaucasia, Dagestan (vicinity of Khasavyurt), Eastern Transcaucasia. General dis- tribution: Balkans (Macedonia and northward), Central Europe (ex- treme southeast). Described from Georgia. Type in Berlin. Note. Judging from the herbarium material at our disposal, C. adamii Willd. is quite common in Central Ciscaucasia and in the east- ern part of Western Ciscaucasia. In Eastern Transcaucasia it is found in the lesser Caucasus in the Kura River basin, reaching in the south- 574 572 east as far as the Agdam District of the Azerbaidzhan SSR (village of Kuzanla). Subgenus 21. Calcitrapa (Adans.) Hayek, Centaurea-Art. Osterr.- Ung. (1901) 2, 11 p. p.—Calcitrapa Adans. Fam. II (1763) 116; Lam. Fl. Fr. II (1778) 29 p. p.; Juss. Gen. pl. (1789) 173; Dostal, Klic (1958) 734 p. p.—Happophaestum S.F. Gray, Nat. Arrang. Brit. Pl. II (1821) 443.—Centaurea sect. Calcitrapa (Adans.) DC. Prodr. VI (1837) 596; Boiss. Fl. or. III (1875) 619; O. Hoffm. in Pflanzenfam. IV, 5 (1893) 331.—Centaurea subgen. Centaurea Schmahh FI. II (1897) 120 p. p.—Centaurea subgen. Calcitrapa sect. Eucalcitrapa Hayek b) Carduiformes Hayek, op. cit. 3—Capitula medium-sized, with many florets, solitary at tips of lateral branches and reduced branchlets, nu- merous on plant. Involucre oblongly ovate, ovate or globosely ovate, 6—14 mm in dia, glabrous; involucral bracts coriaceous, greenish (yel- lowing on drying), smooth, from round-ovate outer to oblong-ovate and oblong inner; outer and middle bracts with (3)5—7 palmately spread- ing spines, of which central spine is much longer (10-30 mm) and thicker than others (3—5 mm long); inner involucral bracts with small, membranous, weakly irregularly toothed appendage. All florets tubu- lar, pinkish-violet, heterogeneous: central florets bisexual; peripheral ones sterile, lacking staminodes, scarcely larger, with 5—6-fid corolla; stamen filaments flattened, covered with papilliform hairs; style weakly bifid. Achenes obovate to oblong-ellipsoidal, 3—4 mm long, initially scatteredly hairy, mature ones glabrescent; pappus absent or shorter than achene, white, persistent, double; outer pappus many-rowed, con- sisting of flat, serrate-scabrous bristles, elongated from short outer to inner; inner pappus 1/4—1/3 as long as outer, of single row of bristles. Biennial herbs; stems upright, strongly divaricately branched, leafy to tip; cauline leaves sessile, not decurrent, usually pinnately incised, basal and lower cauline short-petiolate. Type of subgenus: C. calcitrapa L. The subgenus comprises 14—16 species, of which only two grow in our country. is) SAchenestwith pappus ee. ee See 176. C. iberica Trev. +" Achenesdlackinopappus i 2-e.3. 2h. oe 177. C. calcitrapa L. 176. C. iberica Trev. in Spreng. Syst. vet. 3 (1826) 406; DC. Prodr. VI, 597; Ldb. Fl. Ross. II, 2, 711; Boiss. Fl. or. III, 690 p. p.; Hayek, Centaurea-Art. Osterr.-Ung. (1901) 13; in Fedde, Repert. Beih. XXX, 2, 793 (Prodr. Fl. Balc.); O. and B. Fedtsch. Perech. Rast. Turk. IV, 272; Grossh. Fl. Kavk. IV, 209; Opred. Rast. Kavk. 495; Sosn. in SS) 575 Fl. Gruzii, VIII, 562; Stank. and Tal. Opred. Vyssh. Rast. ed. 2, 414; Czer. in Fl. Gruzii, VII, 275.—C. calcitrapa auct. non L.: MB. FI. taur.-cauc. II, 356 p. p., II, 594 and auct. al. fl. Cauc. et As. Med. — C. calcitrapoides auct. non L.; DC. op. cit. 597 p. p.; Ldb. op. cit. 711 and auct. fl. Cauc. et AS. Med.—C. calcitrapa var. iberica (Trev.) Schmalh. Fl. II (1897) 122, excl. sun.—C. pallescens var. iberica f. genuina Gugler in Ann. hist.-nat. Mus. Nat. Hung. VI (1908) 209.— C. pallescens ssp. iberica var. genuina (Gugler) Arenes in Compet. Rend. Séances soc. Biogeogr. 237 (1950) 137.—Calcitrapa iberica (Trev.) Schur. Enum. Pl. Transsilv. (1866) 409.—Leucantha iberica (Trev.) A. and D. Léve in Bot. Notis. CXIV, 1 (1961) 44.—Ic.: Hayek, Centaurea-Art. Osterr.-Ung. (1901) Taf. 1, fig. 4; Javorka and Csapody, Iconogr. Fl. Hung. (1933) 545, fig. 3897.—Exs.: GRF No. 1877; Herb. Fl. cauc. No. 296; Hayek, Cent. exs. crit. fasc. I, No. 47; Fl. exs. austro-hung. No. 3436; Schultz, Herb. norm. No. 2652; Sintenis, It. trans-casp.-pers. 1900-1901, No. 961. Biennial. Stem 20—100 cm high, erect, ribbed, strongly divaricateiy branched from middle or below, scabrous, pale green, less often gray- ish-green, usually not densely covered with flexuous articulate hairs, sometimes glabrous. Leaves green, scabrous-floccose or almost sca- brous from fine, quite stiff hairs, covered with golden, punctate, sessile glands; basal leaves lyrately pinnately parted to incised, petiolate, withering early; cauline leaves pinnately incised, sessile, not decurrent on stem, upper ones mostly undivided or subentire, their leaf segments narrowly lanceolate-linear, narrowly lanceolate, oblongly lanceolate or almost oblong, mostly with remote acute teeth, terminating in small, white, thin, cartilaginous cusp. Capitula solitary at tips of lateral branches and reduced branchlets, numerous on plant. Involucre ovate or broadly ovate, 13-16 mm long and 10-14 mm in dia; outer and middle involucral bracts with (3) 5—7 palmately spreading spines, of which central spine is much longer (10—30 mm) and thicker than oth- ers (3 mm long); inner bracts with very small, membranous, weakly irregularly toothed appendages. Florets pinkish-violet. Achenes 3—4 mm long, 1.2—1.5(1.7) mm wide; pappus 2.0—2.5(3.0) mm long. Flow- ering May to August (September). Along irrigation ditches, banks of mountain streams, roadsides, and wastelands, in foothills and low- sometimes mid-montane zone, less often on slopes and foothill plains—European Part: Crimea (south); Caucasus: all regions; Soviet Central Asia: mountainous Turkmenia, Amu-Darya, Pamiro-Alai, Syr-Darya, Tien Shan, Lake Balkhash Region (foothill areas of Semirechye), Dzhungaria-Tarbagatai (western part of Dzhungarian Alatau). General distribution: Dzhungaria- Kashgaria (Kuldzha), Indo-Himalayas (Kashmir, Punjab), Iran, Arme- 576 574 nia and Kurdistan, eastern Mediterranean (Asiatic part), Balkans-Asia Minor. Described from Armenia. Type in Bonn or Bratislava. 177. C. calcitrapa L. Sp. pl. (1753) 917; DC. Prodr. VI, 597 p. p. excl. y. brevicaulis; MB. Fl. taur.-cauc. II 356 p. p.; Ldb. Fl. Ross. II, 2, 711; Boiss. Fl. or. III, 689; Schmalh. Fl. II, 122 p. p.; Hayek, Centaurea-Art. Osterr.-Ung. 11; in Fedde, Repert. Beih. XXX, 2, 793 (Prodr. Fl. Balc.); Brequet, Monogr. Centaurees Alpes Maritimes, 162; Fedtsch. and Fler. Fl. Evrop. Ross. 1032 p. p.; Stank. and Tal. Opred. Vyssh. Rast. ed. 2, 414.—C. calcitrapa var. eu-calcitrapa Stojan. and Acht. in Stojan. and Stef. Fl. B’lg. ed. 3 (1948) 1209.—Rhaponticum calcitrapa (L.) Scop. Fl. Carn. ed. 2, II (1772) 136.—Calcitrapa stellata Lam. FI. Fr. II (1778) 34; Dostal, Klic (1958) 734.—C. hippophaestum Gaertn. De. fruct. II (1791) 376.—Hippophaestum vulgare S.F. Gray, Nat. Arrang. Brit. Pl. II (1821) 443.—Ic.: Fedtsch. and Fler. op. cit. Figs. 1051 and 1052; Stojan. and Stef. op. cit. Fig. 1253; Hayek, op. cit. (1901) Taf. I, fig. 3; Javorka and Csapody, Iconogr. Fl. Hung. (1933) 545, fig. 3896; Hegi, Ill. Fl. VI, 2 (1929) Taf. 272, fig. 3; fig. 668b, c; Dostal, op. cit. tab. 280, fig. 2496.—Exs.: Hayek. Cent. exs. crit. fasc. I, Nos. 45 and 46; FI. exs. reip. Bohem.-Sloven. No. 1182; Billot, Fl. Gall. et Germ. exs. No. 1902; Schultz., Herb. norm. No. 2757; Todaro, Fl. Sic. exs. No. 913. Biennial. Stem 15—60 cm high, erect, ribbed strongly divaricately branched from middle or below, quite scabrous, pale green, with leaves covered with more or less scattered, flexuous, articulate hairs, rarely subglabrous. Leaves green, sometimes, mostly in youg stage, densely woolly, with golden, punctate, sessile, glands, pinnately incised; basal leaves petiolate, withering early; cauline leaves sessile, not decurrent on stem; upper ones deeply lobed to entire, their segments narrowly lanceolate-linear or narrowly lanceolate with remote acute teeth, less often lobed, terminating in small, white, thin, cartilagineous cusp. Capitula solitary at tips of lateral branches and reduced branchlets, numerous on plant. Involucre oblong ovate, 13-15 mm long and 6— 8(10) mm in dia; outer and middle involucral bracts with 5—7 pal- mately spreading spines, of which central spine is much longer (10-18 mm) and thicker than others (4-5 mm long); inner bracts with small, membranous, weakly irregularly toothed appendages. Florets pinkish- violet. Achenes about 3 mm long and 1.5—1.7 wide; pappus absent. Flowering June to September. Weedy places and dry slopes.—European Part: Crimea (south). General distribution: Mediterranean, Atlantic and Central Europe, Balkans-Asia Minor, Madeira Island, Canary Islands, North America (introduced). Described from western Europe. Type in London. Si oo 573 Note. The report (Hooker f. FI. Brit. Ind. III (1882) 385) of the occurrence of C. calcitrapa L. in northwestern India (Kashmir and Punjab) is extremely doubtful and needs verification. Subgenus 22. Tetramorphaea (DC.) Czer. comb. nova.— Tetramorphaea DC. in Guill., Archiv. Bot. II (1833) 331; Prodr. VI (1837) 609.—Centaurea sect. Tetramorphaea (DC.) Boiss. FI. or. III (1875) 684; O. Hoffm. in Pflanzenfam. IV, 5 (1893) 329.—Centaurea subgen. Calcitrapa sect. Eucalcitrapa a) Tetramorphae (DC.) Hayek, Centaurea-Art. Osterr.-Ung. (1901) 3—Capitula quite small, with many florets, enveloped by apical leaves, solitary at tips of short branches, numerous on plant. Involucre ovate, 6—7 mm in dia (without recurved spines), scarcely arachnoid-hairy or glabrous; involucral bracts coria- ceous, greenish (yellowing on drying), smooth, from broadly ovate outer to oblongly linear innermost; appendages of outer bracts larger, not decurrent on bract, green, leafy, those of middle bracts white, strongly recurved, long (10-22 mm), stiff spine; appendages of inner- most bracts small, membranous, irregularly toothed. All florets tubu- lar, lilac-pink, heterogeneous; central florets bisexual; peripheral ones sterile, lacking staminodes, fewer, not large, with (3)4-fid corolla; stamen filaments flattened, covered with papilliform hairs; stigma weakly bifid. Achenes ellipsoidal-obovate or ellipsoidal, 2.0—-2.3 mm long, glabrous, immature ones with occasional hairs; pappus as long as achene or slightly longer, white, persistent, double, outer pappus many- rowed, consisting of irregular rows of fewer, scarcely serrate-scabrous bristles, gradually elongated from outer to inner; inner pappus much shorter than outer, of single row of very small narrow oblong-lan- ceolate scales smooth or indistinctly erose-toothed above, connate at base in ring. Annual herbs; stems upright, divaricately branched mostly from base, white, leafy to capitula; cauline leaves sessile, broad, amplexicaul-cordate with stem scabrous from flexuous, articulate, long cilia, stiffly, pectinate-ciliate, spinous-toothed, sometimes partly with small teeth, undivided; basal and lower cauline leaves petiolate, usu- ally lyrately divided or lyrately pinnately divided, withering early. Type of subgenus: C. bruguieriana (DC.) Hand.-Mazz. The subgenus includes two species, which grow in Southern Transcaucasia, Soviet Central Asia, and West Asia (from northwestern India to Mesopotamia inclusively). 178. C. belangeriana (DC.) Stapf in Denkschr. Acad. Wien, I (1885) 66, n. v.; Bornm. in Beih. Bot. Centralbl. LIX, Abt. B (1939) 305; Czer. in Fl. Turkm. VII, 275.—C. phyllocephala var. persica Boiss. Diagn. pl. or. ser. I, 6 (1845) 134.—C. phyllocephala B. 576 i] WE ~= oS — VA, 4 . We (Zoe 577 Plate XXVII. 1—Stizolophus coronopifolius (Lam.) Cass.; 2—Centaurea rhizantha C.A.M.; 3—Phalacrachena calva (Ldb.) Mjin. 579 ay % belangeri Boiss. FI. or. III (1875) 684.—C. phyllocephala auct. fl. As. Med. et Cauc. non Boiss.—C. bruguieriana var. belangeri Parsa, F1. Iran. III (1943) 687, sphalm. “bruguiera.”—Tetramorphaea belangeriana DC. Prodr. VI (1837) 609; Karjagin in Fl. Azerb. VIII, 440.—T. bruguieriana auct. non DC.: Grossh. Opred. Rast. Kavk. (1949) 490.—Exs.: Sintenis, It. trans-casp.-prs. 1900-1901, No. 360; Bornm. It. Pers.-turc. 1892-1893, Nos. 4079 and 4082. Annual. Plant (5)10-50 cm high, very strongly divaricately branched usually from base, quite slender, white, with leaves densely or more or less sparsely covered with flexuous, articulate, long cilia. Leaves covered with golden, punctate, sessile glands, acutely scabrous- toothed, sometimes some of them with small teeth; basal and lower cauline leaves larger, to 10(13) cm long (excluding petiole), elliptical or oblongly lanceolate, undivided to lyrately pinnatifid, less often lyrately divided, petiolate, withering early, others smaller, oblongly ovate, mostly undivided, sessile, broadly amplexicaul-cordate. Capitula enveloped by apical leaves, solitary at tips of short branchlets, numer- ous on plant. Involucre ovate, 9-10 mm long and 6—7 mm in dia (excluding recurved spines); appendages of outer involucral bracts large, green, leafy, of middle bracts white, strongly recurved, long (10-22 mm) stiff spines; appendage of innermost bracts small, membranous, irregularly toothed. Florets lilac-pink. Achenes 2.0—2.3 mm long and 0.8-1.0 mm wide; pappus 2.0—3.0(3.3) mm long. Flowering May to July, (Plate XXIV, Fig. 2). Dry, sandy, clayey, and stony places, loessic mounds, in foothill plains, foothills, and lower mountains, to 1,200 m; as weed in crop fields, old fields and wastelands, roadsides.—Caueasus: Southern Transcaucasia (Nakhichevan ASSR); Soviet Central Asia: mountain- ous Turkmenia, Karakum (southeastern), Pamiro-Alai, Syr-Darya. Gen- eral distribution: Iran, Indo-Himalayas (Punjab, Peshawar). Described from northern Iran. Type in Geneva. Note. Var. erecta Winkl. ex. O. and B. Fedtsch. (Perech. Rast. Turkm., IV, 272), named by Winkler and described by O. and B. Fedtschenko, apparently has no taxonomic significance. GENUS 1625. Phalacrachena Iljin': * Iljin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, VII, 3 (1937) Silt. "Treatment by N.N. Tzvelev. *From the Greek word phalacros—naked, and Latin word achena—seed; so named because achenes in the species of this genus lack a pappus. 580 578 Capitula heterogamous, medium-sized, with many florets, solitary at tips of stems. Involucre broadly ovate to subglobose, 10-18 mm in dia, glabrous or subglabrous; involucral bracts many-rowed, imbricate, herbaceous, almost coriaceous, with 4-7 weak veins, gradually elon- gated from broadly ovate outer to oblong innermost; appendage of outer and inner bracts as broad membranous border, more or less decurrent on bracts (sometimes to 1/2 to 1/3 their length), and usually narrower than bracts, brownish, acuminate or short-spinulescent, pectinately ciliate, in innermost bracts larger, oblong-ovate, recurved, almost not decurrent, with small teeth or irregularly ciliate. Recep- tacle setose, its bristles numerous, whitish. Florets various shades of pink, heterogeneous; central florets (disk florets) tubular, bisexual, 1.5—2.0 times as long as involucre; peripheral florets one-rowed, ster- ile, large, tubular-infundibuliform, lacking staminodes, with 4—5-lobed corolla; anthers with short basal appendages their apical appendages free above, lanceolate; stamen filaments weakly flattened, densely covered with short papilliform hairs; stigma short- or almost to middle bifid. Achenes oblong, 4—6 mm long, very weakly compressed, weakly arcuate, sparsely hairy only near hilum, mature achenes often entirely glabrous, slightly narrowed at apex and terminating in concave or al- most flat areole, lacking pappus, bordered with very weakly developed crown with rounded margin. Perennial herbs with long, more or less branched rhizome, and erect, simple or weakly branched stems; leaves short-petiolate or subsessile, entire or more or less pinnately lobed, covered with short papilliform hairs and more or less developed arach- noid-tomentum, usually punctate-glandular. Type of genus: P. inuloides (Fisch. and Schmaih.) Ijin. The two very distinct species of this small genus, which are quite closely related to the genus Centaurea L., are endemics of the USSR and are distributed in the south of the European part of the USSR and in Kazakhstan. 1. Plants to 40-50 cm high, with entire oblong-lanceolate leaves .......... safe vase ex RN VM, a ngs 1. P. inuloides (Fisch. ex Schmalh.) Ijin + Plants to 15-20 cm high, with pinnately lobed sometimes some of them with undivadedileaves wee.) s..s0-e- 2. P. calva (Ldb.) Ijin 1. P. inuloides (Fisch. ex Schmalh.) [jin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, VII, 3 (1937) 51; Grossh. Opred. Rast. Kavk. 501.—Centaurea inuloides Fisch. ex Schmalh. FI. II (1897) 126; Claus in Beitr. Pflanzenk. Russ. Reichs, VIII (1851) 231, nom. nud.— Phaeopappus inuloides Fisch. ex Nym. Consp. Fl. Eur. (1878-1882) 418, nom. nud.; Boiss. Fl. or. Suppl. 312, nom. nud. 58 a 579 Perennial. Plants to 40-50 cm high, covered with very short papil- liform hairs and spinules, usually with slight mixture of thin arach- noid-tomentum, scabrous. Stems solitary or few, erect, usually simple, with solitary capitulum at tip, less often with few lateral branches. Leaves oblong lanceolate, basally narrowed into short, winged petiole, or subsessile, subacute, entire, dark green, with numerous punctate glands. Capitula on more or less thickened, peduncles, leafless at tips. Involucre 12—18 mm in dia and 15—20 mm long; appendages of in- volucral bracts as broad, triangular border, acuminate, or spinulescent, with 8—10 cilia on each side, in innermost bracts larger, ovate, de- flected sideways shortly and irregularly-fimbriate. Florets pink; stigma weakly bifid. Achenes 5—6 mm long, lacking pappus. Flowering May to July. Alkaline and saline meadows, clayey slopes, old fields.—Euro- pean Part: Black Sea Region, Lower Don, Lower Volga. Endemic. Described from Krasnoarmeisk (former Sarepta.) area. Type in Leningrad. 2. P. calva (Ldb.) Iljin in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, VII, 3 (1937) 4.—Centaurea calva Ldb. in DC. Prodr. VI (1837) 583; Fl. Ross. II, 2, 703; Kryl. Fl. Zap. Sib. XI, 960. Perennial. Plants to 15—20 cm high, covered with very short papil- liform hairs and thin arachnoid-tomentum, sometimes subglabrous. Stems solitary or few, erect to almost prostrate, strongly shortened, usually simple, with solitary capitulum at tip. Leaves highly variable in shape, usually pinnately lobed with lobes of different lengths and width, often some of them narrowly linear and entire, narrowed toward base into short, winged petiole or subsessile, acuminate or with short cartilaginous cusp, from subglabrous, green to grayish-tomentose, with inconspicuous punctate glands. Capitula often surrounded by approxi- mate apical leaves. Involucre 10—14 mm in dia, and 13—16 mm long; appendages of involucral bracts as broad triagular border, short decur- rent on bract, acuminate, with 6-10 cilia on each side, lighter in color than appendage; in innermost bracts much longer, ovate, deflected laterally, toothed or irregularly ciliate. Florets pink; stigma bifid al- most to middle. Achenes 4-5 mm long, lacking pappus. Flowering May to July (Plate XXVII, Fig. 3). Sandy and gravelly banks of water bodies, alkaline areas of steppes, clayey and stony slopes.—Western Siberia: Upper tobol (southeastern part), Irtysh (southern part); Soviet Central Asia: Aralo-Caspian (north- eastern part), Lake Balkhash Region (northern part). Endemic. De- scribed from Eastern Kazakhstan. Type in Leningrad. 582 580 Note. In specimens from the northwestern part of the range, often almost all the leaves are entire, with involute margins; possibly these form a distinct ecogeographical race. GENUS 1626. Carthamus L.':? Sp. pl. (1753) 830; DC. Prodr. VI (1837) 611.—Kentrophyllum Neck. Elem. Bot. I (1790) 155, 86. Capitula homogamous, with many florets, solitary or corymbosely clustered, oval or oblong, 2—6 cm long, 1.5—2.5 cm in dia. Involucre many-rowed; involucral bracts variable, outer leafy, spiny-toothed, sometimes broadened in lower part, appressed or squarrose, twice as long as florets, slightly longer, equal or sometimes even shorter, others scarious, yellowish-green, sometimes with pearly luster on inside; middle bracts oval or ovally lanceolate, inner ones lanceolate; all undivided, spinescent or middle bracts with pectinate-ciliate append- ages or crenate-toothed. Receptacle flat, often with long scales or bristles. All florets bisexual, very rarely few outer ones sterile, yellow, stramineous, orange-yellow, red, purple, purple-pink, less often whit- ish; corolla regular or more or less regular, 5-fid, to 4.5 cm long; limb to 1.2 cm long; stamen filaments free, short, glabrous or hairy above; style branches short, almost connate. Achenes 4-angled-oval, or 4- angled obovate, sometimes with prominent ribs, smooth or, especially in upper part, rugose, glabrous, whitish, sometimes with spot 3—6 mm long, with lateral, often oblique hilum; sometimes pappus in lacking all florets or only in peripherals ones, often consisting of isolated scales or bristles; pappus scales unequal, outermost scales very short, sinuate, whitish, middle longer, sometimes 2—3 times as long as achene, serrate-toothed, acuminate, whitish, ochreous or blackish-purple; in- nermost scales often reduced, obtuse or acuminate, less often as long as middle or even longer, whitish; bristles ciliate-serrate, all equal or unequal, fragile. Annual plants, less often biennial or perennial. Stem cylindrical, straight, above cymosely or divaricately branched, very rarely prostrate, thick, sometimes slender, often especially in upper part floccose, woolly or even arachnoid-hairy, usually glaucous, fer- ruginous, sometimes glabrous, whitish, smooth, weakly lustrous, 0.3— 1.5 m high. Leaves alternate, sessile, semiamplexicaul, sometimes amplexicaul, lowermost leaves often narrowed into thin petiole; all 'Treatment by S.A. Shostakovsky. °The name originates either from the Arabic kurtum—the ancient name of dyer's saffron, cultivated in the Orient for its red dye, or from the Greek word katairo— cleanse, since the species of this genus are used as a laxative. 583 581 leaves coriaceous, less often somewhat stiff, lowermost usually deeply pinnately parted, others pinnately lobed, spiny-toothed, less often subentire, lacking cusp, often glandular, pubescent, sometimes glabrous or subglabrous, smooth, up to 15 cm long and to 5 cm wide. Lectotype of genus: C. tinctorius L. The genus is exclusively Mediterranean (sensu lato), with its range extending from Soviet Central Asia and western India to the Canary and Madeira islands and from Ethiopia to Central Europe, inclusively. Five species are found in the USSR. 1. Plants glaucous; florets purple-pink .................... 2. C. glaucus MB. + Plants glabrous or pubescent but not glaucous; florets yellow, pale Se Moy meme aes os heh ae ne ih eal Ra de Ua eal ab edled abies 2 2. Plants more or less pubescent (woolly, arachnoid-hairy, glandular) eenantnane ll Greansereltonnnh i biak te. wicila callie oo! he at Lissa. Koel eee 3h ey) PuplantssplabROUSie+ ventions). caielyl eulels 0). teh.. ayo) eel amber, eas, 4. 3. Achenes always with pappus; outer and inner pappus scales usually distinctly shorter than middle ones. Outer involucral bracts ovate- lanceolate, not widened, subentire. Basal leaves lyrately pinnately parted, with ovate unpaired lobes, spiny-toothed; cauline leaves oval or oblong, pinnately parted, spiny toothed ..... 1. C. lanatus L. + Achenes usually without pappus, less often inner pappus scales longer than outer and middle. Outer involucral bracts lanceolate or oblong, widened above, with long yellowish spines along margin. Leaves oblong or oval, smooth, toothed with thin stramineous spines te MES tad dak naan Gece amen Me anbes, Jeep 3. C. oxyacanthus MB. 4. Plants wild; leaves distinctly spiny-toothed; florets light yellow .... wren Netwedicull wedi earl at ech d SMO M ls, ante 4. C. gypsicola Ijin + Plants cultivated or rarely wild; leaves short-spiny-toothed or subentire; florets red, orange-red, less often yellow ...................... sat dpa Mee al ails ad akacaeai dil sstezawinwade nym aeet el ui 5. C. tinctorius L. Section 1. Kentrophyllum (Neck.) Fiori and Paol. Fl. anal. Ital. III (1903-1904) 350.—Annuals or biennials. Plants woolly, arachnoid- hairy, glaucous, mealy, often ferruginous. Florets yellow or purple- pink. Pappus always well-developed, consisting of scales of various forms; outermost scales very short, sinuate, middle longer, acuminate, innermost as long as middle or shorter. Type of section: C. lanatus L. 1. C. lanatus L. Sp. pl. (1753) 830; Boiss. FI. or. III, 706; Schmalh. Fl. II, 131; Fedtsch. and Fler. Fl. Evrop. Ross. 1033; Grossh. Fl. Kavk. 58 aS 582 IV, 222: Pavlov, Fl. Tsentr. Kazakhst. II], 329.—C. tauricus MB. FI. taur.-cauc. II (1808) 285.—Kentrophyllum lanatum DC. Prodr. VI (1837) 610; Ldb. Fl. Ross. II, 714.—K. tauricum C.A.M. Verezeichn. (1831) 66.—Ic.: Sibth. and Sm. Fl. Gr. IX, tab. 841; Rchb. Ic. FI. Germ. XV, tab. 746. Annual, biennial. Plants to 1 m high, woolly, grayish-green, arach- noid- or almost arachnoid-hairy, especially above. Stem erect, solid, cymosely or divaricately branched above. Leaves glandular-glutinous; basal leaves somewhat stiff, lyrately pinnately parted; cauline leaves coriaceous, almost amplexicaul, oval or oblong, pinnately parted, spiny- toothed with conspicuous veins. Capitula solitary, at tips of on stem and lateral branches, to 3.5—4.0 mm cm long, 2.0—2.5 cm in dia. Outer involucral bracts leafy, spiny-toothed, ovate-lanceolate, as long as flo- rets or slightly longer; inner ones scarious, lanceolate, undivided or crenate-toothed from middle, spinescent, shorter than florets. Achenes 4-angled-oval, whitish or yellowish, almost lustrous, sometimes punc- tate or rugose above, to 5 mm long; outermost pappus scales very short, whitish, sinuate; middle ones 2.0—2.5 times as long as achenes, ochreous, serrulate-toothed, acuminate; innermost scales reduced, some- times (in plants from Soviet Central Asia) as long or almost as long as middle ones, whitish. Flowering July to August. Pasturelands, arable lands, and fields; common weed.—European Part: Bessarabia, Black Sea Region, Lower Don, Crimea; Caucasus: All regions, except alpine; Soviet Central Asia: Syr-Darya, Tien Shan, Pamiro-Alai, mountainous Turkmenia. General distribution: Central Europe (south), Mediterranean, Balkans-Asia Minor, Iran. Described from southern France. Type in London. Note. C. Janatus L. x C. oxyacanthus MB. = C. turkestanicus M. Pop. (7r. Uzb. Univ., Vyp. 14, 1941) is a known hybrid. According to M.G. Popov, it differs from C. Janatus L. by roundish, four-angled, pale, peripheral achenes without a pappus, with transverse grooves and an almost straight hilum. 2. C. glaucus MB. Tabl. prov. Casp. (1798) 58; Fl. taur.-cauc. II, 284; Boiss. Fl. or. III, 707; Schmalh. FI. II, 131; Fedtsch. and Fler. FI. Evrop. Ross. 1034; Grossh. Fl. Kavk. IV, 221.—Kentrophyllum glaucum Fisch. and Mey. ex DC. Prodr. VI (1837) 611; Ldb. FI. Ross. II, 713.—K. anatolicum Boiss. Diagn. pl. or. ser. I, 6 (1845) 113. Annual. Plants 30-80 cm high, glaucous, grayish-floccose. Stems erect, cymosely branched above. Leaves coriaceous, amplexicaul, oval or oblong-lanceolate, spiny-toothed. Capitula terminal, clustered, to 3 cm long, 1.5—2.0 cm in dia. Outer involucral bracts as long as florets, leafy, spiny-toothed; inner ones shorter than florets, scarious, lanceolate, 584 583 undivided, spinescent. Florets purple or purple-pink. Achenes 4-angled, oval, small, whitish, indistinctly transversely rugose above, to 4 mm long; pappus scales ochreous, outermost scales very short, sinuate; middle ones 2-3 times as long as achene, acuminate, serrulate-toothed, gradually thickened; inner scales reduced, fimbriate. Flowering June to September. Dry places and abandoned fields, common weed.—European Part: Crimea; Caucasus: all regions, except forest and alpine zones. Gen- eral distribution: Asia Minor, Iran. Described from Eastern Caucasus. Type in Leningrad. Section 2. Carthamus.—Annual plants. Stem glabrous, smooth, whitish, less often with uppermost ’ arachnoid-woolly especially above. Florets light yellow, orange-red, or red. Pappus lacking or consisting of serrulate-ciliate, unequal scales, shorter than achene; sometimes middle scales longer than achene. Type of section: lectotype of genus. 3. C. oxyacanthus MB. Tabl. prov. Casp. (1798) 108; FI. taur.- cauc. II, 283; DC. Prodr. VI, 612; Boiss. Fl. or. II, 709; Ldb. FI. Ross. II, 715; Grossh. Fl. Kavk. IV, 221; Pavlov, Fl. Tsentr. Kazakhst. Mi329. Annuals. Plants to 70 cm high. Stem straight, quite hard, divaricately branched from middle or above, smooth, whitish, glabrous below, pubescent above. Leaves sessile, weakly amplexicaul, some- what stiff oblong or oval, toothed, with stramineous thin spines, as long as width of leaf, spinescent, with prominent veins, glabrous or especially upper leaves grayish-pubescent, glandular; lower leaves of- ten pinnately parted. Capitula crowded, 2.5 cm long, 1.5 cm in dia. Outer involucral bracts whitish beneath, attenuate toward tip, leafy, with long, light yellow spines as long as florets; inner bracts scarious, lanceolate, undivided, spinescent, sometimes bracts arachnoid-hairy (var. arachnoideus Bge.). Florets light yellow. Achenes obovoid, 4 angled, truncate at apex, smooth, weakly lustrous, light yellow, spot- ted, black, to 6 mm long; pappus lacking or very rarely consisting of few serrulate-ciliate, unequal bristles, shorter than achene. Flowering June to August. Sandy places and fields, common weed.—Caucasus: Ciscaucasia (east), Eastern Transcaucasia; Soviet Central Asia: Tien Shan, Pamiro- Alai, mountainous Turkmenia. General distribution: Armenia and Kurdistan, Iran. Described from Eastern Ciscaucasia (Beshbarmak Mountain). Type in Leningrad. 586 584 4. C. gypsicola Iljin in Izv. Bot. Sada Akad. Nauk SSSR, XXX, 3-4 (1932) 357; Pavlov, Fl. Tsentr. Kazakhst. III, 330. Annual. Plant to 60 cm high. Stem simple or branched, especially above, glabrous, smooth, whitish. Leaves weakly amplexicaul, some- what stiff, lanceolate or oval, spiny-toothed, with prominent veins, entirely glabrous, somewhat lustrous. Capitula solitary, globose, 2 cm in dia. Outer involucral bracts leafy, spiny-toothed, in upper part al- most horizontally spreading, as long as florets; inner ones scarious, oval or lanceolate, yellow, at apex spiny, appressed, shorter than flo- rets, glandular. Florets light yellow. Achenes obovate or oblong-obo- vate, weakly compressed, weakly 4-angled above, smooth, lustrous, light yellow, speckled gray or speckled-black, to 6 mm long; pappus lacking or sometimes its bristles few, short and finely serrate-ciliate, shorter than achene, unequal. Flowering June to August. Always on gypsiferous clays and saline soils —Soviet Central Asia: Aralo-Caspian, Lake Balkhash Region (western); Caucasus: Southern Transcaucasia (Armenia, near border with Turkey). Endemic? Described from Turkey. Type in Leningrad. Note. This species, described by M.M. Iljin, was first reported by Schrenk from the specimens collected by Borschov from western Kazakhstan and by Schrenk himself from the Sary-Su River basin. Actually this species and C. oxyacanthus MB. are similar to each other in their general apearance, whitish stem, structure of the leaves, involucral bracts, achenes, and pappus. The two species, although readily distinguished in typical cases, were confused for a long time (cf. Iljin, op. cit.). It is also interesting that their ranges, at least within Soviet Central Asia, are sharply demarcated: C. gypsicola—endemic to the northern parts of Soviet Central Asia, is found nowhere south of 42°N. Lat., while C. oxyacanthus grows only in the southern parts of Soviet Central Asia and nowhere reaches north of 42°N. Here we have not considered the Armenian localities of C. gypsicola, which are very interesting but still inadequately studied. 5. C. tinctorius L. Sp. pl. (1753) 830; DC. Prodr. VI, 612; Ldb. Fl. Ross. II, 715; Boiss. Fl. or. III, 709; Fedtsch. and Fler. Fl. Evrop. Ross. 1033; Grossh. Fl. Kavk. IV, 221; Pavlov, Fl. Tsentr. Kazakhst. III, 329; Hanelt in Kulturpflanze, [X.—Ic.: Rchb. Ic. Fl. Germ. XV, tab. 746; Fedtsch. and Fler. op. cit. Fig. 1054. Annual. Plants to 1 m high. Stem erect, cymosely branched above, glabrous, sulcate, white. Leaves sessile, weakly amplexicaul, coria- ceous, oval or ovately lanceolate, short-spiny-toothed or subentire, awnless, glabrous, smooth, somewhat lustrous. Capitula solitary, ovate, 3 cm long, 2.5 cm in dia. Involucral bracts shorter than florets; outer 587 585 ones broadened toward base, scarious, yellowish-green, in upper part attenuate, green, broadly oval, short-spiny, outer most entirely recurved; inner ones scarious, ovate lanceolate, smooth, spinescent. Achenes obovate, 4-angled, truncate at apex, with prominent ribs, smooth, white, 4.5-7.0 mm long; pappus lacking or very rarely consisting of few, unequal, short, serrulate-ciliate bristles, shorter than achene. Flower- ing June to July. In cultivation, sometimes naturalizing in southern regions of Eu- ropean part of USSR (in Ukraine, Lower Volga, Urals), as well as in Caucasus and Soviet Central Asia. Described from southern France. Type in Leningrad. The following hybrids are known: C. lanatus L. x C. tinctorius L. (cf. Hanelt, loc. cit.). According to the data of Hanelt, almost all hybrids of these two species are sterile. Another hybrid—C. oxyacanthus MB. x C. tinctorius L.—has been reported by Popova (Popova in Zap. Leningr. S.-Kh. Inst. Nov. ed. 2 [1939] 127). Note. Unknown in the wild. According to recent information (Kult. FI. SSSR, VII, and according to Hanelt), two centers of diversification of forms of C. tinctorius are known: 1) India-Soviet Central Aisa and West Asia; and 2) Palestine-northeastern Africa. The first should be considered primary and the other secondary. At present, the plant is found in cultivation in many countries, from the Canary and Madeira islands to Tibet, China, and Japan, inclusively. There are also reports of its cultivation in America and Australia. Economic Importance. The only species of the genus of consid- erable economic importance. Carthamin is obtained from its flowers, which is used as a red dye for fabrics and without a mordant. Carthamin is a glucoside with the general formula C,,H,,O,,. Safflower was a valuable dying agent in ancient India, China, Egypt, and also in me- dieval times in Southern Europe, where the plant was introduced, apparently by Arabs. At present, the dying agent of safflower is used in the food industry as a substitute for saffron. Its seeds yield an oil which is closer in properties to the best of the sunflower and hemp oils. The oil is used in cooking, in industry for preparing non-yellow- ing white enamels, and in perfumery. 588 586 GENUS 1627. Cnicus L.'? LeSpi pli 53) 1826: Few peripheral florets in capitulum sterile, filiform, not conspicu- ous, with 2—3-fid corolla, rest bisexual with bilabiate, corolla, one of its lobes narrow, long, with one tooth, and other with 4 short teeth. Basal appendage of anthers scarious, long; stamen filaments densely papillate. Style branches short, obtuse, mostly oblong-ovate, with short papilliform hairs on outer side and ring of longer hairs at base. Pappus persistent, double, consisting of fewer elastic, somewhat stiff, bristles, longer in outer row and very short in inner one; achenes with toothed margin and numerous parallel raised ribs, with oblique hilum. A monotypic genus. 1. C..benedictus L. Sp. pl. (1753) 826; DC. Prodr. VI, 606; Ldb. Fl. Ross. II, 713; Schmalh. Fl. II, 130; O. and B. Fedtsch. Perech. Rast. Turk. IV, 278.—Centaurea benedicta L. Sp. pl. ed. 2 (1763) 1296.—Carbenia benedicta Benth. in Benth. and Hook. f. Gen. pl. II (1873) 482.—C. cnicus Winkl. in herb. Annual. Plant 5-50 cm high, branched from base, often lacking main stem, but with equally long, prostrate or ascending branches, bearing solitary terminal capitulum sometimes at ground level. Stem and branches densely hairy with long deciduous, beaded hairs; sticking together, mostly horizontally spreading, later less dense. Leaves green on both sides, with similar hairs, sometimes later becoming glabrous and with scattered sessile glands, pinnately parted into triangular or mostly oblong-triangular, acuminate lobes, lobes with spines along margin and notches separating them, somewhat stiff, fragile, with con- spicuous but thin veins; leaves sessile and decurrent on stem, lower ones strongly narrowed toward base into winged decurrent petiole; basal leaves petiolate, uppermost leaves forming leafy involucres around capitula and enveloping them. Outer involucral bracts oblong-ovate or oblong, with more or less thin spine at apex surrounded by long, flexu- ous, thin hairs, usually originating from beaded stalk; middle bracts longest, oblong or lanceolate, with relatively hard, long, pectinate, ochreous or brownish spine at apex, with similar hairs and often ad- ditionally with scarcely visible spinules; inner bracts slightly shorter, lanceolately linear, with scarious border, with scarcely developed pec- tinate appendage at apex. Corolla of peripheral sterile florets filiform, 'Treatment by M.M. Iljin. ?A Dioscoridean name of the plants, derived from the Greek word knixein—to scratch, injure; so named for its prickly habit. 589 587 pale; central florets bisexual, weakly yellowish, 13-17 mm long, with corolla tube 8—11 mm long and longer lobe (3.0—3.5 mm), shorter lobes 1.5—2.0 mm long. Pappus consisting of 9-12 hard, elastic, scarcely toothed, ochreous bristles 4-11 mm long; outer row bristles much shorter, covered with cobwebby hairs; achenes oblong-cylindrical, 7.0— 8.5 mm long, somewhat thick, hollow, lustrous, ochreous, or even dark-brown, with regular parallel ribs, and toothed edge above. Flow- ering May; fruiting June. Clayey deserts, on gravelly loams, loessic and clayey hills, waste- lands, old fields, garbage dumps, and crop fields——European Part: Black Sea Region (also cultivated throughout south of Ukraine); Caucasus: Eastern, Western, and Southern Transcaucasia; Soviet Cen- tral Asia: Lake Balkhash Region, Dzhungaria-Tarbagatai, Tien shan, Syr-Darya, Amu-Darya, mountainous Turkmenia, Kyzyl-Kum(?), Kara- Kum(?), Pamiro-Alai. General distribution: Mediterranean, Balkans- Asia Minor, Armenia and Kurdistan, Iran-Afghanistan, Indo- Himalayas(?), Dzhungaria-Kashgaria. Described from western Europe. Type in London. Economic Importance. A medicinal plant, entered into the pharmacopaea; its active principle is a bitter substance—carbobenedictin and cnicin (= Centrauran), used as an appetizer. Moreover, it contains tannins, some quantity of resin, gums, sterols, and glucosides. Its seeds contain 24-29% oil with specific gravity of 15/15—0.9262; the refrac- tive index is 1.4653 at 25°C; the saponification number is 191 and the iodine number is 141 (S.L. Ivanov). At the Ulyanovsk experimental field 26 centners of seeds were obtained from one hectare. Tribe 13. Mutisieae Cass. in Dict. Sc. nat. VIII (1817) 395, XX (1821) 379, XXXIII (1824) 462, LX (1830) 584; Opusc. phyt. I (1826) 327; Less. in Linnaea, V (1830) 131 and 237 p. p. excl. subtrib. Facelideae Less.: O. Hoffm. in Pflanzenfam. IV, 5 (1893) 333.—Trib. Carlineae Cass. in Dict. Sc. nat. VII (1817) 109 p. p., X (1818) 156 p. p., XX (1821) 357 p. p., XLVII (1827) 497 p. p., LX (1830) 569 p. p.; Opusc. phyt. I (1826) 292 p. p., II (1826) 202 p. p—Ordo Perdicieae Link, Handb. I (1829) 728.—Trib. Mutisiaceae Les. Synops. Comp. (1832) 92 p. p. excl. subtrib. Facelideae Less.: Benth. in Benth. and Hook. f. Gen. pl. II (1873) 214.—Trib. Mutisiees Baill. Hist. des. Plantes, VIII (1882) 69.—Fam. Mutisiaceae Bessey in Ann. Miss. Gard. II, 2 (1915) 164.—Capitula heterogamous or homogamous. Involucral 2-many rowed, involucral bracts obtuse, subobtuse, or acuminate. All florets bilabiate, bisexual, or peripheral ones pistillate, longer. Recep- tacle flat or slightly convex, locular or alveolate, mostly glabrous. Anther with short basal appendage. Style without tuft (collar) of pollen collecting hairs, bilobate. 590 588 Herbs (in USSR), shrubs, rarely trees, distributed mainly in America, particularly in South America; some genera are found in Africa and Asia. Subtribe 1. Mutisinae O. Hoffm. in Pflanzenfam. IV, 5 (1893) 340.—Subtrib. Mutisieae Less. in Linnaea, V (1830) 241 p. p., VI (1831) 98 and Synops. Comp. (1832) 93 p. p.—Subtrib Lerieae Less. in Linnaea, V (1830) 350 and Synops. Comp. (1832) 120.—Subtrib. Barnadesiae Benth. in Benth. and Hook. f. Gen. pl. II (1873) 215.— Subtrib. Onoserideae Benth. op. cit. 215 p. min. p.—Subtrib. Gerbereae Benth. op. cit. 217 p. p.—Subtrib. Nasauvieae Bebth. op. cit. 218 p. min. p.—cf. description of the tribe. KEY TO GENERA OF SUBTRIBE MUTISINAE* JoawiCapitwlavallyalikeveere pad. lekeerell tok Us seer ly af Oeil a + Capitula variable; in spring, heterogamous, in autumn, homoga- mous, with cleistogamous florets ..............:::eeeeees Leibnitzia Cass. 2. Peripheral florets with long limbs, surpassing 2—3-rowed involucral ad J DEI erties et aes Poneten! dap ee es coeaiia Gerbera Cass. + Peripheral florets with short limbs, shorter than many-rowed in- volucrelof imbricate bracts wens. eee. See Uechtritzia Freyn. GENUS 1628. Gerbera Cass.’ ? Cass. in Bull. Soc. Philom. Paris (1817) 34.—Arinica L. Sp. pl. (1753) 885 p. p. Capitula heterogamous. Involucre 1-, 2-, less often 3-rowed. Flo- rets with bilabiate, corolla; peripheral florets one-rowed, pistillate, with long limb surpassing involucre and usually brightly colored, and 2 short lobules of upper lobe rolled into ring; disk florets bisexual, very small, with corolla covered by pappus, its lobes almost equal in length, upper lobe with 2 lobules, lower one with 3 teeth, all coiled in ring; pollen tube very long exserted from corolla; anthers with filiform basal appendage, 1/3 as long as anther; style with flat, broadly elliptical lobes. Perennial herbs, with simple stems, each bearing single capitu- lum and only having basal leaves. *Constructed by E.G. Pobedimova. 'Treatment by E.G. Pobedimova. 2Named in honor of the 18th-century German naturalist, T. Gerber, who worked in Russia. 589 Type of genus: G. linnaei Cass. A genus with South African species for the most part. The number of species and their distribution in Asia is difficult to ascertain, since, after Cassini’s paper cited above, botanists included in the genus spe- cies that clearly do not belong to it (for example, G. schimperi Sch. Bip., in which the achenes have a long beak.). In its present circum- scription, the genus Gerbera is an aggregate genus needing mono- graphic treatment. Only one cultivated species is known in the USSR. G. jamesonii Bolus in Gard. Chron. I (1889) 772, fig. 122; Hook. f. in Bot. Mag. (1889) tab. 7087.—Ic.: Bolus, loc. cit.; Hook. f. loc. cit. Perennial. Rhizome short, with numerous roots. Leaves only basal, oblong, obtusely or acutely lobed with undulate lobes, unequal teeth, 13-25 cm long and 5-8 cm wide, petiolate; petioles 15—20 cm long. Floriferous shoots to 20 from each rhizome, 25—60 cm high, hard, glabrous. Capitula solitary, 7-15 cm in dia. Involucre campanulate, 7— 10 cm in dia, narrowed toward base; bracteal leaves lanceolate, ap- pressed. Peripheral florets one-whorled, about 30, with narrow, long limbs, with 3 teeth, yellowish in lower part, orange or flame-red above, with very short tube, upper lip [lobe] of corolla very short, bilobate, coiled into ring; disk florets small, with short, roundish ring of both corolla lobes. Achenes of peripheral florets and disk florets identical, terate, pubescent; pappus white, with scarcely toothed bristles. Flow- ering 2nd half of October to December. Caucasus: Eastern, Western and Southern Transcaucasia (in USSR known only in cultivation). Described from cultivated plants from Kew; seeds collected from South Africa (Transvaal). Type in Kew. Note. This species was introduced into cultivation long ago in gardens all over the world. In open ground, it can be cultivated only in the subtropics, and northwards—in green houses. It is one of the most beautiful ornamental plants, with the ray florets having very long, brightly and extremely diversely colored limbs. Often hybrids of this species are cultivated because they are more resistant (Chochua in 7r. Sukhum. Bot. Sada, VI [1951] 169). 592 590 GENUS 1629. Leibnitzia Cass." ” Cass. in Bull. Soc. Philom. Paris (1817) 34; in Dict. Sc. nat. XXV (1822) 421.—Anandria Less. in Linnaea, V (1830) 346; DC. Prodr. VII (1838) 40; Ldb. Fl. Ross. II (1846) 768; Turcz. in Bull. Soc. Nat. Mosc. XXI, 3 (1848) 87 (FI. baic.-dahur.).—Gerbera auct. non Cass.: Sch. Bip. in Flora, XXVII (1844) 782 p. p. quoad sect. Anandria Walp. Repert. VI (1846-1847) 316; Maxim. in Bull. Soc. Nat. Mosc. LV li Gl 879) 29. Capitula variable: in spring-heterogamous; in autumn—homoga- mous with cleistogamous florets. In spring capitula peripheral pistil- late florets, ligulate; disk florets smaller, bisexual with almost equal short lobes and projecting staminal tubes; style scarcely exserted from tube. In fall capitula all flerets bisexual, small with short, almost equal lobes, lower lobe broad with 3 teeth, and upper ones cut almost to base into linear lobes; staminal tube scarcely visible at throat of flo- rets; style included in staminal tube, less often capitula consisting of florets with shortened lobes. Perennial herbs with rosettes of basal leaves, small during spring flowering and large during autumn floweing time. Type of genus: L. anandria (L.) Turcz. Besides the two species reported here for the flora of the USSR, this genus includes the Himalayan species, L. kuntzeana (A. Br. and Aschers.) Pobed. comb. nova (=Gerbera kuntzeana A. Br. and Aschers. in Cat. sem. hort. Berol. [1871] App. 3), and the southern Chinese L. ruficoma (Franch.) Pobed. comb. nova (=Gerbera ruficoma Franch. in Morot. Journ. Bot. II [1 March 1888] 68). 1. Plant 15-30 cm high; leaves large, lyrate, (6)9—10(13) cm long, (2.0)2.5—3.0(4.0) cm wide, with roundish, sometimes broadened terminal lobe and 3-4 pairs of lateral lobes somewhat smaller than terminal lobe, glabrous or weakly arachnoid-hairy beneath, less often floccose. Disk florets with staminal tube included in corolla ....... Aided nel utes phenyl dese 2. L. knorringiana (B. Fedtsch.) Pobed. + Plant 5—18 cm high; leaves small, weakly lyrate, (1.5)2.0-5.0(7.0) cm long, (0.7)1.5—2.5(2.8) cm wide, with ovate or oblong terminal lobe, and 2-3 pairs of small lateral lobes several times smaller than terminal lobe, white-tomentose beneath. Disk florets with staminal tube greatly-exserted from corolla’............ 1. L. anandrai (L.) Turcz. 'Treatment by E.G. Pobedimova. 2Named in honor of Leibnitz, an eminent 17th-century scientist and philosopher. >These species have been compared only in an early stage of development, be- cause there are no complete specimens in the herbarium. Soil Plate XXVIII. 5)! Leibnitzia 2 and Schmalh.) Pobed; 1—UVechtritizia kokanica (Rgl. knorringiana (B. Fedtsch.) Pobed. 593 592 1. L. anandria (L.) Turcz. in Scheglov', Ukaz. Otkryt. VIII, I, 3 (1831) 404; Bess. in Flora, XVII, 1, 17; Nakai in Journ. Jap. Bot. XIII, 852; Kitagawa, Lineam. Fl. Mansh. 456; Kryl. Fl. Zap. Sib. XI, 2964.—Tussilago anandria L. Sp. pl. (1753) 865.—T. lyrata Willd. Sp. pl. III, 3 (1803) 1936.—Perdicium anandraia R. Br. in Ait. Hort. Kew. ed. 2, V (1813) 84.—P. tomentosum Thunb. FI. Jap. (1784) 920 (sec. Less.).—Leibnitzia cryptogama Cass. in Dict. Sc. nat. XXV (1822) 422.—L. phaenogama Cass. ibid.—Chaptalia anandria Spreng. Syst. veg. 3 (1826) 504; Ldb. FI. alt. IV, 89.—C. lyrata Spreng. ibid.— Anandria radiata Less. in Linnaea, V (1830) 241 and 346.—4A. discoidea Less. ibid. 347.—A. bellidiastrum DC. Prodr. VII (1838) 40; Maxim. Prim. Fl. Amur. 176; Turcz. in Bull. Soc. Nat. Mosc. XXI, 3, 87(Fl. baic.-dahur.).—A. bellidiastrum a. vernale Turcz. in DC. loc. cit—A. bellidiastrum 8. autumnale Turcz. in DC. loc. cit.—A. dimorpha Turcz. in Bull. Soc. Nat. Mosc. XI (1838) 95 (Cat. pl. baic.- dahur. No. 695); Ldb. Fl. Ross. I, 768.—Gerbera anandria Sch. Bip. in Flora, XX VII (1844) 782; Walp. Repert. VI, 316; Kom Fl. Manchzh. III, 762; Kom. and Alis. Opred. Rast. Dalnevost. Kr. II, 1086.—Ic.: Pobedimova in Bot. Zhurn. SSSR, XXX, 5 (1945) 220, 221, and 223.— Exs.: GRF Nos. 1931, 1932, and 3499. Perennial. Rhizome very short, thick, brown, with large fascicle of string-like, fleshy, light brown roots. Plants with two growth stages— spring and autumn. Leaves in the spring stage ovate or oblong-ovate, less often weakly lyrate, cordate, (1.5)—2.0—4.0(5.0) cm long and (0.7)1.5—2.5(2.8) cm wide, green and weakly arachnoid hairy above, white-tomentose beneath, almost entire, with short, thick, inconspicu- ous cusp or irregularly toothed, with more frequent and larger cusp; petioles (1.0)1.5—3.0(3.5) cm long, white-tomentose. Scapes 1, less of- ten 2, (5.0)5.5—15.0(18.0) cm long, simple, erect, each bearing single capitulum, tomentose, developing simultaneously with bracts 3—5 mm long and 0.25—0.5 mm wide, less often distant. Capitula (0.6) 1.0— 1.5(2.8) cm wide. Involucre 2-rowed; outer bracts short, narrow, lin- ear; inner ones somewhat broader; all bracts obtuse, reddish-violet along margin, more or less floccose, peripheral florets pistillate, bila- biate, with short, initially white, later pinkish ligules at end of flow- ering time, with 1—2—3 unequal teeth at apex, upper lip in form of 2 short teeth; staminal tube included in corolla; anthers with smaller quantity of pollen, with 2 acute, short, basal appendages; style ex- serted from staminal tube, bifid, its lobes papillose; disk florets with both lips coiled in ring; staminal tube long, exserted from corolla; 'The almost non-existent volumes of this publication are preserved only in the Public Library, Leningrad. 593 594 anthers also almost lacking pollen; ovary covered with short, upward- spreading, appressed hairs. Mature achenes undeveloped in spring form. Flowering second half of April to June. Leaves in the fall stage lyrate, (5)9—15(18) cm long and (3.0)3.5— 5.5(6.5) cm wide, terminal lobe broad, ovate, acute, scarcely cordate and with 3—4 pairs of smaller lobes, pale green beneath, glabrous or weakly arachnoid-hairy, less often whitish-pubescent; petiole weakly winged, arachnoid-hairy, (1.0)3.5—7.5(10.5) cm long. Scapes one or many, 25—60 cm high, with single capitulum, floccose-tomentose, with numerous, linear bracts (9)13—30(40) mm long, 0.5—1.0 mm wide. Ca- pitula 2.5-4.0 cm in dia, opening only at fruiting. Involucral bracts narrower than in spring capitula. All florets bisexual with small co- rolla shorter than pappus bristles; its lower lobe short, with 3 obtuse teeth, upper lobe bifid; staminal tube inserted in corolla; anthers with abundant pollen, basal appendages of anthers short. Achenes on short stalk, oblong, 5—6 mm long, 0.75—1.0 mm wide, with short beak and prominent ribs, pubescent, especially in upper part; pappus longer than achene, with numerous toothed bristles connate at base in ring. Flow- ering July to September. Dry open mountain slopes, grassy patches, in mixed and oak for- ests.—Eastern Siberia: Angara-Sayans, Dauria; Far East: Zeya-Bureya, Ussuri, Sakhalin. General distribution: Mongolia, China, Japan, Ko- rean Peninsula. Described from Siberia. Type in London. 2. L. knorringiana (B. Fedtsch.) Pobed. comb. nova.—Gerbera knorringiana B. Fedtsch. in Izv. Bot. Sada, XV, 1 (1915) 3; O. and B. Fedtsch. Perech. Rast. Turk. VI, 346. Perennial. Rhizome short, woody, brown, with light brown fleshy, more or less slender roots, in upper part covered with remnants of previous years’ petioles of leaves. Leaves only basal, lyrate, large (6)9—10(13) cm long, (2.0)2.5—3.0(4.0) cm wide, terminal lobe almost round or scarcely broadened, slightly longer than 3—4 pairs of lower lobes, gradually decreasing toward leaf base, dark green and glabrous above, pale, glabrous or finely arachnoid-hairy beneath, less often floc- cose, with large, irregular teeth and short obtuse cusp; petiole 3—7 cm long, often winged in upper part; scapes 1 or 3, 15—30 cm high, floc- cose, especially densely below capitula; bracteal leaves small, scat- tered. Capitula one on each scape, narrow, (7)10—12(15) cm in dia, involucre 2—3-rowed; involucral bracts: outer short, lanceolate, ob- tuse, inner almost twice as long as outer; all glabrous, sometimes vio- let along margin and at apex; receptacle locular, scarcely convex; pe- ripheral pistillate florets with 3 toothed lower tip (ligule), with upper tip biparted into narrowly linear short lobes; style bilabiate, ciliate, 595 594 exserted from corolla; disk florets with almost equal lobes, lower lobe broader with 3 large teeth, upper one not divided up to base, with weakly coiled labellae; staminal tube included in floret; style not ex- serted or with small linear, appressed basal appendages, stamen fila- ment shorter than anthers, attached in upper third of tube; ovary with short appressed erect hairs; mature achenes unknown. Flowering May to June. (Plate XXVIII, Fig. 2). High-mountain spruce forests (under canopy of spruce).—Soviet Central Asia: Tien Shan. Endemic. Described from western Tien Shan (Lake Sary-Chilek). Type in Leningrad. Note. This species is known only in the flowering stage. It was collected in August 1945 in the fall stage by V. Vipper in Arkit, Karagatun-Sai in a spruce-alder forest under. a canopy of spruce, at 1,700 m, unfortunately, only in the vegetative state with longer leaves than im the spring form (cf. Plate XXVII, Fig. 2). The absence of capitula on these plants does not permit us to resolve definitively whether this species belongs to the genus Leibnitzia; however, the great similarity in the structure of the capitula with L. anandria forces us to include it in this genus. GENUS 1630. Uechtritzia Freyn" * Freyn in Oest. Bot. Zeitschr. XLII (1892) 240.—Gerbera §3. Uechtritzia Beauverd in Bull. Soc. Bot. Geneva, ser. 2, II (1910) 43. Capitula homogamous. Involucre subglobose; involucral bracts many-rowed, imbricate, usually white-tomentose. All florets bisexual; peripheral florets one-rowed, with slightly longer, 3-toothed, lower and narrow, bifid rounded ring of upper lobe of corolla; anthers with short appendages at base and with very little pollen; style bifid, with flat elliptical lobes; disk florets slightly smaller, all corolla lobes coiled in ring; staminal tube exserted from corolla, style scarcely visible. Achenes lacking beak, oblong, more or less pubescent. Perennial herbs, with simple stems bearing single capitulum and only basal leaves. Type of genus: U. armena Freyn. Freyn described this genus as monotypic. However, after studying the Mediterranean species “Gerbera kokanica,” we were convinced that it also belongs to the genus Uechtritzia on the basis of the struc- ture of the capitulum and the bisexual florets, and the subglobose invclucre with imbricate involucral bracts, which is not characteristic 'Treatment by E.G. Pobedimova. *Named in honor of the Polish botanist Rudolph Uechtritz. 596 597 505 of the genus Gerbera. Moreover, apparently, even the ranges of these genera are well separated: Gerbera is tropical whereas Uechtritzia is more northern. However, until the species composition of the genus Gerbera is established, it also is not possible to define more precisely its overall range. 1. Leaves ovate, linear-ovate, or broadly ovate, obtuse, cordate, al- most entire or very rarely with remote, scarcely visible, obtuse fee clay herent teem eor an MUA eeenet Wed, MENRED. Seapets 1. U. armena Freyn + Leaves lyrate, linear-obovate; terminal lobe large, ovate, obtuse, lateral lobes 3-4 pairs, smaller, almost round or weakly elongated ck cers Rea Re 2. U. kokanica (Rgl. and Schmah.) Pobed. 1. U. armena Freyn in Oest. bot. Zeitschr. XLII (1892) 240— 242.—Gerbera armena Beauverd in Bull. Soc. Bot. Geneve, ser. 2, II (1910) 43.—Exs.: Herb. Fl. cauc. No. 50. Perennial. Rhizome thick, woody, brown, covered with brown scaly leaves, at apex with remnants of petioles of previous year’s leaves, with thick string-like light brown roots. All leaves in basal rosette ovate, linearly or broadly ovate, (7.5)10.0—11.0(12.5) cm long, (6.0)6.5— 8.0(8.5) cm wide, obtuse, cordate, almost entire or with occasional, irregularly spaced, inconspicuous, obtuse teeth, green above, glabrous, lustrous, with very distinct veins or irregularly weakly floccose-tomen- tose, densely tomentose beneath, long-petiolate; petiole 8-11 cm long, more often weakly winged, floccose-white-tomentose. Scapes usually 1, less often 2, 55-70 cm high, one-headed, straight, simple, floccose- tomentose, densely pubescent below capitulum, leafless, occasionally in upper half with remote, very small, 4-6 mm long, 0.5 mm wide, linear bracteal leaves, green above, white-tomentose beneath. Capitula (4.0)4.5—5.0(5.5) cm in dia. Involucre many-rowed, imbricate; outer involucral bracts lanceolate, acuminate, white-tomentose on outer side, fluff, readily detaching and exposed upper part reddish, inner bracts oblong, obtuse, abruptly acuminate; all spinescent, with thick reddish glands on more especially in upper half. Receptacle flat, alveolate, with filiform lacerate scales on edges of alveoli. Peripheral florets bilabiate, with erect, broad lower lobe (ligule) with 3 apical teeth and upper lobes bifid in thin lobules linear, coiled in ring, and clavate; staminal tube short with style strongly exserted, bifid, papillate; in disk florets corolla lobes variable, all lobes coiled in ring; staminal tube very long, strongly exserted from corolla, stamen filaments at- tached in middle of corolla tube, shorter than anthers; basal appendage of anthers linear, longer, toothed; style bifid, slightly exserted from staminal tube. Achenes 7 mm long, 2 mm wide, oblong, terete, brown, 598 596 densely covered with short appressed hairs; pappus very long, 12 mm long, with numerous scarcely toothed bristles connate at base in ring. Flowering July to August. Pine forests on mountain slopes. So far not found in USSR.— General distribution: Armenia and Kurdistan (former Kars Region). Described from Eastern Anatolia (Sapikor-Dag Mountain). Type in Brno; isotype in Leningrad. 2. U. kokanica (Rgl. and Schmalh.) Pobed. comb. nova.—Ger- bera Kokanica Rgl. and Schmalh. in Izv. Obshch. Lyubit. Estestv. Antrop. i Etsn. 34, 2 (1882) 53; O. and B. Fedtsch. Perech. Rast. Turk, IV, 304. Perennial. Rhizome thick, woody, dark brown, covered at apex with brown remnants of petioles of previous year’s leaves, with thick, string-like, light brown roots. All leaves in basal rosette, oblong-obo- vate, (7)12—15(20) cm long, (3)5.0—7.5(8.5) cm wide, lyrate, lustrous and glabrous above, densely white-tomentose beneath, irregularly toothed with distant prickles and very short spines; terminal lobe ovate, obtuse, large; lateral lobes 3—4 pairs, almost round or weakly attenu- ate; petiole short, 2-7 cm long, more or less winged; scape one, less often 2, (20)25-50(65) cm high, terminating in single capitulum, straight, simple, floccose-tomentose, densely white-tomentose below capitulum; bracteal leaves uniformly distributed on scape, remote, erect, very small, (3)5(7) mm long, 0.25—0.5 mm wide, lanceolate-linear, acute, reddish-green above, white-tomentose beneath. Capitula (2.8)4.0— 5.0(6.0) cm in dia. Involucre many-rowed, imbricate; involucral bracts lanceolate, gradually acuminate, white-tomentose on outer side, gla- brous inside, green, with reddish-lilac margin, and occasional thick, short glands. Receptacle flat, scarcely convex, locular. All florets bi- labiate, bisexual; in peripheral florets lower lobe short, erect, with 3 teeth, not longer than involucre; upper one bifid with narrowly linear lobules coiled in ring, clavately thickened; staminal tube short, usually with little pollen; appendages at base of anthers short; style long, strongly exserted from corolla, bifid, with short, flat, oval lobes papil- late; in disk florets both lobes coiled in ring; staminal tube very long, exserted from corolla, pinkish-lilac, with normally developed pollen; appendage at base of anthers very long, sometimes longer than an- thers, linear, weakly toothed; stamen filaments, not longer than stami- nal tube, inserted in lower third of corolla; style scarcely exserted from staminal tube. Achenes 4.5 mm long, 1.5—2.0 mm wide, oblong, cylindrical, weakly compressed laterally, densely white-pubescent with long silkish, erect hairs, lacking beak; pappus much longer than achene, 597 with tiny and short-toothed appressed bristles, connate at base in ring. Flowering July to September (Plate OMIM, Fig:"1). Rocks, on mountain slopes in woody-shrubby communities.—So- viet Central Asia: Tien Shan, Pamiro-Alai. Endemic. Described from Shakhimardan. Type in Leningrad. 599 ADDENDA XXVII DIAGNOSES PLANTARUM NOVARUM IN TOMO XXVIII FLORAE URSS COMMEMORATARUM Diagnosis of New Species Mentioned in Twenty-eighth Volume June 1963 CARDUUS L. 1. C. hajastanicus Tamamsch. sp. nova (sect. Carduus). Planta 50-100 cm alt., glaucescenti-viridis; caulis a basi divaricato- ramosus, interrupte alatus, alis spinosis, parce araneoso-tomentosus; folia anguste oblanceolata, sessilia, non profunde et inaequaliter pinnatilobata; lobi triangulares, in spinam flavam longam 8—9 mm lg. attenuat. Pedunculi monocephali; calathidia solitaria, 3-3.5 cm lg., 2.5—3 cm It.; involucrum subglobosum, basi impressum; involucri phylla inaequilonga, basi dilatata et appressa, subglabra, externa et media in acumen robustum spinosum attenuata, externa reflexa, interna anguste lanceolata, purpurea; flores atro-purpurei vel rubescentes, involucri phyllis longiores; corolla 19—19.3 cm lg. Habitat in declivibus lapidosis et herbidis. Typus. Armenia, ad lac. Sevan, prope p. Tzamakabert, 19 VIII 1953, S. Tamamschjan. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur. Affinitas. C. onopordioidi affinis, sed caule divaricato-ramosissimo, pedunculo vix nutante, calathidiis plus minusve elongatis (non globosis), floribus atro-purpureis (non roseis) valde differt. 2. C. furiosus Tamamsch. sp. nova (sect. Carduus). Planta valde spinosa, 30-35 cm alt., lucide viridis; caulis solitarius, simplex, dense foliosus; folia vix coriacea, spinosissima, pinnatilobata; lobi dentati, spinosi, spina flava robusta valde pungente terminati; folia caulina supra lucida, subtus atro-virentia, vix et partim perplexe araneoso-tomentosa. Calathidia subglobosa, 3—3.5 cm lg., 3—4 cm It.; involucrum globosum, subglabrum vel vix araneosum; involucri phylla pappo et corolla longiora, exteriora mediaque robusta, basi plus minusve lata, in acumen spinosissimum subito attenuata, reflexa; corolla rosea, 22-23 mm lg. 600 599 Habitat in lapidosis siccis. Typus. Irania, Badalan. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur. Affinitas. C. onopordioidi affinis est, sed calathidiis subglobosis, involucri phyllorum forma sat bene differt. 3. Sect. Carduastrum Tamamsch. sect. nova. Calathidia magna vel mediocria, subglobosa. Involucri phylla sem- per angusta, conspicue vel inconspicue appresse imbricata, plus minusve herbacea vel lignosa, rigida, apice valde pungentia. Pedunculi longissimi, usque ad apicem alati, alis spinosis, vel exalati, non spinosi, griseo-lanati. Typus sectionis: C. crispus L. 4. C. nikitinii Tamamsch. sp. nova (sect. Stenocephalus Rouy). Planta 5—15(18) cm alt.; caulis vix costatus, usque ad calathidia spinulosus et dense foliatus, simplex vel rarissime in parte superiore parce ramosus. Pedunculi albo-lanati; calathidia cylindrica, parva, pauciflora (flores in numero 8-9); involucri phylia subglabra, virescentia, exteriora late ovata, apice membranaceo-marginata, acuminata, in aculeolum rectum transeuntia, media anguste ovata, in- tima lineari-lanceolata, membranacea, semper apice fusco-violacea; pappus sordide albus, corolla et achenio longior. Habitat in declivibus siccis et schistosis in Turcomania. Typus. Turcomania, montes Kopetdagh orientalis, Kaachka, 27 IV 1957. leg. V. Nikitin et Isczenko. In Herb. Inst. Bot. Acad. SC. URSS (Leningrad) conservatur. Affinitas. A C. pycnocephalo, qui in Europa meridionali habitat, involucri phyllorum forma, pubescentia et aculeolo non reflexo differt; a C. arabico, C. beckeriano et C. cinereo involucri phyllis habituque valde distinguitur. CIRSIUM Miil. 5. Sect. Caucasigena Charadze sect. nova.—Calathidia nutantia. Involucrum plus minusve arachnoideo-pubescens; involucri phylla in- tegra vel aculeolato-ciliata, apice in aculeum elongatum attenuata; involucri phyllorum exteriorum aculei reflexi vel patentes. Flores rubri, rarius albi; corollae tubus angustus parti dilatatae subaequilongus vel eam superans. Folia supra aculeolato-arachnoidea. Typus sectionis: C. caucasicum (Adams) Petrak. 601 600 6. C. gagnidzei Charadze sp. nova (sect. Caucasigena Charadze). Perennis; caules ascendentes, ad 1 m alt., subcostati sulcatique, purpurei, superne breviter ramosi, ad apicem foliosi; folia oblonga, supra viridia, dense minute setosa, subtus glaucescentia, plus minusve arachnoideo- pilosa, nervis prominentibus subglabris; folia inferiora in petiolum inciso-alatum margine aculeatum angustata, ad nervum medium inciso- pinnatipartita, segmentis ovato-triangularibus bipartitis margine aculeolatis, lobis dentibusque in aculeum luteolum 5—7 mm lg. acutatis; folia caulina sessilia, basi auriculato-dilatata, semiamplexicaulia, 15 cm lg., 7 cm It., ad medium inciso-pinnatipartita, segmentis ovato- triangularibus bipartitis, lobis in aculeum firmum acutatis; folia superiora diminuta, inciso-pinnatipartita, suprema sub calathidiis solitaria, ea paulum superantia. Calathidia subsessilia, in numero 2—3 ad apices ramorum congesta vel solitaria, oblonga, denique ovato- globosa, inflorescentiam corymbosam formantia; involucrum 10-12 mm in diam., inferne leviter superne dense arachnoideo-pilosum; involucri phylla virescentia, lanceolata, carinata, a medio in aculeum tenuem luteolum erectum vel patentem attenuata, externa parva, glabra, media elongata, arachnoideo-pilosa, interna lineari-lanceolata, aculeo elongato filiformi falcato; flores purpurei; corolla 20 mm lg., parte dilatata fere ad medium dissecta; tubus angustus partem dilatatam subaequans; pappus sordide albus, pilis internis apice scabris elongatis. Habitat in regione subalpina, in pratis altiherbosis Caucasi occidentalis. Typus. Transcaucasia occidentalis, systema fluminis Mzymta ad fontes fl. Laura, 17 VII 1962, R. Gagnidze. In Herb. Inst. Bot. Acad. Sc. RSS Georgicae (Tbilissi) conservatur. Affinitas. A speciebus proximis calathidiis parvis et lobis foliorum inferiorum latis differt. 7. C. longiflorum Charadze sp. nova (sect. Caucasigena Charadze). Perennis; caules erecti, ad 150 cm alt., a medio plus minusve ramosi, ad apicem foliosi, subsulcati, purpureo-colorati, sub- glaucescentes, superne sparse araneoso-pilosi; folia ambitu elliptica vel oblongo-elliptica, a caulinis inferioribus apicem versus sensim diminuta, 4-17 cm lg., 7—1.5 cm It., ultra medium profunde late inciso- pinnatipartita, segmentis elongatis lanceolatis vel triangulari-lanceolatis 2—3(5)-partitis, laciniis apice in spinam firmam 5-10 mm lg. luteoloalbidam acutatis; folia supra viridia, setis aculeatis luteolo-albidis et pilis minutissimis sparsis obsita, subtus dense griseo-tomentosa, nervis prominentibus tenuiter pubescentibus; folia suprema sub calathidiis 2-3, parva, lanceolata, involucro breviora vel ei aequilonga. Calathidia numerosa, plus minusve nutantia, breviter pedunculata, 602 601 oblonga, plerumque 1.5—2 cm lg., cum floribus 3—3.5 cm lg., in numero 2-3 congesta vel solitaria, inflorescentiam laxam corymboso- paniculatam formantia; involucrum subglabrum, sparse araneosum, plerumque atro-purpureum; involucri phylla lanceolata, navicularia, dense imbricata, dorso carinata, apice in aculeum robustum luteolum patentem attenuata, margine minutissime ciliata, scabra, interna lineari- lanceolata, membranacea, in acumen tenue elongatum fusco-coloratum falcatum attenuata, margine scabra, minutissime acute serrata; flores purpurei; corolla 20—22 mm lg., parte dilatata ad medium quinquepartita; tubus angustus, partem dilatatam subaequans; pappus 16 mm lg, sordide albus, pilis internis apice leviter clavato-dilatatis serrulatis; achenium 4.5 mm lg., 2.3 mm lt. Habitat in regione montana media, in declivibus australibus Caucasi Magni. Typus. Transcaucasia occidentalis, systema fluminis Mzymta, in viciniis <>, in pratis, 29 VIII 1962, A. Charadze. In Herb. Inst. Bot. Acad. Sc. RSS Georgicae (Tbilissi) conservatur. Affinitas. A C. gagnidzei foliis profunde pinnatis, segmentis angustis et eclethidiis elongatis numerosis differt. 8. C. ezerkessicum Charadze sp. nova (sect. Caucasigena Charadze). Perennis; caules elati, ad 2 m alt., ascendentes, a medio ramosi, superne atro-purpurei, subsulcati, ad apicem foliosi, subglabri, pilis sparsis flexuosis adpressis; ramuli erecti, subpatentes; folia supra plus minusve dense adpresse aculeolato-setosa, ad nervum crispato-pilosa, subtus pallide viridia, sparse arachnoideo-pilosa vel tenuiter griseo- tomentosa, caulina sessilia, utrinque attenuata, inferiora auriculato- dilatata, semiamplexicaulia, oblongo-elliptica, 25 cm lg., 10 cm It., ultra medium inciso-pinnatilobata, segmentis remotis triangulari-ovatis 2—3(5)-lobatis margine aculeolato-ciliatis, lobis in aculeum tenuem 4-8 cm lg., acutatis; folia cetera inferioribus similia, apicem versus sensim diminuta, media circa 17 cm lg., 8 cm It., inferiora oblongo- lanceolata, suprema sub calathidiis 4-5 cm lg., ea vix superantia. Calathidia haud numerosa, ovato-oblonga, 15—22 mm lg., cum floribus ad 3 cm lg., nutantia, terminalia solitaria vel bina, breviter pedunculata, axillaria valde diminuta, inflorescentiam breviter corymboso- paniculatam formantia; involucrum inferne glabrum, superne subarachnoideum; involucri phylla externa oblonga, concava, dorso subcarinata, luteolo-viridia, rarius purpurea, in apicem breviter aculeatum subfalcatum reflexum acutata, margine plus minusve dense aculeolato-ciliata, aculeolis erectis luteolis, interna membranacea, lineari-lanceolata, in mucronem tenuem fere filiformem reflexum 603 602 acutata, margine minute arguteque serrata, scabra, superne sub- arachnoidea vel glabra; flores lilacino-rubri; corolla ad 22 mm lg., tubo angusto ad 10 mm lg., parte dilatata ad 1/3 dissecta; pappus sordide albus, pilis internis fere ad apicem pinnatis, apice subdilatatis scabris. Habitat in regione subalpina, in partis althiherbosis. Typus. Caucasus occidentalis, Guzeripl, trajectus <>, in declivitate m. Abago, 2000 ms. m., 12 VIII 1962, R. Gagnidze. In Herb. Inst. Bot. Acad. Sc. RSS Georgicae (Tbilissi) conservatur. Affinitas. A C. gagnidzei Charadze et C. longifloro Charadze involucri phyllis externis margine aculeolato-ciliatis, apice in aculeum breviorem acutatis differt. 9. C. Buschianum Charadze sp. nova (sect. Caucasigena Charadze). Perennis; caules ad 1 m alt., subsulcati, araneoso-pubescenter, ascendentes, superne ramosi; folia inferiora mediaque magna, ad 30 cm lg., 13—-15(16) cm It., late elliptica, basi angustata, caulina basi auriculato-dilatata, semiamplexicaulia, apicem versus angustata, acutata, ad medium pinnatipartita, segmentis ovatis rarius bipartitis vix sursum directis, dentibus apice aculeatis, aculeis 2-4 mm lg.; folia supra setis adpressis dense obsita, subtus plus minusve dense albo-tomentoso- pilosa, caulina superiora abbreviata, 4.5—9.5 cm lg., 2-5 cm It., suprema sub calathidiis linearia. Calathidia terminalia nutantia, ad apicem ramorum elongatorum in numero 2-3, inflorescentiam laxam corymbosam formantia; involucrum inferne adpresse arachnoideo- pilosum, superne laxe araneosum, ad 4.5 cm It.; involucri phylla externa valde abbreviata, media lanceolata, falcata, reflexa, plus minusve dense aculeato-ciliata, in aculeum brevem rarius longum firmum lutescentem reflexum angustata, interna lineari-lanceolata, membranacea, in acu- men elongatum aculeatum reflexum angustata; flores lilacino-rubri; corolla (23)25—27 mm lg., parte dilatata ad 17 mm lg.; pappus sordide albus, pilis internis apice subdilatatis scabris; achenia ad 5 mm lg. Habitat in regione subalpina Caucasi orientalis. Typus. Ossetia meridionalis, prope p. Ermani, Delwars, 2000 m s. m., 10 IX 1935, E. et N. Busch. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur. Affinitas. A speciebus proximis ex affinitate C. ossetici (Adams) Petrak, foliis majoribus et latioribus et involucri phyllis externis valde abbreviatis falcatis fere tomentoso-pilosis differt. 10. Subsect. Armata Charadze. subsect. nova (sect. Epitrachys DC.).—Calathidia plerumque magna, rarius mediocria, erecta, foliis 604 603 supremis plus minusve elongatis obvallata, solitaria, in numero 2-3 congesta vel inflorescentiam racemosam formantia. Involucrum plerumque dense araneoso-pilosum vel tenuiter araneosum; involucri phylla integra, levia vel scabra, rarius margine sparse aculeolata, plerumque a medio in aculeum elongatum erectum vel patentem attenuata. Typus subsectionis: C. Jappaceum (MB.) MB. 11. C. megricum Charadze sp. nova (sect. Epitrachys DC.). Perennis; caules 30-200 cm alt.; simplices, ascendentes, albo- tomentoso-pilosi, ad apicem foliosi; folia elongato-oblonga, supra aculeolis minutis plus minusve dense obsita et tenuiter arachnoidea, subtus albo-tomentoso-pilosa; folia caulina inferiora profunde pinnatisecta, segmentis elongato-lanceolatis acuminatis, basi in petiolum alatum dentatum plus minusve longum attenuatis, inaequaliter bipartitis, lobis dentibusque in aculeum firmum 10-15 mm lg. acutatis, caulina superiora sesilia, semiamplexicaulia, profunde pinnatipartita. Calathidia terminalia axillariaque subsessilia, inflorescentiam brevem racemosam formantia; involucrum subglabrum vel araneoso-pilosum, 1.2—2 cm It.; involucri phylla externa linearia, virescentia, cetera membranacea, ab ovatis ad lanceolata, in acumen subfalcatum reflexum in aculeum gracilem attenuatum abrupte acutata, inferne glabra, superne araneoso- pilosa, margine subscabra; flores rosei; pappus luteolo-albidus; achenia fusco-colorata, matura ignota. Habitat in regione subalpina Transcaucasiae australis, in pratis. Typus. Armenia, distr. Megri, supra pag. Gjalur, in pascuis, 2300— 2400 ms. m., 8 VIII 1932, I. Karjagin (sub. C. isophyllo (Petrak) Grossh. x x C. schelkownikowii Petrak). In Herb. Inst. Bot. Acad. Sc. RSS Azerbaidzhaniae (Baku) conservatur. Affinitas. A C. anatolico (Petrak) Grossh. calathidiis confertis et segmentis foliorum latioribus differt; a C. schelkownikowii Petrak calathidiis parvis, indumento adpresso, segmentis foliorum acuminatis differt. 12. Sect. Lamyropsis Charadze sect. nova.—Calathidia breviter pedunculata. Involucri phylla externa apice triangulari-lanceolata, membranacea, attenuata, in apicem sensim angustata, apice partem dilatatam plerumque 2-3 plosuperante, interna externa aequantia vel eis breviora. Pappus e pilis internis gracilibus fragilibus fere ad apicem pinnatis in acumen filiforme attenuatis constans. Planta perennis, foliis plus minusve coriaceis, supra levibus, subglabris, subtus albo- tomentosis, segmentis lateralibus reductis, in aculeum firmum acutatis. Typus sectionis: C. sinuatum (Trautv.) Boiss. 605 604 13. C. badakhschanicum Charadze sp. nova (sect. Echenais (Cass.) Petrak). Perennis; caules ad 1 m alt., ascendentes, sulcati, inferne sparse crispatopilosi, superne lanuginosi, ramosi, ramis axillaribus plus minusve elongatis; folia caulina glaucescentia, supra sparse pilosa, subtus secundum nervos primarios lanato-pilosa, media ad 10 cm lg., 3 cm It., superiora valde diminuta, ovata, lanceolata vel oblongo- lanceolata, omnia ad medium inciso-pinnatilobata, lobis 3—5-partitis, basi auriculato-dilatata, semiamplexicaulia, margine inaequaliter crenato-dentata, aculeis numerosis luteolis inaequilongis; lobi foliorum in aculeum longum 12-15 mm lg. acutati; folia suprema sub calathidiis parva, lanceolata, pinnatidentata, aculeis luteolis firmis longis 8—10 mm lg., involucro aequilonga vel id vix superantia. Calathidia ad api- ces caulis et ramorum 2—5 approximata, subsessilia involucrum glabrum, rotundato ovatum, 2—3.5 cm in diam., demum late apertum, phyllis subhorizontaliter reflexis; phylla-externa mediaque ovato-lanceolata, vel lanceolata, internis subbreviora, a medio in acumen membranecum attenuata, in aculeum longum tenuem acutata, margine aculeata, aculeis divaricatis luteolis, interna lineari-lanceolata, apicem versus in acu- men subdilatatum margine undulatum acuminatum subreflexum attenuata; flores rosei, 17 mm lg., tubo angusto 7.5 mm lg., parte dilatata fere ad medium dissecta; pappus luteolo-albidus, ad 13 mm lg., pilis tenuissimis, internis apice scabris elongatis. Habitat in regione montana superiore. Typus. Tadzhikhstania, Pamiro-Alaj, Schugnan, in valle Gunt, Ryvak, 29 VII 1901, O. et B. Fedtschenko. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur. Affinitas. Ab omnibus speciebus sectionis Echenaidis (Cass.) Petrak ex Asia Media involucri phyllis externis margine longe aculeatis, involucri phyllis internis apicem versus in acumen membranaceum angustum acuminatum attenuatis, foliis margine aculeis longioribus tenuioribus armatis differt. 14. Subsect. Montana Charadze subsect. nova (sect. Cirsium). Calathidia haud numerosa, ad apices caulis et ramorum lateralium congesta. Folia magna, lata, integra vel late pinnatilobata, glabra vel subtus arachnoideo-vel fere tomentoso-pilosa. Typus subsectionis: C. pseudopersonata Boiss. et Bal. Area geographica subsectionis: Europa austro-orientalis, Caucasus occidentalis, Asia Minor (Anatolia borealis). 15. Subsect. Aciculata Charadze subsect. nova (sect. Cirsium). Calathidia mediocria, foliis supremis margine longe aciculatis eis 605 brevioribus vel longioribus obvallata. Involucri phylla ab ovatis ad lineari-lanceolata in aculeum brevem vel mucronem acutata. Typus subsectionis: C. obvallatum (MB.) MB. Area geographica subsectionis: Caucasus, Asia occidentalis. ONOPORDUM L. 16. O. prjachinii Tamamsch. sp. nova (sect. Onopordum). Planta 60—100 cm alt., lucide virenti-grisea; caulis superne ramosus, usque ad apicem alatus; alae sursum gradatim angustatae, margine sinuato-spinulesae; floia caulina media 13—25 cm lg., 2.5—4 cm It., decurrentia, apice acuminata, spinula terminata, margine inaequaliter pinnati-sinuata; lamina foliorum utrinque griseo-arachnoidea. Calathidia complanate globosa, 5 cm in diam., ad apices ramorum divaricatorum solitaria; involucrum subglobosum; involucri phylla basi albido- tomentosa, superne glabra, virescentia, fructificatione reflexa, spinulosa, omnia aequilonga; corolla pallide rosea vel albida; pappus pallide aureus, setis inaequilongis basi scabrido-setulosis apice glabris. Habitat in fruticetis montium Asiae Mediae. Typus. Asia Media, in valle fl. Vachsch, prope Cziljak in fruticetis, 26 V 1939, M. Prjachin. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur. Affinitas. Habitu O. acanthio L. similis est, sed involucri phyllis, calathidiis solitariis et eorum magnitudine differt. Ut videtur species hybrida. 17. O. frickii Tamamsch. sp. nova (sect. Erecta Rouy). Planta non vel vix armata; caulis 90-100 cm alt., costatus, in parte superiore vel ad medium longe multiramosus, ramis vix divaricatis, ut caulis spinosoalatis; ala griseo-arachnoidea, 0.5—0.8(10) mm It., spinis brevibus luteis praedita; folia caulina sessilia, late lanceolata, 12 cm Ig., 2 cm It., decurrentia. Calathidia solitaria, plus minusve parva, 3 cm lg., 1 cm It., oblongo-cylindracea; involucri phylla omnia imbircata, adpressa, inaequilonga, exteriora 0.6—-1.2 cm lg., ovato-lanceolata et lanceolata, media lineari-lanceolata, intima e basi oblongata in aculeum fusco-violaceum transeuntia; achenia 5-6 mm lg., 2-3 mm Ilt., complanate ovata; pappi setae scabrido-ciliatae, achenio duplo breviores. Habitat in Armenia. Typus. Armenia, Laar prope Erivan, in agris, n° 693, Frick. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur. 60 oO 606 Affinitas. Nostra species nova nulla specie caucasica generis Onopordi similis est; calathidiis parvifloris elongatis (non globosis), habitu et caule gracili parce vel non aculeato valde differt. 18. O. seravschanicum Tamamsch. sp. nova (sect. Erecta Rouy). Planta robusta, sed non elata, albido-cinerea; caulis 1 cm in diam., sulcato-striatus, albo-tomentosus, divaricato-ramosus; folia basalia breviter petiolata, petiolis alatis crenulato-dentatis, caulina media superioraque sessilia, 12—12.5 cm lg., 2—2.5 cm It., pinnatilobata, lobis triangulatis, spina tenui flava terminata; foliorum omnium lamina su- pra valde tomentosa, subtus tomentosissima et rugulosa vel crispata; folia ramulorum gradatim diminuta, sub calathidiis 2 cm lg., 0.5 cm It., lanceolata, integerrima vel denticulata, spinulosa. Calathidia oblongo- globosa, in inflorescentiam racemosam aggregata; involucrum subglobosum; involucri phylla extrema parva, 0.5—0.6 cm lg., ovato- lanceolata, media lineari-lanceolata, vix reflexa, 2—2.5 cm lg., in spinam transeuntia, basi glabriuscula, superne vix araneoso-pubescentia, gladulosa, margine ciliato-glandulosa, intima basi anguste linearia, apice subulata, violacea, margine ciliata; corolla albida, minute glandulosa; pappi setae barbellatae. Habitat in montibus altis Asiae Mediae. Typus. Seravschan, Rars (Maida-Kischtage pass.), alt. 9-10 000 pd., 16 VII 1893, V. Komarov. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur. Affinitas. Habitu O. polycephalo Boiss., quod in Tauro Cilicico habitata, simile, O. /eptolepidi appropinquat, sed involucri phyllorum et calathidiorum forma valde differt. SERRATULA L. 19. Sect. Iljinia Boriss. sect. nova.—Plantae humiles, acaules vel caulibus 2—20 cm alt., sed tum foliis integris; folia rosulata, radicalia et caulina lyrata, petiolis plus minusve longis, utrinque 2—5-lobata. Calathidia 1.5—3.5 cm in diam., sessilia vel pedunculata, pedunculis plus minusve elongatis; involucri phylla 7—9-seriata. Typus sectionis: S. lyratifolia Schrenk. 20. S. modesti Boriss. sp. nova (sect. /Jjinia Boriss.). Planta perennis; radices funaliformes, fuscae; rhizoma robustum, longum; caules 10-20 cm alt., simplices, basi reliquiis foliorum vetustorum obtecti; folia radicalia glabra, 8-15 cm lg. et 2.5—4 cm It., petiolis (1)5—10 cm lg., margine acute serrata, interdum basi utrinque 607 607 uni-bilobata, solida et coriacea, apice acutiuscula, acumine brevi molli tenuique. Calathidia 2.5—-3.5 cm in diam., pedunculis foliis fere aequilongis; involucrum 2-3 cm in diam., basi planiusculum, non angustatum, parti superiori aequilatum; involucri phylla 7—9-seriata, exteriora et media ovata, gradatim prolongata, anguste nigro-marginata, viridia, apice acumine caduco, interna lanceolata, appendice membranacea; receptaculum dense setosum, setis levibus 5 mm lg.; corolla purpurea, 25 mm lg., parte inferiore filiformi-tubulosa ca. 13 mm lg., parte superiore anguste campanulata 13 mm lg., lobis linearibus 7 mm lg.; tubus antherarum corolla subbrevior, superne lobatus, lobis liberis obtusis, basi appendicibus linearibus brevissimii praeditus; filamenta libera, parce papillari-pubescentia; stylus corolla paulo longior, breviter bilobatus; achenia ovata, 5 mm lg. et 3 mm It., sublevia, tenuis sime sulcata; pappus ca. 16 mm lg., densus, flavidus; pappi setae appress brevissime plumosae. Habitat in declivibus lapidosis siccis et collibus jugi alatau Dshungarici. Typus. Kasachstania, distr. Dzharkent (nunc Panfilov), flum, Kapkak, in declivibus lapidosis et siccis, 13 VII 1912, V. Saposhnikow et B. Schischkin. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur. Affinitas. A S. lyratifolia Schrenk caule evoluto, foliis integris, calathidiis et floribus majoribus differt. 21. Sect. Leuzeopsis Boriss. sect. nova.—Plantae monoeciae; fo- lia coriacea, magna, integra vel praecipue laminarum parte inferiore pinnatilobata, nervis prominentibus, margine obtuse crenato-dentata. Calathidia magna, 3—4 cm It., subglobosa, floribus hermaphroditis; involucri phylla coriacea, glabra, appendice reflexa aculeata S—6 mm Ig.; achenia ovata, ad basin subito angustata; pappus longus, 11-15 mm lg.; achenia ovata, ad basin subito angustata; pappus longus, 11— 15 mm lg., achenio duplo longior. Typus sectionis: S. serratuloides (Fisch. et Mey.) Takht. 22. Sect. Demetria Boriss. sect. nova.—Plantae monoeciae; caules late alati; folia magna, decurrentia, margine non profunde sinuato- dentata et aculeata. Calathidia magna, 2.5—3.5 cm in diam., globosa vel ovata; involucri phylla coriacea, subglabra, spinis longis 7-10 mm lg. erectis; corollae pars dilatata parte inferiore angusta 1.5-plo longior. Typus sectionis: S. caucasica Boiss. 608 608 RHAPONTICUM Adans. 23. Sect. Eleutherochaetum Soskov sect. nova.—Pappus flavescenti-rufescens vel flavescenti-cremeus, setis breviter plumosis rigidis basi in anulum non connatis, setae pilis latitudine eius axis duplo-triplo longioribus. Achenia 8-12 mm lg., 3-4.5 mm It., flavescenti-helvola vel flavescenti-brunnea. Corolla flava. Typus sectionis: R. integrifolium Wink. CENTAUREA L. 24. C. razdorskyi Karjag. sp. nova in litt. (subgen. Centaurea). Perennis, pallide virens, saepe glaucescens; caulis basi reliquiis squamatis vel plus minusve fibrosis brunnescentibus vaginarum floiorum emarcidorum vestitus, glaber, erectus, ramis paucis elongatis, 40—70 cm alt. et altior; folia radicalia caulinaque inferiora petiolata, in axillis et rachidum parte inferiore arachnoideo-lanata, caulina ambitu oblonga vel elongato-oblonga, 8—30 cm lg., 3-13 cm It., glabra, inferiora secus rachidem laxe et longe albo-lanata, segmentis numerosis lanceolato- linearibus vel lanceolatis 1-7 cm lg. acutis vel acuminatis secus rachidem decurrentibus margine crebre serrulatis, serrulis cartilagineo- mucronatis, vel plus minusve pinnatifida, laciniis triangularibus triangulari-lanceolatis vel lanceolatis; folia superiora sessilia, sensim diminuta, pinnatisecta, laciniis angustis integerrimis, suprema linearia, integerrima, acuminata. Calathidia ovata, 1.5—2.5 cm It.; involucri glabri phylla adpressa, subcoriacea, lutescentia, lineis S—9 longitudinalibus subparallelis fuscis ornata, extrema late ovata vel subrotundata, obtusissima, margine tenuiter membranacea, interiora oblonga, appendice ovata tenuiter membranacea lacera non emarcida; flores laete citrici, marginales aucti; achenia 7-8 mm lg., oblonga, subtetragona, levia, ad apicem vix transverse rugulosa, areola obliqua lutea marinata; pappus scabridus, cinnamomeus, achenio aequilongus. Fl. VI-VII, fr. VII-VIII. Habitat in declivibus lapidosis siccis Caucasi orientalis (Kobystan). Typus. Azerbaidzhania, Kobystan, mons Agh-burun, 12 VI (fl.) et 14 VII (fr.) 1952, I. Karjagin. In Herb. Inst. Bot. Acad. Sc. RSS Azerbaidzhaniae (Baku) conservatur. Species in memoriam cathedrae botanicae Universitatis primae in Azerbaidzhania a. 1923 fundatoris professoris V. Th. Razdorsky nominatur. Affinitas. Valde affinis C. kasakorum Ijin et C. talievii Kleop. e cyclo C. ruthenicae Lam., sed a prima forma segmentorum foliorum et 609 609 eorum rachide laxe albo-lanato, a secunda caule glabro et foliorum pubescentia differt. 25. Subsect. Fischerianae Czer. subsect. nova (subgen. Cyanus (Juss.) Hayek sect. Protocyanus Dobrocz.).—Appendices involucri phyllorum ea omnino vel partim obtegentes, margine fimbriatae, fimbrillis longis argentatis vel albis; corolla florum radii lilacino- rosea, purpurea, pallide flava, raro albida vel alba. Radix obliqua vel subverticalis, comparative longa, superne in rhizoma vulgo ramosum stolonibus et radicibus adventitiis funaliformibus destititum abiens; rami rhizomatis oblique ascendentes, breves vel plus minusve elongati, sat tenues, folia rosulata et caules floriferos in numero 1—3(5) emittentes; caules humiles vel mediocres, simplices vel in parte superiore parce compresse ramosi, vulgo sat robusti, ascendentes, arcuatim curvati, interdum in parte inferiore fere humifusi, raro erecti, basi vaginis nigro- vel purpureo-brunnies valde dilatatis petiolorum foliorum radicalium normaliter evolutorum ad tempus florendi haud marcescentium omnino tecti; folia oblique sursum directa, rigidiuscula, a lyrato-pinnatisectis ad integerrima; pubescentiae areneosae pili basi valde dilatati (sub lente!). Plantae in regionibus subalpina, alpina, rarius montana media montium Caucasi, Asiae Minoris, Iraniae Borealis (jugum Elburs) et partis meridionalis peninsulae Balcanicae Distributae sunt. Typus subsectionis: C. fischeri Willd. 26. C. willdenowii Czer. sp. nova (subgen. Cyanus (Juss.) Hayek sect. Protocyanus Dobrocz. subsect. Fischerianae Czer.). Planta perennis, tota viridulo-grisea, pubescentia araneosa vel araneosolanata tecta, ob tomentum tenue araneosum rarius grisea vel griseola, pilis basi valde dilatatis (sub lente!); caules non numerosi, (8)15—35 cm alt., plus minusve robusti, interdum comparative graciles, ascendentes, arcuatim curvati, raro suberecti, vulgo simplices; folia lanceolata, oblongo-lanceolata, rarius lineari-lanceolata vel oblongo- oblanceolata, apice a breviter acuminatis ad obtusiuscula, ab integerrimis ad profunde sinuato-pinnatipartita, rosulata, radicalia et caulina inferiora petiolata, cetera sessilia, plus minusve breviter deccurentia; folia caulina media 4—10(15) cm lg. et 0.4—2.5(3.5) cm It. Calathidia vulgo solitaria; involucrum ovatum vel globoso-ovatum, 18— 22(25) mm lg. et 12—15(18) mm It., praecipue glabrum; appendices involucri phyllorum exteriorum mediorumque semiorbiculares vel tri- angulares, phylla partim obtegentes, margine longe fimbriatae, fimbrillis argentatis 1.5-2.5 mm lg.; appendices phyllorum interiorum superne pallide fuscae vel albido-fuscatae, minores, fimbriatae; corolla florum disci violaceo-lilacina, lilacino-rosea, raro alba vel albida, 15-19 mm 610 610 lg., radii purpurea, roseolo-purpurea, raro alba, vel albida, 6—8-partita, 28-37(40)mm lg.; tubus corollae florum radii 13-20 mm lg., limbo 14-17(20) mm lg., lobis a lanceolato-linearibus ad lineares vulgo acuminatis; achenia ellipsoidalia, 4.5-5 mm lg. et 2.3-2.5 mm It.; pappus 2-3 mm lg., internus externo quadruplo brevior. Fl. VI—-VIII. Habitat in pratis subalpinis alpinisque (ad alt. 2700 m s. m.) Ciscaucasiae, Daghestaniae, Transcaucasiae orientalis australisque. Typus. Caucasus septentrionalis, vallis fluminis Chyzny prope ostium fluminis Tujala, 2000 ms. m., 7 VIII 1931, E. et N. Busch. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur. Affinitas. A proxima C. fischeri Willd. s. str. involucris minoribus ovatis vel globoso-ovatis, floribus minoribus et foliis breviter decurrentibus bene differt. 27. Subsect. Stricticaules Czer. subsect. nova (subgen. Cyanus (Juss.) Hayek sect. Protocyanus Dobrocz.).—Appendices involucri phyllorum ea partim obtegentes, margine fimbriatae, fimbrillis ab albidis ad nigras; corolla florum radii cyanea vel coerulea. Rhizoma verticale, brevissimum, stolonibus destitutum, radicibus adventitiis funaliformibus non numerosis tectum, rosulam foliorum haud emittens; caules vulgo solitarii, mediocres vel alti (ad 80 cm alt.), superne parce compresse ramose, rarius simplices, subgraciles, erecti, interdum basi vix ascendentes; folia oblique sursum directa, sat rigida, subangusta, integrrima, non raro radicalia et caulina inferiora utrinque uni-tridentata vel lobata; pubescentiae araneosae pili basi valde dilatati, rarius non dilatati (sub lente!). Plantae in planitiebus silvo-stepposis, stepposis et in regionibus praemontana et montana inferiore Europae mediae australisque nec non in regionibus australibus partis europaeae URSS distributae sunt. Typus subsectionis: C. stricta Waldst. et Kit. 28. Subgen. Czerniakovskya Czer. subgen. nov.—Calathidia mediocria, multiflora, ad apices caulis et ramorum lateralium solitaria, in inflorescentiam laxam corymbiformem vel subcorymbiformem agegregata; involucra ovata, ovalia vel oblongato-ovalia, 8-15 mm It., leviter araneosa vel subglabra; phylla_ coriacea, viridula (in sicco flavescentia), superne plus minusve striata, externa orbiculari-ovata et ovata, interna oblongato-lanceolata et oblongato-linearia; appendices phyllorum externorum mediorumque membranaceae, haud decurrentes vel decurrentes, triangulares vel lanceolato-triangulares, apice seta brevi elastica vel rigidiuscula instructae, margine pectinato-ciliatae; appen- dices phyllorum internorum membranaceae, fimbriato-ciliatae vel inaequaliter dentate; flores omnes tubulosi, flavi vel pallide flavo-rosei, 611 heterogeni, disci hermaphroditi, radii steriles, staminodiis albis in numero 5 in tubo corollae occultatis, non aucti, corolla 5—6(7)-partita; filamenta complanata, papillis tecta; stylus breviter bilobatus; achenia oblongato-ellipsoidalia, 7-8 mm lg., parce pilosa; pappus achenio aequilongus vel duplo longior, rufidulus, non deciduus, duplex, externus e setis sat multiseriatis planis serratis inaequilongis constans, internus externo 1.5—2-plo brevior, uniseriatus, paleaceus, paleis (10)12—15 plus minusve latis oblongatis vel oblongato-lanceolatis serrato-scabridis rigids. Herbae perennes, caulibus parce ramosis vel simplicibus ad apicem foliosis et foliis vulgo integerrimis tomentosis vel araneosis subsessilibus. Typus subgeneris: C. kopetdaghensis Ijin. 29. C. holophylla Socz. et Lipat. sp. nova (subgen. Psephellus (Cass.) Schmalh.). Planta perennis, caulis erectus, simplex, 30-60 cm alt., foltis radicalibus 1.5-plo longior, subglaber, sub calathidio parce albo- araneosus; folia supra viridia, subtus tenuiter albo-tomentosa, radicalia et caulina integra, margine denticulata, denticulis raris acuminatis, oblongato-ovata, basi cordata, apice acuta; folia radicalia magna, 10— 15(20) cm lg. et 4.5 cm It., longe petiolata, petiolis 10—20(30) cm lg., caulina minora, brevius petiolata; folium apicale basi cuneatum, breviter petiolatum. Calathidia solitaria; involucrum ovatum, 2.5 sm It.; appen- dices involucri phyllorum magnae, triangulari-lanceolatae, brunneae, nervo uno pullato, basi subnigrae, margine hyalinae, fimbrillis longis, lucide fuscis; flores violacei, marginales non aucti. Fl. VII-IX. Habitat in pinetis partis occidentalis Caucasi Majoris. Typus. Caucasus septentrionalis, pars superior systematis fluminis Kjafara affluentiae fluminis Zelenczuk, in pineto Kyzylezuk, ca. 1800 ms. m., 2 IX 1945, V. Soczava et V. Lipatova. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur. Affinitas. A speciebus omnibus sectionis Hypoleucarum (sosn.) Sosn. caule subglabro, foliis integris margine remote denticulatis differt. 30. C. ciscaucasica Sosn. sp. nova (subgen. Psephellus (Cass.) Schmalh.). Planta perennis; caules 20-30 cm alt., dense araneoso-tomentosi, vulgo simplices, rarius leviter ramosi; folia supra viridia, sparse araneoso-pubescentia, subtus plus minusve dense albo-tomentosa; fo- lia radicalia breviter petiolata, lyrato-pinnatipartita, lobo apicali magno ovato basi plus minusve cuneato vel rotundato apice acuminato margine remote et obscure denticulato, lobis lateralibus utrinque 2—3 minoribus ovatis brevissime petiolulatis; folia caulina non numerosa, vulgo inte- 61 = 612 gra, ovata, inferiora petiolata, superiora sessilia. Calathidia magna; involucrum globosum, 2 cm It.; involucri phylla plus minusve dense lanato-pubescentia; appendices involucri phyllorum parvae, lanceolatae, nigrescenti-fuscae, fimbrillis flaveolis vel albidis, appendicis latitudine longioribus; flores rosei, marginales aucti. Fl. V—VI. | Habitat in declivibus herbaceis montium Ciscaucasiae. Typus. Ciscaucasia, in vicinitate opp. Kislovodsk, declives ad vallem fluminis podkumok, mons Koltzo, 2 VII 1911, A. Gordjagin. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur. Affinitas. A C. abchasica (Alb.) Sosn. foliorum segmentis lateralibus bi-trijugis ad basin valde angustatis, involucri phyllis plus minusve dense lanato-pubescentibus bene differt. 31. C. kolakovskyi Sosn. sp. nova (subgen. Psephellus (Cass.) Schmalh.). Planta perennis; caules 15-35 cm alt., simplices; folia ob pilos appressos araneosos, supra griseola, subtus caerulescenti-griseo- tomentosa; folia radicalia cum petiolo ad 25 cm lg., ambitu oblongato- ovata, pinnatipartita, lobo apicali fere non aucto oblongato-ovato 6—14 cm lg. integro vel basi uni-bifido margine remote denticulato apice acutiusculo basi plus minusve rotundato petiolulato, lobis lateralibus utrinque 3—8 ad basin laminae gradatim decrescentibus ambitu ovatis vel plus minusve ellipticis obtusiusculis basi rotundatis superioribus confluentibus inferioribus petiolulatis; folia caulina diminuta, non numerosa, superiora integra, oblongato-lanceolata. Calathidia mediocria; involucrum 1.5 cm It.; appendices involucri phyllorum oblongato- lanceolatae, fuscatae, breviter pubescentes, fimbrillis albidis; flores rosei, marginales leviter aucti. Fl. VIJ—VIII. Habitat in declivibus schistosis calcareis siccis in regione alpine Abchasiae. Typus. Abchasia, jugum Gagrinsky, mons Chyrka, 27 VIII 1947. A. Kolakovsky et V. Jabrova. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur. Affinitas. A C. leucophylla MB. foliis subtus caesio-tomentosis (non albo-tomentosis), involucri phyllis appendicibus oblongo- lanceolatis fuscatis praeditis differt. 32. C. schistosa Sosn. sp. nova (subgen. Psephellus (Cass.) Schmalh.). Planta perennis; caules ascendentes, (7)10—15(30) cm alt., simplices vel parce ramosi, araneoso-lanati, fere ad calathidia foliosi; folia supra leviter araneoso-lanata, subtus albo-tomentosa; folia radicalia ‘pinnatisecta vel lyrato-pinnatisecta, segmento apicali magno elliptico 612 613 vel ovato integro vel lobato, segmentis lateralibus minoribus ovatis vel oblongato-ellipticis decurrentibus, interdum partim integra, elliptica; folia caulina paulo diminuta, pinnatisecta, segmentis brevibus oblongatis vel ellipticis integris vel leviter partitis obtusis breviter acuminatis. Calathidia mediocria; involucrum 15-20 mm It.; appendices involucri phyllorum oblongato-ovatae, albo-scariosae, hyalinae, dorso stria plus minusve lata brunnea ornatae, fimbrillis albis, appendicis latitudini aequalibus vel ea vix brevioribus, interdum ex toto albae; appendices phyllorum intimourum triangulares, fimbriatulae, macula brunnea notatae, rarius albidae; flores purpurei, marginales aucti. Fl. V. Habitat in declivibus schistosis montium Transcaucasiae orientalis. Typus. Caucasus orientalis, Azerbaidzhania, distr. Divitschi, in faucibus Isti-su inter pag. Arabdagnja (Chaltan-dagnja) et thermas Isti- su, in schistosis, 31 V 1937, I. Karjagin et M. Schevljakov. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur, isotypus in Baju. Affinitas. A C. hymenolepide Trautv. foliis supra griseolis (non viridibus), calathidiis majoribus, involucro 1.5—2 cm It. (non 1 cm It.) differt. 33. C. meskhetica Sosn. sp. nova (subgen. Psephellus (Cass.) Schmalh.). Planta perennis; caules numerosi, humiles, 5—15(25) cm alt., ascendentes vel decumbentes, ramosi, basi villosi, dense tomentosi, ad apicem foliosi; folia supra griseolo-viridia, scabra et araneosa, subtus albo-tomentosa; folia radicalia breviter petiolata, ambitu oblongato- ovata, bipinnatipartita, caulibus fere aequilonga, interdum eis longiora, segmentis ovatis vel lanceolatis sessilibus pinnatifidis; folia caulina pinnatipartita, segmentis anguste lanceolatis, suprema pinnata, segmentis . linearibus, interdum integra, lanceolata. Calathidia mediocria; involu- crum vulgo ovatum, 1-1.5 cm lt.; appendices involucri phyllorum pallidae, lucide stramineae, fimbrillis albidis, appendicis latitudini vulgo aequalibus; flores rosei, marginales vix aucti. Fl. VI—VIII. Habitat in declivibus siccis et collibus argillosis in regione montana media Transcaucasiae orientalis. Typus. Meskhetia, colles argillosi gypsacei contra opp. Achaltziche, ad ripam sinistram fluminis Poskhov-czai, 4 VII 1947, D. Sosnovsky, Kutateladze et I. Mandenova. In Herb. Inst. Bot. Acad. Sc. RSS Georgiae (Tbilissi) conservatur, isotypus in Leningrad. Affinitas. A C. carthalinica (Sosn.) Sosn. foliis radicalibus cauli subaequilongis, involucri phyllorum appendicibus pallidis lucide stramineis fimbriatis fimbrillis albidis appendicis latitudini aequalibus vel ea longioribus differt. 613 614 34. C. substituta Czer. sp. nova (subgen. Jacea (Juss.) Hayek sect. Jacea). Planta perennis, tota vulgo griseolo-viridis vel griseola, pubescentia araneosa plus minusve densa tecta, rhizomate lignoso abbreviato, ra- dices funaliformes emittente; caules solitarii vel non numerosi, ad 80 cm alt., erecti, costatosulcati, sub calathidiis leviter incrassati, a medio vel supra medium sat valde ramosi, ramis elongatis, lilacino-purpurei, praecipue superne ob aculeolos acutos vel pilos rigidos vreves scabri, folia anguste lanceolata, oblongato-oblanceolata, anguste oblongato- lanceolata, interdum caulina superiora sublineari-lanceolata, apice ab acuminatis ad acutiuscula, integerrima, rarius radicalia et caulina me- dia remote denticulata vel haud profunde sinuato-lobata, utrinque pilis densiusculis vel sparsis rigidis curvatis multicellularibus tecta; folia radicalia et caulina inferiora petiolata, sat cito marxescentia, cetera sessilia, ad basin subangustata. Calathidia ad apices caulis et ramorum lateralium solitaria, in inflorescentiam corymbiformem vel racemoso- corymbiformen aggregata; involucrum globoso-ovatum, rarius subglobosum, 16-18 mm lg. et 12—16(18) mm It., glabrum, basi griseolo-araneosum; appendices involucri phyllorum membranaceae, magnae, phylla omnino obtegentes, albidae, in parte media pallide fuscatae vel fuscatae, raro ex toto pallide fuscatae; appendices phyllorum exteriorum mediorumque orbiculares, convexae, laceratae, margine irregulariter dentatae vel fimbriato-dentatae; appendices phyllorum interiorum ab orbicularibus ad ovatas, leviter laceratae, integrae vel apice irregulariter denticulatae; flores lilacino-purpurei; achenia 3 mm lg. et 1-1.3 mm It.; pappus nullus. Fl. VII-VIII. Habitat in pratis et fruticetis, ad margines silvarum in regionibus australibus partis europaeae URSS. Typus. Tauria, Hortus Nikitensis, prope semitam, 28 VII 1922, S. Stankov. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur. Affinitas. Nostra species nova C. pannonicae (Heuff.) Hayek affinis est, sed ab ea differt involucris globoso-ovatis vel subglobosis 12— 16(18) mm It. (non ovatis (8)10—12 mm It.), caulibus foliisque densius pubescentibus et caulibus magis ramosis. 35. C. turgaica Klok. sp. nova (subgen. Heterolophus (Cass.) Dobrocz.). Caules floriferi 1-10 ex axillis foliorum radicalium oriundi, procumbentes vel ascendentes, 5—25 cm lg. vel alti, simplices vel vix ramosi, ramulis 1—2, albido-tomentosi; folia radicalia rosulantia, 3.5-12.5 cm lg., pinnatisecta, segmentis lateralibus 2—7-paribus oblanceolatis vel ellipticis 1.5—20 mm lg. et 1.5—7(11) mm It. acutatis vel obtusis margine sicut apicale remote denticulatis; segmentum, 614 615 apicale usque ad 16 mm It.; folia caulina apicala integra; folia omnia subtus albo-tomentosa, supra griseola. Calathidia in caule 1—3; involu- crum subglobusum 15—22 mm lg., 14-22 mm It.; appendices phyllorum externorum lanceolatae, fimbrillis sub apiculo utrinque 1—2, extremorum subdenticulatae, reflexae, ca. 2 mm lg.; appendices phyllorum mediorum phylla omnino obtegentes, atriusculo-fulvae necnon brunneo-aurantiacae vel cinnamoneae, suborbiculares vel late ovato-triangulares, 5-§ mm lg. et It., margine regulariter fimbriatae, fimbrillis 1.5—2(3) mm lg., apiculum superantibus; phyllorum internorum appendix oblongo-ovata vel subovalis, 1.5—-3.5 mm lg., margine irregulariter denticulata, infra medium lilacino-cinnamomea; flores purpurei; achenia 5—6 mm lg., 1.7—3 mm It., pappo ca. 1.5 mm lg. Fl. V—VII. Habitat in steppis et declivibus montanis. Typus. Ditio Turgaica, in viciniis opp. Kustanaj, steppa stiposa, 17 V 1904, S. Fedossejev. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur. Affinitas. A proxima C. sibirica L. species haec magis xerophila caulibus subduplo brevioribus debilioribus, foliis duplo minoribus, segmentis foliorum radicalium oblongioribus, involucro calathidiorum subgloboso apice haud complanato, appendicibus phyllorum obscurioribus et brevius fimbriatis necnon habitatione australiore recedit. 36. C. sergii Klok. sp. nova (subgen. Heterolophus (Cass.) Dobrocz.). Caules floriferi axillares 2-3 (raro plures), procumbentes vel ob- lique ascendentes, (8)10—20 cm lg. vel lati, simplices vel ramificati, ut folia arachnoideo-tomentosi; folia radicalia brevipetiolata, 6-15 cm lg., pinnatisecta, segmentis 7-13; segmenta lateralia oblongo-lanceolata vel elliptica, 5-32 mm lg., 5-11 mm It., acutata, apice attenuata cal- losa glabra; segmentum apicale usque ad 50 mm lg. et 15 mm It.; folia caulina inferiora pinnatipartita, segmentis saepissime 5, superiora in- tegra; folia omnia ab utroque latere ob pubescentiam arachnoideo- lanulosam albida, margine remote crenulato-denticulata. Calathidia in caule 1—4; involucrum oblongo-ovoideum, 15-20 mm lg., 7-10 mm It., basi arachnoideo-lanulosum; phylla ovata vel oblonga, appendicibus vix tecta, eis longiora, sub appendicibus pubescentia; appendices angustae, fulvae, haud attingentes, in phyllis externis brunneolae, anguste lanceolatae vel lineari-lanceolatae, ca. 3 mm lg. et 1 mm It., integerrimae, plus minusve reflexae, in mediis lanceolatae necnon oblongo-ovato-lanceolatae, 2.5-5 mm lg., 1-2 mm Ilt., margine irregulariter fimbriatulae, fimbrillis utrinque 2-7 usque ad 1 mm lg., acumine fimbrillis longiore, rarius subdenticulatae vel integerrimae, in internis ovales vel oblongo-obovatae, minutissime eroso-denticulatae; 615 616 flores roseo-purpurei; achenia 6.5—7 mm lg., ca. 2 mm It., pappo 2-3 mm lg. fumagineo. Fl. VI—VII. Habitat in declivibus lapidosis vel arenosis austro-sibiricis. Typus. Sibiria, in collibus arenosis ad fl. Irtysch, Altai, C.A. Meyer. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur. Affinitas. A C. sumensi Kalen. europaea species haec asiatica involucro magis oblongo, appendicibus phyllorum mediorum longioribus et latioribus fulvis (non atro-cinnamomeis), acheniis longioribus (non 44.5 mm lg.), pappo eis subduplo longiore differt. 37. C. ossethica Sosn. sp. nova (subgen. Lopholoma KCASs: ) Dobrocz. sect. Lopholoma). Planta ad 80-100 cm lat., pilis brevibus papillaribus et crispatis intermixtis tecta, tomentello araneoso tenui, leviter scabra, viridis; caules solitari1 vel non numerosi, erecti, simplices vel parce ramosi; folia pinnatipartita vel pinnatilobata, inferiora sat longe petiolata, superiora sessilia vel subsessilia, interdum integra; lobi laterales utrinque 2—5, oblongati, integerrimi vel plus minusve dentati; segmentum apicale lateralibus vulgo majus. Involucrum 19—23 mm It., 14-25 mm lg., glabrum vel subglabrum; appendices involucri phyllorum exteriorum mediorumque tenuiter coriaceae, obtuse triangulares, 2—3.5 mm lg. (sine margine decurrente), nigro-fuscae, margine ciliatae, ciliis albidis 2-3.5 mm lg., apice acumine brevissimo (ad 1—1.5 mm lg.); involucri phylla intima appendicibus suborbicularibus margine profunde fimbriatis; flores rosei, marginales paulum aucti; achenia S—6 mm lg., breviter sparseque pilosa; pappus 4.5-6 mm lg., pappi interioris setae paleiformes, exterioris setis longissimis 2.5—3-plo breviores. FI. VII- IX. Habitat in declivibus lapidosis in regionibus media et superiore montium Caucasi. Typus. Ossetia australis, in fauce Ediss, ca. 2050 m s. m., 6 IX 1933, E. et N. Busch. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur. Affinitas. A proxima C. scabiosa i acheniis et oe longioribus et area geographica differt. 38. C. kubanica Klok. sp. nova (subgen. Acrolophus (Cass.) Dobrocz.).—C. maculosa auct. fl. cauc. p. p. non Lam.—C. rhenana Dumb. in Dokl. Akad. Nauk Arm SSR V, 2 (1946) non Boreau. Caules solitarii, erecti, 20-110 cm alt., costulato-sulcati, inferne foliiscum subarachnoidei, superne scabridi, ramosi; folia radicalia petiolata, pinnatisecta, segmentis incisis, ‘marcéscentia; folia caulina brevipetiolata (infima) et sessilia, 1-8 cm lg., pinnatisecta, segmentis 617 lateralibus 1—11-paribus integris vel in foliis inferioribus incis, subtus et margine scabrido-setosiuscula; foliorum lobi terminales lineares vel oblongi, 2—27 mm lg., 0.3—5 mm Ilt., acuminati et subaculeati; folia sessilia, basi auriculata, auriculis integris vel dissectis; folia apicalia integra, lineari-oblonga, incurvata. Inflorencentia communis oblongo- paniculata, ramis 0.5—14 cm lg. parciuscule foliosis, calathidio unico terminatis; involucrum ovoideum vel late ovoideum, 12—15 mm lg., 7— 12 mm It., bicolor; phylla ovata usque ad linearia, 3-11 mm lg., 1.5— 3.2 mm It., 5—7-nervia, virescentia, glandulis sessilibus luteis minutis obsita; appendices late ovatae necnon (in phyllis internis) ellipticae, 1.5—2.5 mm lg., 1-4 mm It.; appendices phyllorum exteriorum mediorumque parte media integra triangulari vel ovato-triangulari ni- gra, acumine 0.5—1.5 mm lg. molli margine regulariter fimbriato, fimbrillis 5—10-paribus 0.3—2 mm lg. basi nigrescentibus superne rufescentibus apice capilliformibus tortuosis, interiorum ellipticae vel ovato-lanceolatae, denticulatae vel integerrimae; flores purpureo-rosei; corollae florum marginalium 20-21 mm lg., centralium 14-15 mm lg.; achenia oblonga, subobliqua, 2.7—3.2 mm lg., 1.2—1.5 mm It., pilosiuscula, matura nigrescentia, flavido-striata; pappus 1—2.3 mm lg. Fl. VII-VIII. Habitat in steppis praemontanis vel montanis caucasicis. Typus. Kubania, steppae montanae ad ostium fl. Dzemagat, 1907. N. Busch.—Regio Kubanica, steppa montana ad ripam dextram fl. Allu- Kama, in declivibus inferioribus inter ostia, 1907, E. et N. Busch. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur. Affinitas. Ab occidentali C. rhenana Boreau species haec caucasica inflorescentia communi paniculata (haud corymbosa), fimbrillis appendicum rufescentibus (non nigrescentibus) differenda est. 39. C. sophiae Klok. sp. nova (subgen. Acrolophus (Cass.) Dobrocz.). Caules ca. 50-60 cm alt., in costis obtusis leves, densiuscule arachnoidei, ramosissimi; folia caulina inferiora pinnatisecta, segmentis linearibus ca. 1-2 mm It., margine scabrida; folia apicalia linearia, 4— 15 mm lg., 0.5—1 mm It. Calathidia apice ramorum 2.5—9 mm longorum solitaria; involucrum oblongo-ovoideum, 9-12 mm lg., 5-6 mm It.; phylla ovata usque ad linearia, 2-8 mm lg., 1.2—2.5 mm It., 5-nervia, fulvida; appendices albido-fulvidae, in phyllis externis mediisque transversim ovales, 1.5—2 mm lg., 2.5—3.5 mm It., margine crenulatae et apice tenuiter laceratae; internorum appendices oblongo-ovato-lan- ceolate, ca. 1.5—2 mm lg., subintegerrimae; flores rosei; corollae florum marginalium ca. 15 mm lg., centralium ca. 13 mm lg.; achenia oblonga, 3.2—-3.6 mm lg., 1.2—1.6 mm It., matura atro-olivaceo-brunnea, pappo 2-3 mm lg. Fl. VII-VIII. 617 618 Habitat in arenosis tanaiticis. Typus. Ditio Septentrionali-Caucasica, regio Salskiensis, 3—4 km SW versus a st. Tzymljanskaja, terrasa (tanaitica) prima, silva inundata, in pulvino ad Tanaim, 10 VIII 1927, S. Gorschkova. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur. Affinitas. Species haec a C. majorovii Dumb. involucro oblongiore (non ovato-conoideo), caule levissimo (nunquam superne in costis scabro), a C. arenaria MB. (incl. C. wolgensis DC.) involucro latiore, caulibus crassioribus et densius arachnoideis, ramis floriferis valde brevioribus, foliis margine scabris, nunquam levibus recedit. 40. C. steveniana Klok. sp. nova.—(sect. Acrolophus (Cass.) Dobrocz.).—C. caprina auct. fl. taur. non Stev. Caules usque ad 80 cm alt., basi subteretiusculi levesque, superne in costis scabrido-setosiusculi, foliiscum arachnoidei, subdivaricato- ramosi; folia radicalia necnon caulina inferiora petiolata, bipinnatisecta, subtus setulis aculeoliformibus 0.2—0.3 mm lg. dense obsita, lobulis terminalibus lineari-oblohgis oblongo-lanceolatis vel obovatis usque ad 20 mm lg. et 3-4 mm It.; folia media sessilia, pinnatisecta, segmentis lateralibus 1—3-paribus integris vel incisis. Calathidia apice ramulorum 3-12 cm longorum solitaria; involucrum oblongo-ovoidem, (9)10—12 mm lg., 5—6 It.; phylla ovata usque ad linearia, appendicibuscum 3—9.5 mm lg. 1—-2.5 mm It., 3—5-nervia; appendices orbiculares neconon ellipticae, 1.2—2.7 mm It., acumine 0.2—1.2 mm lg. molliusculo; phyllorum externorum mediorumque appendices margine fimbrillis regularibus 2—5-paribus albi dis ca. 0.5—2 mm lg. ornatae, sub eis paleaceo-auriculatae, auriculis latis denticulatis vel laceris; appendicum pars media solida, ovato-lanceolata, cinnamomea vel sordide purpurea; flores roseo-purpurei; corollae florum marginalium ca. 14 mm, centralium 11—12 mm lg.; achenia 2.5—3 mm lg., 1—-1.4 mm It., albida, pappo nullo vel vix evoluto ca. 0.5 mm lg. Fl. VI-VIII. Habitat in sabulosis vel declivibus glareosis litoralibus ad Pontum Euxinum. Typus. Ditio Taurica, Eupatoria, in arenis litoralibus ad Pontum Euxinum, 6 VII 1956, M. Kotov. In Herb. Inst. Bot. Acad. Sc. RSS Ucrainiae (Kioviae) conservatur. Affinitas. C. caprina Stev. vera planta hybrida inter nostram et C. diffusam Lam. verosimiliter est. A C. ovina Pall. et aliis caucasicis proximis species nostra pappo achenii nullo vel subnullo (nunquam 1- 1.5 mm lg.) recedit. 41. C. koktebelica Klok. sp. nova (sect. Acrolophus (Cass.) Dobrocz.). 619 Caules nonnulli, erecti, 25-70 cm alt., basi subteretiusculi, supra costatosulcati, in costis scabri, flocculoso-arachnoidei, serius glabrati, a collo divaricato-ramosissimi; folia radicalia necnon caulina inferiora petiolata, pinnatisecta vel pinnatifida, arachnoidea vel arachnoideo- tomentella, subtus et margine setosiusculo-scabra; lobi terminales (segmenta) 2-15 mm lg., 1-4 mm It., acuminati; folia media pinnatifida, lobis linearibus acutatis vel acuminatis; folia apicalia linearia, acuminata. Calathidia partim sessilia vel subsessilia, congesta, partim apice ramulorum usque ad 4.5 cm longorum solitaria; involucrum oblongo-ovoideum, ca. 9 mm lg., 3.5—4 mm It.; phylla oblongo-ovata usque ad anguste linearia, 2.5-8.5 mm lg. (cum appendice), 0.8—1.7 mm It., 1—5-saepissime trinervia, virescentia, media internaque partim purpurea; appendices orbiculares, 1.5-2.5 mm lg., 1.5—-3 mm Ilt., acumine 0.5—1.3 mm lg., in phyllis mediis margine fimbrillis regularibus 4—6-paribus 1—-1.5 mm lg. ornatae, basi auriculis paleaceis denticulatis obsoletis auctae; appendicis triangularis pars media integra, flavida, basi saepe macula obscura ornata, cinnamomeopurpurea; flores purpurei; corollae florum marginalium 12-13 mm, centralium 9-10 mm lg.; achenia 3.1—3.2 mm lg., ca. 1.5 mm It., olivaceo-cinnamomea, pappo subnullo vix 0.1—0.2 mm lg. Fl. VIII—X. Habitat in steppis tauricis cimmericis. Typus. Tauria orientalis, regio Sudakiensis, prope pag. Planerskoje, in artemisieto, 6 X 1957, M. Klokov. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur, syntypi — Kioviae. Affinitas. A proxima C. steveniana Klok. calathidiis partim congestis et sessilibus (nunquam omnibus apice ramulorum solitariis), involucro breviore et angustiore abhorret; a C. ovina Pall. et aliis caucasicis proximis pappo achenii subnullo differt. 42. C. aemulans Klok. sp. nova (subgen. Acrolophus (Cass.) Dobrocz.). Radix palaris robusta; caules solitarii, erecti, 30-60 cm alt., sulcato- costati, in costis setosiusculi, foliiscum densiuscule arachnoidei, a collo divaricato-ramosissimi; folia radicalia necnon caulina infima petiolata, pinnatisecta, segmentis pinnatifidis vel partim integris; lobi terminales oblongi, oblongo-lanceolati vel obovati, 3-15 mm lg., 1-4 mm It., apiculati, subtus et margine setulis aculeoliformibus 0.2—0.3 mm lg. dense obsiti; folia media superioraque sessilia, pinnatifida, lobis oblongo-vel lineari-lanceolatis aculeato-acuminatis; folia apicalia in ramulis terminalibus oblongo-linearia vel linearia, 1.5—2 mm lg. et 0.3—0.4 mm It. Calathidia plus minusve in ramulis secundariis congesta, subsessilia, rarius apice ramulorum usque ad 2 cm longorum solitaria; involucrum subcylindricum, 10-12 mm lg., 3—3.5 mm It., stramineum, 61 oo 620 parum arachnoideum; phylla oblonga usque ad angusto-linearia, 2.5— 10 mm lg. (cum appendice), 0.5—1.5 mm It., nervis obsoletis 1-3; appendices 2-6 mm lg., acumine pungente 0.7—4 mm lg. et fimbrillis rigidis 0.3-1.7 mm lg. 3—7-paribus; acumina pungentia, divaricata et in phyllis exterioribus recurvata; flores purpurei, marcescentes pallidi, corollis 10-12 mm lg.; achenia 2.3-2.8 mm lg., 0.9-1.1 mm It. glabra, epapposa. Fl. VII-X. Habitat in steppis tauricis cimmericis. Typus. Tauria orientalis, regio Sudakiensis, prope pag. Planerskoje, in artemisieto, in declivibus, 10 X 1957, M. Klokov. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur, syntypi—Kioviae. Affinitas. A proxima C. diffusa Lam. calathidiis congestioribus saepius subsessilibus, involucro longiore (non solum 8—10 mm longo), acuminibus appendicum longioribus robustioribus et magis recurvatis, floribus purpureis (nunquam pallide roseis vel albidis) primo aspectu differt. 43. Sectio Pseudophalolepis Klok. sect. nova (subgen. Phalolepis (Cass.) Dobrocz.)—Herbae biennes. Appendices involucri phyllorum fere omnino membranaceae vel paleaceae, basi tantum triangulari vel fascia mediana angusta solidiusculae. Pappus achenio subaequilongus, interdum paulum longior vel manifeste brevior. Corollae roseae, pallide purpureae, cremaceae vel interdum flavae. Typus sectionis: C. gerberi Stev. Sectio paleopontica, relicta. 44. C. vankovii Klok. sp. nova (subgen. Phalolepis (Cass.) Dobrocz. sect. Phalolepis).—C. nikitensis Illar. in Bot. Mat. Gerb. Bot. Inst. Akad. Nauk SSSR, XVIII (1957) 305 p. p. quoad plantam jailicam. Caules 2-10, assurgentes, ca. 30-35 cm alt., ramosi, densiuscule arachnoidei, superne in costis scabri; folia radicalia necnon caulina inferiora 4-10 cm lg., petiolata, pinnatisecta, segmentis integris vel partim 2—3-fidis, arachnoidea, subtus setosiuscula, margine scabrido- serrulata; lobi folii laterales saepissime lineari-lanceolati, 9-25 mm lg., 1-2.5 mm It., acuminati; lobus (vel segmentum) apicalis eis submajor; folia superiora basi trifida, lobulis lateralibus abbreviatis, apicali magno usque ad 3 mm It.; folia apicalia sub calathidio disposita integra, linearia, 5-15 mm lg., ca. 1 mm It., supra sublevia, subtus scabra. Calathidia apice ramorum haud longorum solitaria; involucrum ovoideum, 14-16 mm lg., 10-14 mm It.; appendices orbiculares vel late ovatae, parte media solida ovato-triangulari atro-cinnamomea, marginali flavido-paleacea tenuiter denticulata vel subfimbrillata; phyllorum mediorum appendices ca. 3-4 mm lg., apice plane 619 621 emarginatae, acumine 0.5—0.7 mm lg. molli, externorum longius acuminatae, internorum acumine nullo; flores laete purpurei; achenia oblonga, ca. 4—4.2 mm lg., pilostuscula, matura nigrescentia, pappo 1.5—-1.8 mm lg. Fl. VI-VII. Habitat in declivibus montanis tauricis. Typus. Tauria, Jajla (ajpetrica), in locis lapidosis, 17 VI 1907, K. Levandovskij. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur. Affinitas. A proxima C. sterili Stev. (incl. C. nikitensi lar. p. max p.) praemontana species nostra jailicola caulibus humilioribus (nun- quam usque ad 100 cm alt.), involucre calathidii late oroideo (non oblongo-ovoideo) et majore (non 10-14 mm lg. et 6-10 mm It.), acheniis majoribus (non 2.5—3.5 mm lg.) et obscurioribus bene distincta. Praeterea C. sterilis Stev. saepissime biennis, nostra vero perennis est. 45. C. novorossica Klok. sp. nova (subgen. Phalolepis (Cass.) Dobrocz. sect. Phalolepis).—C. sterilis auct. fl. cauc. non Stev. Herba biennis; caules 1—3, erecti vel ascendentes, 20—70 (et ultra) cm alt., ramosi, costati, foliiscum densiuscule flocculoso-arachnoidei, suprene in costis scabri; folia radicalia necnon caulina inferiora petiolata, bipinnatisecta, lobis terminalibus angustis, sub anthesi saepissime marcescentia; folia media ca. 2—6 cm lg., sessilia, subtus et margine (sicut omnia) scabrido-setosiuscula, supra parum scabrida; lobi eorum laterales anguste lineares vel lineari-lanceolati, 5-25 mm lg., 0.5-1.5 mm It.; lobus apicalis usque ad 32 mm lg. et 2.5-3 mm It.; folia superiora pinnatifida, minora, apicalia sub calathidiis integra vel interdum basi subdentata, linearia, 7-12 mm lg., 1.2—-1.5 mm It., acuminata. Calathidia apice ramorum solitaria, in caule (1-3) 7-18, floribuscum 17—20 mm lg.; involucrum ovoideum, 12—15 mm lg., 8— 12 mm It., albidum, opacum, vix nitens; appendices in phyllis externis orbiculari-rhomboideae, acumine usque ad 1.5 mm lg. molli terminatae, margine sicut omnes tenuiter denticulatae, in phyllis mediis orbicu- lares, ca. 3.5-4 mm lg., ceteris paulum majores, margine lacerae, acumine nullo, basi in parte media macula cinnamomea triangulari notatae, in phyllis internis ovales; flores laete purpurei; achenia oblongo- cuneata, ca. 44.5 mm lg., pilosiuscula, matura nigricantia, pappo 1.5— 3.3 mm lg. Fl. VI-VI. Habitat in declivibus lapidosis praemontanis occidentali- transcaucasicis. Typus. Caucasus, prope opp. Novorossijsk, 6 VII 1892, V. Lipsky. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur. Affinitas. Species haec a C. sterili Stev. longiuscule distat, a magis affini C. sarandinakiae Ullar. Tauriae orientalis incolae lobis foliorum 620 622 terminalibus angustioribus, involucro angustiore (non 10—16 mm It.) albido (nunquam flavido), appendicibus basi maculatis (haud concoloribus) minoribus clare differt. 46. C. georgica Klok. sp. nova (subgen. Phalolepis (Cass.) Dobrocz. sect. Phalolepis).— C. pseudodeusta auct. fl. cauc. p. p. non Hayek. Herba biennis; caules solitarii vel pauci, erecti, 40-70 cm alt., a medio divaricato-ramosi, foliiscum haud dense arachnoidei, inferne leves, superne in costis scabri; folia radicalia necnon caulina inferiora petiolata, bipinnatisecta, lobis terminalibus linearibus vel lineari- lanceolatis angustis, sub anthesi saepissime marcescentia; folia media 1.5—4 cm lg., sessilia, pinnatisecta, segmentis pinnatifidis, subtus et margine aculeolato-scabra, supra scabrida; segmenta vel lobuli terminales lineares usque ad lineari-lanceolatos, ad + 10 mm lg., 0.5— 1.5 mm It.; lobus apicalis ad 2—3 mm It., folia apicalia integra, linearia, acuminata. Calathidia apice ramorum elongatorum primariorum vel breviusculorum secundariorum, in caule (3) 12-30, floribuscum 17-19 mm lg.; involucrum ovoideum, ca. 12—13 mm lg. et 8—9 mm It., flavido- cinnamomeum vel cinnamomescenti-flavum, nitens; appendices parte media solida, flavo-cinnamomea, marginali paleacea, flavida, tenuiter denticulata vel subfimbrillata, in phyllis externis ambitu late ovatae, apice in emarginatura acumine ca. 1 mm lg. terminatae, in mediis orbiculares, apice rotundatae, acumine nullo; flores laete purpurei; achenia ca. 3-7 mm lg., brunnescenti-olivacea, pappo ca. 1.5 mm lg. Fl. VI-VI. Habitat in declivibus lapidosis praemontanis Transcaucasiae occidentalis. Typus. Transcaucasia, Borshom, 19 VI 1865, G. Radde. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur. Affinitas. A C. novorossica Klok. involucro obscuriore differt. 47. C. latiloba Klok. sp. nova (subgen. Phalolepis (Cass.) Dobrocz. sect. Phalolepis). Caules solitarii vel pauci, erecti 20—70 cm alt., divaricato-ramosi, foliiscum arachnoidei, superne in costis aculeolato-scabri; folia radicalia necnon caulina inferiora petiolata, usque ad 8—12 cm lg., pinnatisecta, segmentis pinnatifidis ambitu ovatis, subtus setosiuscula; lobuli terminales oblongo-ovati vel obovati, 2-15 mm lg., 1.5-5 mm Ilt., folia media sessilia, 2-4 cm lg., pinnatisecta, segmentis lateralibus 1-—3- paribus oblongis 3-20 mm lg. 1-4 mm It. apicem versus paulum dilatatis, apicali majore, subtus sicut superiora brevisetosa, margine setosiusculo-serrulata; folia apicalia integra, oblonga, 8—25 mm lg., 2— — 623 6 mm It., obtusiuscule apiculata. Calathidia apice ramorum solitaria, in caule 7—40, floribuscum 16—21 mm lg.; involucrum late ovoideum vel subglobosum, 11-14 mm lg., 10-15 mm It., flavidum vel cinnamomescenti-flavidum, nitens; appendices phyllorum parte media solida flava vel laete cinnamomea, marginali paleacea flavida vel flavido-albida subintegerrima, in phyllis externis late ovatae, apice plane emarginatae, acumine subnullo, in mediis orbiculares, ca. 5—6 mm lg., in phyllis internis ellipticae, minutae; flores laete purpurei; achenia ca. 2.5 mm lg., olivaceo-brunnea, pappo ca. | mm lg. Fl. VI-VII. Habitat in declivibus necnon pratis montanis transcaucasicis. Typus. Transcaucasia orientalis, distr. Kuba, Kusary, 30 VI 1900. S. Grigorjev. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur. Affinitas. A C. transcaucasica Sosn. segmentis lobulisque foliorum valde latioribus, involucro calathidii haud complanato abhorret. 48. Sect. Polypodiifoliae Tzvel. sect. nova (subgen. Microlophus (Cass.) Hayek).—Plantae monocarpicae, vulgo biennes, pilis brevissimis papilliformibus et longioribus crispatis tectae; folia radicalia et caulina inferiora pinnatilobata, lobo terminali lobis lateralibus subaequali; folia apicalia vulgo prope basin calathidiorum posita; calathidia plus minusve corymbose congesta. Typus sectionis: C. polypodiifolia Boiss. 49. C. rhizanthoides Tzvel. sp. nova (subgen. Rhizocalathium (Tzvel.) Tzvel.). Planta acaulis vel subacaulis, viridis, plus minusve crispate pilosa; folia lyrata, longe petiolata, laminis vulgo solum prope basin paucilobatis (lobis 1-3 a quovis latere) saepe partim integerrimis. Calathidia vulgo solitaria, in centro rosularum foliorum posita; involu- cra 12-20 mm It.; appendices involucri plyllorum lanceolatae, retroflexae, marginibus ciliate, vulgo ciliis 1-5 a quovis latere praeditae; flores lucide flavi; achenia 4-5 mm lg., pappo 1—2 mm lg. Habitat in pratis montanis et declivitatibus lapidosis Georgiae australis et Armeniae occidentalis. Typus. Armenia, in declivitatibus lapidosis ad vallem lac. Sevan prope pag. Sevan, 13 VII 1957, n° 875, S. Czerepanov et N. Tzvelev. In Herb. Inst. Bot. Acad. Sc. URSS (Leningrad) conservatur. Affinitas. Species C. rhizanthae C.A.M. proxima, sed floribus lucide flavis (non luteis) et foliis solum prope basin paucilobatis vel integerrimis differt. ADDITAMENTUM Addition While the twenty-eighth volume was in press, the next volume of the Flora URSR [Flora of the Ukrainian SSR] was published, in which M.V. Klokov described the following new species of the genera Carduus L. and Cirsium Mill. 1. Carduus bicolorifolius Klok. in Fl. URSR, XI (1962) 566, 499, Fig. 99 (C. personata auct. fl. ucr. non Jacq.). 2. Carduus incanus Klok. ibid. 566, 502, Fig. 100. The author describes this species from Moldavia and puts it close to C. crispus L. 3. Carduus fortior Klok. ibid., 567, 504, Fig. 101 (C. acanthoides auct. fl. ucr. et ross. non L.). 4. Carduus tyraicus Klok. ibid. 568, 507, Fig. 102 (C. hamulosus auct. fl. ucr. non Ehrh.). 5. Carduus tauricus Klok. ibid. 569, 511, Fig. 103 (C. hamulosus auct. fl. taur. non Ehrh.). 6. Carduus attenatus Klok. ibid. 569, 517. According to the au- thor, this species occupies a systematic position intermediate between C. nutans and C. thoermeri Weinm. 7. Cirsium lipskyi Klok. ibid. 570, 533 (C. arachnoideum auct. fl. taur.). 628 INDEX ALPHABETICUS’ nominum specierum atque synonymorum plantarum in tomo XXVIII Florae URSS commemoratarum ACONOS AGAns iil s eesti atreteesosenione 228 SUSI SCOP ieee sseckvccesctecessexeas eset se 231 Acantholepis Kar. and Kir., sect......41 VA COED GD) CowSeCt..ccscsc desserts sss stsentecoss 150 Acaulia Petrak, subsect .................. 201 Aciculata Charadze, subsect. 197, 605 ACFrOCentron CASS. .........ccceseees 370, 493 Acrocentron (Cass.) DC., sect....... 497 — Kotschyanum (Heuff.) A. and D. NE GN Cute tote nes daa cece tact sanrceaan ances batt 502 — orientale (L.) A. and D. Léve........ a UR dae gale Sa ue ON 501 —salonitanum (Vis.) A. and D. Léve eco A a Vel a een 498 —scabiosa (L.) A. and D. Léve ... 503 Acrolophus Cass. 1.0.1.0 370, 512 Acrolophus (Cass.) Dobrocz., subgen . Fe PEE Se Rate HO nO 512 Acrolophus, sect. c) Maculosae Hayek al lc re a SA ee 517 — d) Arenariae Hayek.......... 520, 525 — 1) Ojlindraceae Hayek.............. 532 — m) Squarrosae Hayek ............... 535 — squarrosus (Willd.) Nevski ...... 535 Acroptilon Cass. ...........cesceeeeeeeees 343 — angustifolium Cass. ...........000 345 — australe Ijin oe eee 344 — obtusifolium Cass. 1.00... 345 — picris (Pall. ex Willd.) C. A.M .... SN ae eS ae enue eee Oe an tna 345 == FEPENS (We. ) MOG siacvcsresacventsooone ees 345 — SEFVAtuM CASS.......eeeeeeeeeeseeeteees 345 — subdentatum Cass. ........:11ee 345 Aetheopappus Cass. .................0.. 3 3538} — Caucasicus SOSM.. 0... cee 355 — Fedorovii Charkevicz.... .......... .354 — pulcherrimus auct.. ...........00-- 355 — pulcherrimus (Willd.) Cass....... 355 — — var. denticulatus C. Koch. ... 355 — — var. foliosus Somm. and Lev.... BU As Poa Seen} Ae MN, a NE 355 ocseahaceds cotusnuceacossisstseesusieatvae sess tetts 354 — — var. Raddeanus (Trautv.) Bordz. PN Si bese case ces vos cee oa fea Wes 355 —Raddeanus (Trautv.) Sosn. ex Grossh. SNR Rel el a ene 354, 355 —Vvedenskii (Sosn.) Sosn. ........... 354 Aggregata Petrak, subsect................. 69 AllboviamlzvieltiSeCtpe-trcccn-s seer 493 Allfrediay Cassioi cere. ccoteo- cases covicstcesacs 39 SUSE Ghetto csctecaccuelaeeecuse ca ae eas omnes 40 — acantholepis Kar. and Kir... ..... 41 ——cernua (I) (Cassicia cscs. cccssccres see . 40 — dolicholepis Schrenk ex Iljin.... 42 == IKarelini Wed be. . tercetessectsecs caer: . 41 — nivea Kar. and Kir. .............0. ee 42 — squarrosa Tausch... .............000 . 40 — stenolepis Kar. and Kir... .......... 40 — suaveolens RUPT.. ..........:0cee 42. — tianschanica Ruf .............:0. 00 41 Amberboa (Pers.) Less.. ........... ... 523 — sect. Amblyopogon DC. ............ 472 — sect. Chryseis {Cass.) DC... .... 323 — sect. Phaeopappus DC... ........ » 359 — sect. Xanthopsis DC................-. 474 — amberboi (L.) Tzvel... ............. » 331 — biennis (Ijin) Grossh.............. . 326 —sbucharical [ijinet eee eeecee eee ee 332 —= |PIAUCANAUCT + 5 Ts —— Cennuusebatiinvexs le dDaee. eee 40 = CAUPTFITAG INN Pe, Kotacsossmrppocceosonce0 6 130 ==SCINEHEUS) ATENES wcrcns: ecesceeeacecse-s-05s 36 =—=HGIMETE US Vista nnestctestcaseesonetees noe Bees, — — XC. nervosus C. Koch. ... ....39 — — var. amblyolepis Trautv....... 33 Carduus colchicus Alb. ............000+ 30 == COULLMIUS eal beeceeceececs Ol — hamulosus Ehrh... «0.0... cece 22 =e XC. MuUtansel. 2. ii cackercts. is . 39 — — xX C. onopordioides Fisch. ex MB RE AN ie ch aba Ns ie eS es 39 — — B. pseudocollinus Schmalh... 21 — helenioides L. ................. 165, 166 — heterophyllus L. ........000 165, 280 — horridus Adams.. ............... 78, 116 iy strixg ©. AL IM cine cece i tevessses oes 14 == ANCANUS KU OKMS. dot soesensqweakees.c. 621 ——WBNOGUUS I KOlAK. cece. ceccetses ssss2sase ccs 30 —- Karelini B. Fedtsch. ................... 42 — Kerneri Simk.... .............ccc:cceeee . 31 — Knorringianus Tamamsch.. ....... 33 627 — Koslovskyi Sosn. ..........0cc0008 00 30 = /aciniatus dba see ee -s eees 30 ——Wanceolatus Nx ei... nee ees. 135 — lanipes O. and B. Fedtsch........ 44 Se Winkla ceesasine os ladleoens: . 47 —=wlappaceus, MB ...:..3.f2c--csss» O21 carthalinica (Sosn.) Sosn......... 439 carthamoides Benth.............. ... 311 Caspia GTOSSHN. ....... cece ee 534 ECasprecallyin eee 328 caucasica MB. ex Willd. ....... + 399 Centauniuime Wace ee 37/1 centauroides auCt.... .......:c000 498 cerinthifolia Sibth. and Smith...306 cheiranthifolia Willd.. ............. . 399 — var. purpurea Sosn............. 398 cheirolopha (Fenzl) Wagenitz... 564 chlorantha Adams ex Ldb. ..... 399 Cichoraced SteVv.. ...ccccccccecceeeees . 301 — var. taurica Stev... ..........0. 430 circassica (Alb.) Sosn. ............. 425 ciscaucasica SOSN.. ...........00. . 428 coelolepis Fisch. and Mey. ex Heynh. TA teen ee eee. <4 Fae 501 coerulea Willd. ex Ldb.. ........ . 415 colchica (Sosn.) Sosn.... .......... 438 Colina Tes teen arenes: . 497 commutata Stank... .............00.06 446 conglomerata auct... 0... . 456 SE ytA UM) Se ay dete 450 COTLACEA "AUCt neers 506 — Waldst. and Kit. ................. 503 coronopifolia Lam... ............6- . 366 Crupina ls a). ee . 254 crupinastrum MOTIGS.. ............-+- 5253 cyanifolia POI... 1... 5 I? cyanocephala Vel... .....::cc00000 417 cyanoides Berggr. and Wahlb. ..416 CYANUSP LE. Wee ee ee 416 cyrtolepis Ldb. .............cccccee es 510 czerkessica Dobrocz. and Kotov .... snc Tusshgba Wee evecavehste dat oaeeeee ree 405 daghestanica (Lipsky) Czer..... 479 daghestanica Sosn. ex Grossh... 558 — var. lacerata (Lipsky) Czer........ Sa ee See ee 480 dealbataauct-5..220- 429 dealbata Willd... ...... ee . 434 — var. kachetica (Rehm. ex Boiss.) Sosnenvkc ee Se Bae 435 — var. lyrophylla (C. Koch.) Sosn. Venue Laat agate daa ee EO Oe 434 decipiens auct.. ........cceeeeee ee 446 declinata MB................... . 430, 439 Demetrii Dumb. ...............0000 0 332 depressa MBean ent eee 415 diffusa-Lam: scisicatcee ee 532 Dimitriewiae Sosn ............ 0. 491 donetzica KIOk.. ........c:cceceeeees . 547 Dubjanskii [jin. 0.0... eee 550 Edmondii Sosn.. ..........:cccee . 432 elatior (Gaud.) Hayek. ............. 455 elbrusensis Boiss. and Buhse... 396 epirotaHalacsyeeree see 408 erivanensis (Lipsky) Bordz. ... 475 exaltata Adams.. ..........0..:00 . 366 exsurgens SOSMN.. ..........::cseeeeee . 435 finitima BordZ... 2.0... 338 FUSChEri ‘AUCt He ecetree 394 Pascheni Will dite cece... eee 396 — x C. cheiranthifolia Willd. ..397 — XC. Huetii Boiss... ........... . 397 — ssp. cheiranthifolia (Willd.) Sosn. PPE AEM PEM TE Leet Ae A 399 — SSP. cyaned SOSN..........cccceeee oo sevasbustharistebucset atone 398, 410, 411 634 — — ssp. ochroleuca (Willd.) Sosn... s sataneent een eae a ETRE, | Oe 399 — — ssp. purpurea Sosn.... 396, 398 — — var. ochroleuca (Willd.) Grossh. RRR ccc ROE RR: Siete tie SBS ee 399 — — var. purpurea (Sosn.) Grossh.... ayswrerpetdind boca te, euler, A 396, 398 — Frickii Fisch. and Mey. ex Sosn. aeaee nan aoaaate A+ PoE ee 490 =F nitsehiiy Hayek. wer. sseeee eto <0 $07 — fuscimarginata (C. Koch) Juz. ..408 — fuscovariegata Stank................ 408 == PEOMMIGE TRONS cceconcccceecee oat SS) /5 Ol) =—t GERDETL AUCL..., © atiiersteretes chaos 549 === (GEM BE I SWEAY 0) ascde Boscecnasoadseescecene 550 — glastifolia auct.. .......cceceeee ee 339 Bee ey Le ricci ecb scnen ce ans tear ee RE 337 — — var. Biebersteinii (Jaub. and Spach) (Gubler. eae 339 — — var. intermedia (Boiss.) Gugler BORE DT PGA | einen eaves. oreniegds: 338 — — var. Tournefortii (Jaub. and Spach) sGugleraeee se. mood! — glauca Willd... «00... 326 =—WGIEANI ACU. hea: eee 505 — Glehnii Trautv. oe 509 — — var. bicolor Trautv.. ........... .509 — Gontscharovii Iljin... «0.0.0.0... . 382 — grandiflora Pall.. ............0 318 — Grossheimii Sosn. ............006 568 — Gulissashvilii Dumb.................. 531 — hajastana Tzvel.. .......... eee . 383 = JEAN INC) Oeg6, ansccacaceoe: ravtcooese 502 — hololeuca Boiss... ........:ceceeee 564 — holophylla Socz. and Lipat ............. 1, 5 BRA |) Spee Vek etal 426, 610 ——WNOTLOGUM OR Aer eee eeseeseeeeecee ee 417 — Huetii Boiss... 0.0.0... eeeeeceeeeeeee 410 Centaurea hymenolepis Trautv. .... 433 — hypanica Pacz... ....c.:.c.ccseee 544 — hypoleuca DC... «0... 424 — Hyrcana Grossh.. ........eee . 460 — hyrcanica Bornm...............0. 0 460 = WIDENICA MINE Veaycrene ceeeretr-ceceeene =: . 574 — — XC. salicifolia MB. ............ 459 — Iljiniana War... eee . 554 == Lilt (Cer Keo, cece cep ncnceececcnnce . 419 — incanescena (DC.) Sosn........... 473 — Intacta LAD.. .........ceeeeeneeeeees 7550 — integrifolia auct... .........ccceeees 448 — — B. Fedtsch. ......... eee es 315 = tN Ns) Ro . 473 — integrifolia Tausch.. ................ 504 — inuloides Fisch. ex Schmalh.... 580 631 = Wacea auct, ..s).0%..0s. tae 446, 448 Se ACCA, RRA eld eee: 444 = A GHephny glade ).cxtcotes 451 — — xX C. pseudophrygia L. ....... 446 — — ssp. angustifolia auct......... 446 — — ssp. angustifolia var. integra sub- var. pannonica (Heuff.) Guglet...... REALIGN. aed RM thee Shes satus 446 — — ssp. angustifolia var. pannonica (Heuitit®) Hay cker ke. hate 446 — — ssp. eujacea Guglet........... . 444 — — ssp. genuina (Wimm. and Grab.) DOStaly) Pate eee A ceasks.eeas:. . 444 — — ssp. jacea (L.) Gremll... .... 444 — — ssp. oxylepis auct................ 449 — — ssp. pannonica (Heuff.) Soo and Javon Kaliieieted eae el eters ise 446 — — var. commutata auct............ 446 — — var. decipiens auct... ......... . 446 — — var. lacera auct..........e . 446 — — var. oxylepis auct................ 499 — — var. pannonica (Heuff.) M. Pop... TERR Seedy, ee eee 446 — = Var. pratensis auct:iy:::.2.2:.. 446 — — var. salicifolia (MB.) Briquet... CELLONE leh caalth SAIS bl ele 458 — — Q. genuina Wimm. and Grab.... 242 TM UE Mnenerae, ORES 8 444 — — 6. livonica (Weinm.) Weinm. nel Meats, cahane Eevee een OB, vat 446 — karabaghensis (Sosn.) Sosn..... 437 — karatavica B. Fedtsch... .......... 321 — karduchorum Boiss.................. . 473 — kasakorum Ijin... wee 383 — Klokovi Oppermann ex Klok... 543 — kobstanica Tzvel. ......... ee 483 — Koenigii Sosn...........ccceeeees . 492 — koktebelica Klok............. . 528, 617 — Kolakovskyi Sosn. ........ .. 429, 611 — konkae KIOok.. ........ eee . 545 — kopetdaghensis Iljin.. .............. . 420 — Korshinskyi Mljin... .........0.-0060 386 — Kotschyana Hevff... ............0.... . 502 — Kryloviana Serg ......... oe 471 — kubanica Klok. oe 518 — Kultiassovii Ijin... 384 — lancifolia Sieb.. .......... ee . 564 2“jlanigera,.DC........scoseud adetasneunts 396 — lasiopoda M. Pop... .......... 385 — latiloba Kok... 2... 557, 620 — latisquama DC.. ..0......:cececeeee ee 498 — Lavrenkoana Klok.. ................. 526 — Ledebourii (Korsh.) Klok. ..... 472 — leucantha Adama..................-++ 399 632 — leucolepis Ldb............... .. 548, 553 — leucophylla MB... ..............:00 429 — leuzeoides (Jaub. and Spach) Walp. Sea tia oh Re nines tee eee ME 487 — livonica WeInM...............000eee 446 — longiaristata Illar....... . 554 — lyrata B. Fedtsch. .........cc 314 — macrocephala Muss.-Puschk... 333 — maculosd auct .........06. 0: 518, 519 — maculosa Lam.. ................ 512, 517 — — ssp. micranthos (Gmel.) Gugler pe Pegs reneee | AY). bee eannt oie 4528-2 519 — — ssp. rhenana Guglet... ........ 517 — — ssp. rhenana lljin.. ............ . 518 — Majorovii Dumb.. .............00.. . 520 — Maleevii (Sosn.) Sosn.............. 428 — maramarosiensis (Javorka) Czet..... Hesbscoe, 3s AO DO I SRM 392 — margarita-alba Klok.............. ... 544 — MAFZAritaCeA AUC... cece cesses sa dinitoaceiact AAU Bete 544, 547, 548 — margaritacea Ten. ............. 543 — maris-nigri SOSMN. 1.0.00... eee 431 — Marschalliana auct. .........000cc cee disese Pd. ARIE. 3 465, 467, 469, 471 — Marschalliana Spreng. ............... 466 — — var. bessarabica Trautv... ... 466 — — var. intermedia Schmalh..... 470 — — f. Ledebouri Korsh.. .......... . 467 — — f. vulgaris, Trautv. ex Korsh.... verve RNS, SRO), wheel easter . 467 — matthiolifolia Boiss... .............. . 408 — membranacea Lam. ...............+- 318 — mentiens CZef.. .........:cceee . 416 . — meskhetica Sosn.. «0.0.0... . 439 — Meyeriana Tzvel.. ................0 473 — micrantha Gmel. var. recurvata DUMB xi5 easton. PA 519 — micranthos Gmel...............08 519 — minima (Boiss.) B. Fedtsch. ... 349 —— Modestinkedtt:.4.e seas 381 — mollis auct.............. 392, 401, 402 — mollis Waldst. and Kit... ......... 393 — — f. maramarosiensis Javorka 392 635 — — f. ramosa Czako.................. 392 — monanthos GEOTG}..........0ee 318 — montana auc... eee . 394, 408 ==! montana Als. A: deceive. See 391 — — ssp. mollis (Waldst. and Kit) Gugletese cere ees 393 — — — var. ramosa (Czako) Dostal. soveseadaesccdesugedes oa eee Te 392 — — — var. typica f. maramarosiensis (Javorka) Dostal. .............0.. 392 — — var. cyanea auct... oe 410 — — var. purpurascens auct... ... 398 — — Q. cyanea auct.......... . 394, 404 — — Q@. cyanea DC............. 393, 402 — — B. axillaris Schmalh... 402, 404 — — B. axillaris f. ochroleuca (Willd.) Solita associat cierto ere . 399 — — B. nigrofimbria C. Koch.... 394 — — B. purpurascens DC........... 396 — — y. albida DC.... cee 399 aS sO citning, DC Aenean aen 399 — MOSChAILA AUCt.. .......cccceecceeceeeees . 326 Ce il ete ON a eng 6S) — — B. amberboi L. ........ce. 331 — — B. glauca (Willd.) MB....... 326 — nana B. Fedtsch... ..............0000 B99) — Nathadzeae Sosn. ................00025: 490 — NIZrA AUC. wo... oe 450, 453 — NIZreSCENS AUCt... .....eereeseeeeee 446 — — ssp. eunigrescens var. rotundifolia subvar. salicifolia (MB.) Gugler... LANGE). \ivnerben ranean’, | cule saz: 458 — — B. transalpina C. Koch.. ... 453 — — y. vochinensis C. Koch... .. 458 — nigriceps Dobrocz................05 452 — nigrifimbria (C. Koch) Sosn. .. 394 — nigrofimbria auct............0.004. 410 — nigrofimbriata JuZ.. ...........0.. . 408 — nikitensis Wat... ..........:.csceeee 552 2S{Ninde Juz... .... ee. eee 554 =tnitens,, MB ..As.1ieemais . 342 — nitida B. Fedtsch...............000 319 — Novakii Dostal... 0.0... eee 401 — novorossica Klok.. .............00 . 556 — ochroleuca Willd.. ..............00. 41399 — — f. cheiranthifolia (Willd.) Gugler Se daitecatecduocs ME Res tos casa hE Bee aS 399 — — f. genuina Guglet... ............ 399 — odessana Prod... ..........eeeeee $523 =foltensis; Sosn\.<..:...9:.02 4.22220 570 = PORIENtAlIS, es eesrns sees eee . 501 — — XC. apiculata Ldb... .......... 512 — — var. brachylepis Fisch. and Mey. Laschevau oss ai Auouaneassscs io OA ene ee 501 — — var. calocephala Schmalh... 502 — — var. macrolepis Fisch. and Mey... WH sack saatsvpemadts sccce oR NRE ae ae 501 — — var. rubescens (Bess.) Schmalh... ED sleaanes adtveenceuaeeyeeevndeey tees ANAS 502 — ossethica Sosn.. «0.0... cece A505 — ossica (C. Koch.) Boiss... ....... 344 SHOVING AUCts.-cc-c3 20-02. teh ste o eases . 526 +*tovina»Pall.:...2iseek. gee 530 == VoxylepisviaUct = ..20Rtitcstene) ee 449 633 — Paczoskii Kotov ex Klok........ . 549 — pseudovina Illar............. 513 — pallesens ssp. iberica var. genuina — pterocaula Trautv... .............0..- 338 (Guglen)vArenesteeee 2 2)..:.) < 574 — pulchella Ldb... «0.000. 369 — — — var. iberica f. genuina — — Q. viminea (Less.) DC....... 370 GugleRy. eee Ee He A eas, 574 — — 8B. glabra DC. .. ..... ee. A370 — pambakensis (Sosn.) Sosn....... 437 — pulcherrima Willd... .............. 355 — pannonica (Heuff.) Hayek. ...... 446 — — var. intermedia Trautv... .... 355 — parviflora BESS. ........:cccceees 532. “S=¥pulehra Dee kee . 417 — pauciloba Trautv. ........ cee 427 — pygmaea Adams ex Ldb......... . 415 =P PechowAll bier... cctsen eee: 493 — pyrroblephara Boiss................ 464 ==spectinata AUCts.. Li i8h «acacia: 450 — Raddeana Trauty .......... ceo 355 SSS pelia AUct>,.. 20 edth teehann: 498 — Razdorskyi Karjag... ................. 386 =——ppellucidawRechy fess a.ecces js. 322 — recurvata C. Koch.. ....... 510 — pergamacead auct... ......cceceeeees 487 = reflexd AUCt... ccc ceesteecescseeees . 500 — pergamacea DC... ....... 487 = reflexa Lamlty.2: sissies ee 499 — phaeopappa (DC.) Sch. Bip.. . 363 — — X €. adpressa Ldb)..:...... .-. 500 — Phaeopappoides Bordz........ .... 486 — — var. grandiflora Grossh.. ... 500 — phrygia auct ........... 453, 456, 462 — — var. macracantha C. Koch...500 =—ephnyeia i.e vane docs deomtens 449 SSUPEPENS We fo ciieiin etree ece eeebeaaeee . 345 — — x C. pseudophrygia C. A. M.... — Reuteriana Boiss...................... . 396 Se 2 Neda oer Rb is ak . 457 — rhenana Boreau. ........................ 517 — — ssp. austriaca (Willd.) Gugler. .. — rhenana Dumb.. ...........:0:000 . 518 RO PSR Atha Mba Re crete a 450 — rhizantha C. A. M................. 567 —— ssp. pseudophrygia f. intercedens — rhizantha auct .........cccceeeeee vee 568 subf. carpatica (Porc.) Gugler 451 — — C. A. M. xX Tomanthea — — var. subuniflora MB....... ... 454 phaeopappa (DC.) Takht. ex Czer... — phyllocephala auct.. ...........0. OS, «sss RRR DEI peace ane eee 364, 568 — — var. persica BOISS.............. 578 — rhizanthoides Tzvel... ...... 568, 620 — — f. Belangeri Boiss... .......... 578 — rhizocalathium (C. Koch) Boiss. .... — phyllopoda Iljin... .......... mete toa BBS) |S RO Se eae ae 569 = Pichler Boissisanste cee ee 408 — rigida ANGIZ... .......cecceeeeeesees 507 = jpicris, Willd fee Ree 345 — rigidifolia Bess... oe 502 Se PM ANGUS OISS y.10-.sseeeeeee eee eae . 416 — rubescens Bess. ............c:ceeee 502 — pindicola Griseb................ 396 — rubiflora Ilar oe oe 498 — pineticola Ijin... eee . 548 — Ruprechtii (Boiss.) Czer.......... 480 — pinnatifida Schur. .............. 392 — ruthenicad auct... .......ccccecceerceeees 383 — plumosa Aitch. and Hemsl...... 307 — ruthenica Lam... .............. cee 380 = b earpatica) Porc. ee 451 — — var. angustiloba Taliev... ... 381 — polyphylla LAD.. ......ceeceeee . 334 — — var. bipinnatifida Trautv.... 381 — polypodiifolia Boiss.... .............. 563 — — var. genuina f. angustiloba — — XC. Stevenii MB. ............... 565 Ors ney ee ented a ee 381 == PFAtensiS\ AUCt. ..hcsesteescechceee ces = 446 — — var. hispida Korsh.......... ... 386 — proto-Gerberi Klok. .................. 546 — — var. lyrata Trautv................ 381 — proto-margaritacea Klok... ....... 543 — — var. villosa Taliev... ............ 386 — pseudoaxillaris Stef. and Georg.. ... — salicifolia auct... ............. 453, 462 a READ Sokre OAR chcee eect Pee 401 — salicifolia MB............ccee oe 458 — pseudocoriacea Dobrocz.. ........ 506 — — var. glabrata (Litw.) Grossh.. ... — pseudodeusla auct ..........0000. coe We! URI te MON Aeerals subd Shavers Reet ree 453 636 Centaurea pseudoleucolepis Kleop. 542 — — f. glabrata Litw .......00..... 453 — pseudomaculosa Dobrocz.......... 518 — — a. elongata C. Koch.......... 458 — pseudophrygia auct.. ............. . 458 — — B. abbreviata C. Koch....... 453 — pseudophrygia C. A. M. ........ 456 — — y. intermedia C. Koch. ...... 453 — pseudoscabiosa Boiss. and Buhse... = salina Benth yet eee 313 Or as PMN ak Ne RUN) ea Du 508 — salonitana ViS............ccc00. .. 498 634 wu Casadpressamludbyey ane 498 — x C. orientalis L. «0.00.00... 498 — — var. macracantha (DC.) Boiss. and Heldr.23e).f. 280.1). weasels. ..2., . 498 — var. purpurea Ferm. ex Stojan andy Stetien.23.i Ly. Ret iNeed 498 salviifolia (Boiss.) Sosn........... 432 Sarandinakiae Ilar...........0...... S55 Savnanicavklok.2eehs.. .e 524 SGADLOSAL AUCH es.. nee 506, 507 SCabiosaplennd eee. See 503 — var. adpressa (Ldb.) DC.... 507 — var. angustata Ldb............. 507 — var. apiculata (Ldb.) Ldb... 506 — var. coriacea Ldb.............. . 506 — var. integrifolia Korsch... ... 504 — var. melanocephala Rupr... 504 — var. spinulosa (Rochel) Boiss... ete te Re Nhe ance ade A . 508 — var. stereophylla (Bess.) Ldb... saeaesucboeeeubildeomoces el eee ane E 511 Schelkovnikovil Sosn. .............. 412 schemachensis Sosn. ex Grossh. ... vanpat anther cen JOUR, DRIES 558 schistosa Sosn .............. ... 433, 611 segetalis Salisb.. ..........:cccee . 417 SS MMIJUS CaM Zeer eee treneceers 5 Ol Serebrowskyi Sosn... ...........006 413 Senet MlokA.. 2h . 471, 614 STEAM OUIEIN Comrectiee erette gnmtetererseceere cee 564 serratuloides Georg... ..........0 313 SESSHLUSMALLC tA wee eee te oe 567, 568 —— var. integrifolia Bordz. ex Grossh. aaa. Tanto mcs sees eck RE DOO 568 Secusanaa@haix. eater ee 408 Osh) == 408 — var. fuscomarginata (C. Koch) SOSH ROD. TE 408 SEVANENSIS)S OSE ttle. .0e 436 SUD URUCHMAUG taapestsseeo ee 465, 470 sibiricael... RRS. DAISY 2 469 SUDIFICHANIB ee ee ees. Tee 466 — var. diffusa Korsh............... 470 — c¢. Marschalliana Schmalh.. 466 Si GUL AAU Cla poietic Hee. Mee 571 simplicicaulis Boiss. and Huet....... we Pen ite Secictnate ayes waite iatdiins ato etch AO 491 sokolensis (Pawl.) Czer. .......... 401 somchetica (Sosn.) Sosn...... ... 438 Sophiae Klok...............0.. 521, 615 Sosnovskyi Grossh...............:0 . 500 Sosnovskyi Wjin.. .......ccccccccceseees 327 solstitaliswkicknn2L 6: SO 571 — ssp. eu-solstitialis var. Adamii (Wolld))iGuelent. 242.2205 0. 573 — — var. typica Guglet.......... S22 — B. Adamii (Willd.) Heuff.... 573 spectabilis (Fisch. et Mey.) Sch. Bip sic, PROG LOS), BI 361 SPINUIOSA AUCL... ...ceecccerceeseeseees . 508 spinulosa Rochel... ..............0... 507 splendens Lang and Szovits. ... 543 SQUQITOSA AUCL... .occeceeeeeeeeceees 2 SS) squarrosa Willd... .............0.00. 2 5) Stankovii Wlat.:.. .....ccsecccts.c0.e0-00 $53 stenolepis auct....... 454, 455, 456 stenolepis Kerm.... ........::cecee 458 stereophylla Bess... ............00 511 Sterilis#Steve....221..e eee 553 — /X: C.,. machilosal ON eeeaeae 554 Steveniana Klok.............. WS2ZTMONG — x C. diffusa Lam... .......0.... . 528 Steven} MBE SOR ee 7 564 SLGLCLO GA Chines. pe cvetcemeavecetae 404 stricta Waldst. and Kit.. ......... . 402 — ssp. Angelescui Pord. ..... ... 406 — var. Angelescui Grint .... .... 406 strictissima Boiss. and Buhse.. .293 — suaveolens WAIId... .......-ccccccere 331 — subjacea (Beck) Hayek... ........ 446 — substituta CZer. 0... eee 448 — sumensis Kalen. ................00000 0 467 — SZOVItiSiANa AUCt... oc... 563 — Szovitisiana Boiss.. ................. 562 — tadschicorum Tzvel... .........0.6. 368 SS WalievilykKleopy.. Ae. eee . 386 > HanaiticaKilok.. 220... oe 404 — taochia (Sosn.) Sosn.. ............. 435 = tenuiflora, D@.ae ee ee 527 — ternopoliensis Dobrocz.... ........ 401 — Tournefortii (Jaub. and Spach) Wal Disc: sess tvesres.:: fk. BE 37, — transcaucasica Sosn. ex Grossh....558 — trichocephala MB ................ 0... 462 — — Q. angustifolia Fisch. and Mey szaiyeoctguaesstenst ates Ma ES Ded Oe 462 — — B. latifolia Fisch..........0.0.. 460 — trinervja Steph... 0.0.0... . 483 — — x C. Ruprechtii (Boiss.) Czer... sasesnioucec Me a: Re 485 == EriumifettiieA lage. eee 400 — — ssp. variegata var. cana auct.... G cvattivecss AL, BOE BE 408 — — — var. cana (Sibth. and Smith) Stef: and "Georg. 222 eee ee 411 — — var. cana f. pleiocephala (Bornm.) Stef. and Georg... .... 413 — — — var. euxina Stef and Georg... Bie2 IMG. SE A Ae 408 638 — — — var. fuscomarginata (C. Koch) DOstall.o:. Shee. . 404, 408 — — — var. pleiocephala Bornm... 413 — Troitzkyi (Sosn.) Sosn......... ... 427 — tuapsensis Sosn..... 0... 431 — tuba Somm. and Lev............ .... 334 —Muranica lpn we a.. eee 330 — turgaica Klok... 2.0... . 470, 613 — — < @)/Sergil Klok 472 — turkestanica Franch... ............... 387 — tzebeldiana Woron.................. . 459 —wgamica Vjimesy see eee 387 — umbrosa Huet ex Reut.... ........ 417 =MiniflonarLcs ee Se 319 —— UU LOT AP aliagars. erence eee 282 ——sustulata AUC ines » SID =sustulata, DEG... eee 5 oyh0) = aViaNK OVAL O Ks eye te ee corecetce 552 —— VatlegataLam.cin-tss.ccsees eee 408 — variegata auct........ 402, 404, 408 == Vicina LipSkys.ii...:.:...cesesncoster 530 == VIMINEAMLESS He yaw wees cee 369 — virgata var. squarrosa Boiss. .... 535 — vulgaris var. jacea (L.) Godr....444 — Vvedenskyi M. Pop............00005 . 349 — Willdenowii Czer. ......... .. 398, 608 — — XC. cheiranthifolia Willd. 399 == Wolsensis# DCH tse Ee. 522 — Woronowii Bornm. ex Sosn.... 413 — — ssp. eu-Woronowii Dostal... 413 — xanthocephala (DC.) Sosn. .... 476 — xanthocephaloides Tzvel.......... 476 — zangezura Grossh.... «0.0... 411 — zangezuri (Sosn.) Sosn.... ........ 436 — zauandica (Sosn.) Sosn............ 435 Centaureinae O. Hoffm., subtrib... 240 Centaurieae Less., subtrib............ 240 == (Gass tribes. 4.a een 240 Centaurium Cass. ... ........0 370, 377 — sect. Microlophus (Cass.) C. Koch.. hee acess 2 LS a a SER ee ie 560 — behen (L.) C. Koch.. ........0... 5 Oy — ruthenicum (Lam.) C. Koch.... 380 Cephalonolos Neck................- $2, 209 Cephalonoplos (Neck.) DC., sect......... Re ae ARTO ICEO A ree Oar OEE 209 — ochrolepideum Nevsky... .......... 212 — setosum (MB.) Kitam... .......... . 210 Chamaeleon DC., sect... .........00 150 Chamaepeuce DC..........:cecceteee eee 219 = Gynaroides IDC ara. . 137 == (J Ley 0¥,)) yo) C0 ong eeescenncesecetooscon: 139 — echinocephala DC... ...........00.++ 220 Chamaepeuce macrantha O. et B. Fedtsch SUD eh eg teees eee este SN, 49 Ser SCATENK es asses oee eeetees eee 138 — — B. bracteata Rupt. ............0000. 140 = SINUALA ATAU) artes cose cece etek «s 137 — Wachtangii Grossh. ..............:0008 220 Chaptalia anandria Spreng. .............. 593 > WVRALa SPLen eee. ase e ees ees: 593 Characopappeae Sch. Bip. Div. Centauri- NEGESCHMBIP Hts. ceeccoteesceveeo eee ov ee: 240 — Div. Serratuleae Sch. Bip. ............ 240 Chartolepis Cass. ..............:csceeees 335 — subgen. Cheirolepis (Boiss.) Jaub. et Spacht: ... 28s. ee 356 — subgen. Euchartolepis Jaub. et Spach. i Saas ae aca ts ee broe MEE Sore eee 335 — Heeberene auctites a eet 338 — Biebersteinii Jaub. et Spach. ......... 339 — CAppadOCica auct. ..........sceeeeeeees 358 — finitima (Bordz.) jin. ................... 338 — glastifolia AUct. ........scscecereseees 339 — glastifolia (L.) Cass. «0.0.0... eee 336 — intermedia Boiss. ............:::eeeeee 337 — Kotschyi (Boiss. et Held.) Fenzl.... 358 = ilyrata/Boiss i. ceece et a 336 — macrocephala (Muss.-Puschk.) Takht. his IE aE hae Sade decaes Retereeetee 334 — aig (irautv.)|Czen cess. 338 == Salipna @akKoch stance ere 336 — Tournefortii Jaub. et Spach. .......... 337 — Umesaoi Kitamura. .............:0e 336 Cheirolepis Boiss. .............:..:ceeeeee 356 — drabifolia (Sibth. et Smith) Boiss.. 358 — Kotschyi Boiss. et Heldr. ............... 359 — persica BoiSS, «0.00... eeeeeeeeeseeeteees 358 Chiryseis Cassiiaui ntice cette cate. 323 — glauca (Willd.) Cass. ......... 326 == OdOrALd CASS I aivac.s:ccettesien scsteaeess 331 Ciliata Petrak, subsect. ...................66 128 == 2 \CiliataPetrak Ane ee 128 — 3. Fimbriata Petrak. .............000 128 — 1. Serrulata Petrak. ....... 100, 109, 128 Cirsiicarduus lutetianus Arénes. .......... 39 Cirsium Mill. emend Scop. ................. 51 — Subgen. Cephalonoplos Nakai. .... 209 SESECt He ccccden cA Sd 149 — sect. Acarna DC. ..........secceeeeneeeeeeee 218 — sect. Anacantha Iljin. .................... 223 — sect. Apalocentron (Cass.) Kitam. 160 — sect. Breea (Less.) Boiss. .............. 209 SAIsubSeCtts 28 TURAL A Bree ae 164 — abkhasicum (Petrak) Grossh. .......... 86 — Acarna Moench. ..............:c1eeeeee 218 636 == /ACAUIE. AUCt. 0... detilomeedseaev he 204 SiO se teste revateteatk tes 201, 203 == ACAWLEN (IZ. SCOPse eres sean ncuera 205 — — VAT. ACAUITE. oo. eeececscceseetseetteeees 205 — — var. genuina Briq. et Cav. ........ 205 — — B. sibiricum Lab. .........0.ee 201 — — y. Gmelini DC. «0... cess 201 — adjaricum Somm. et Lev. ................ 79 — — ssp. nutans Petrak. ...........0::008 79 —aduncum Fisch. et Mey. ex DC. ...... 97 —— XC. ciliatum (Murr.) Moench... 134 = Tatiine NAUSGM) .eert.ceapsaeeoe cee eet 166 —aggregatum Ldb. .............::eeeeeeeeees 69 — — var. macrophyllum Alb. .............. 70 — alatum (S.G. Gmel.) Bobr. ............ 192 — Albertii Rg]. et Schmalh. .............. 145 =A bertil: WANK ccccersesceepeszecasucecessp< 145 — Albowianum Somm. et Lev........... 189 — Allionii Sch. et Spenn. .................. 205 — anatolicum Petrak. ...........e eee 102 — — var. microcephalum Grossh. ..... 102 — — var. ramosum Grossh. ............+++ 102 =SapiculatumDC: eee. sae eeee er 190 — — ssp. glaberrimum Petrak. ......... 190 — arachnoideum BOSS. .........:.1ceeeeees 96 — arachnoideum (MB.) MB.............. 110 —— var. incanum Lipsky. ............... 111 — argillosum V. Petr. ex Charadze. ... 123 =/argunense DC. 3. cizsaxk a iccameren 210 —— faery io Dl Gxisneseesceccne nc Gs cpscbceooce. coe 200 ==(ArVvense auctor Ax. .2eet, sansseeeee 210 — arvense (L.) Scop. ....eeeeeeeeeseeeeees 213 == arvense Vieth., .......0-.ereees. Leet 211 Sa SS PMCRVEMS Cn see. eee tenaes Jesse bees 213 — — ssp. Hookerianum Petrak. ........ 212 — — ssp. incanum Iljin. ................0.+. 212 — — ssp. rubricaule Novopokt......... 213 — — ssp. setosum Ijin. .............0.020++ 210 — — ssp. vestitum Petrak. ................. 211 — — var. commune Beck................-- 214 — — var. commune f. incanum Beck. 211 — — var. horridum Koch. .............0. 214 — — var. horridum O. et B. Fedtsch . 212 — — var. incanum O. et B. Fedtsch... 211 — — var. mite Koch . ...........eeseeeeenes 214 — — var. obtusilobum Beck. ............ 214 — — var. setosum Koch. ............:20 214 — — var. setosum Kom. ........0--eeee 210 — — f. horridum Vieth. ............ 213,214 — — f. mite Vierh. ...............00005 213, 214 — — c. setosum Schmalh ................. 210 —— a. horridum Wimm. et Grab. ... 213 639 2B mite Wimm. and Grab.... 213 — 6. incanum Ldb..........0.. 0... 211 — 6. vestitum Wimm. and Grab... iM. Ove eter ee, Meet ccna. 211 — Y. integrifolium Wimm and Gralbigiie tt ate essen hn hos. ees co 210 —— V. SCLOSUIM We GD nen. ne eee . 210 — y. setosum f. subulatum Ldb... . bcs sracnss sacs Aeon Ao aU, Camere A weal 210 asiaticum Schischk. .............. ... 176 aspinellum Sojak.... ...c..ccccccceee 96 badakhschanicum Charadze.. .......... sassseaausdudaaiea seal ome ea Raunt . 144, 604 balkaricum Charadze................ 177 Biebersteinii Charadze.. ........... 170 Bornmiilleri Sint. ex Bornm. .... 99 bracteatum Link.. ...........00...66 . 216 bracteosum) DG). oe eee . 93 brevipapposum Tschern ...... .... 204 Buschianum Charadze.. .... 84, 602 — f. aciculare (Somm. and Lev.) Charad Ze.) .........:.:s88..- ete eee ee 84 calvescens Sojak ..........:eeee oe 97 canum (L.) All setae e- eee 169 Cantum (MIB ick ee eee 170 — Y. pannonicum Schmalh.... 171 caput-Medusae Somm. and Lev ..... withastcieeseascesas tients. oe. beets ene 88 carthamoides Link. .............000- ++ 311 caucasicum (Adams) Petrak..... 78 — caucasicum ssp. cladophorum Petrak wna saaetelnndPy atts cemmeneiee Ot bass naomi 78, 79 — ssp. nutans Petrak............ .... 79 — var. Wwedenskyi Petrak....... 79 cephalotes Boiss..............006 e+ 114 chlorocomos Somm. and Lev.... 76 ciliatiforme Petrak ................... 109 ciliato-affine BeSS...........0:000 + 28) ciliatum (Murr.) Moench..... ... 130 Ciliatum SZOWIES......cceccceee vere 133 ciliatum (Murr.) Moench x C. caucasicum (Adams) Petrak .... 134 ciliatum hybridum Fisch...... ... 119 — ssp. Szowitsii Petrak....... ... 133 — var. xantholepis Petrak .. ... 133 — B. microcephalum Lab .. .... 130 congestum Fisch. and Mey........ 90 — B. consanguineum (DC.) Boiss. bake ante SRR acing essa ee ee 91 consanguineum DC..............6. ..6 91 coryletorum Kom .............0:. 005 174 cosmelii (Adams) Fisch. ex Hohen seth sigscviddatisc sabe acces SONG stealer 107 — x C. caucasicum (Adams) Petrak “Su naga a an Atta mtenabaa abba nare ce at 108 — — XC. ciliatum (Murr.) Moench.. 2 ae, Ce ES SN TE 109 — cynaroides (Lam.) Spreng... ... 136 == Cynaroides MIBY. eee cere 137 — czerkessicum Charadze.... 72, 602 — daghestanicum Charadze...... ... 119 — darwasicum Ijin .............00. oe 224 — dealbatum MB..............000 oe 168 — decussatum Janka ................ 0... 277 — depilatum Boiss. and Bal .... ... 185 = MACSCVLOTUM AUCL «err-s-.ce cee: 191 eer EISCNe EX IEIMK, rete oe ose: 192 — var. sinuato-lobata Trautv...192 — var. subintegerrima Trautv ..192 dimorphum Petrak ex Grossh.... duriusculum C. Koch echinocephalum MB.............. ... echinus (MB.) Hand.-Mazz .... elbrusense Charadze............. ... elbrusense Somm. and Ley .... ClOdESHAUCT not ee 192, clodess MBit ates sees 191 Tet Ue Se a 2 MER ae) niet OR 2 149 Petrak 196 — x C. obvallatum f. obvallatum Petrak 196 — x C. obvallatum f. subelodes Petrak 196 - — var. glaberrimum Bornm ... 190 -- — var. sinuato-lobatum O. and B. Fedischt tact eee SZ — — var. subintegerrimum O. and B. Fedische ae yeaa ean 192 — — B. indivisum DC......000.. 0: 195 — — f. setigerum Kryl............ ... 192 = erlODNOrUm AUCte seca eee tee 127 =— efiophorumy SCOpins eee 89 — — ssp. decussatum Petrak var. POlOnicumyPeake see 127 — — var. platyonichium auct. .... 127 — — var. spathulatum auct......... 125 — — var. turkestanicum O. and B. Redischetee ss tate eee 124 — — ¥. turkestanicus Rgl. ...... ... 124 — erisithales (Jacq.) Scop........ ... 163 = CLYLINOLED ISAC Us sere teee eee te 85 — erythrolepis C. Koch. ................. 81 = HESCULENLUTULAUCUS pisses crete eee 205 esculentum (Sievers C. A. M. .201 — ssp. caucasicum (C. A. M.) Petraletere eae. rere tan 203 640 637 — — ssp. Frickii Petrak. ............. 204 — — var. acaulis Trautv. ............ 201 — — var. caucasicum Grossh. .... 203 — — — f. caulescens Grossh..... 203 — — qQ. sibiricum C. A. M. a) Acavle C. A.M 201 Chamaepeuce macrantha O. and B. FeGtsen ec BES 49 i Schrenk ee 138 — — B. bracteata Rupt. .......... ... 140 == SINUALAMMUTAULV secre. tceet- cence eee 137 — Wachtangii Grossh. ............. ... 220 Chaptalia anandria Spreng. ......... 593 == Lyratal Sprenger eee ee 593) Characopappeae Sch. Bip. Div. Centaurineae Sch. Bip........ ... 240 — Div. Serratuleae Sch. Bip....... 240 Chartolepis Cass. .......0...c ee 335 — subgen. Cheirolepis (Boiss.) Jaub. and "Spach isin eae ae 356 — subgen. Euchartolepis Jaub. and Spach hie ein Slwetarnitiees 385) — Biebersteinii auct.........00. 6 338 — Biebersteinii Jaub. and Spach...339 — Cappadocica auc... 358 — finitima (Bordz.) Ijin. ............. 338 == ‘elastifoliavaucives..28- 339 — glastifolia (L.) Cass... 336 — intermedia Boiss. .........0...... 0 337 — Kotschyi (Boiss. and Held.) Fenzl. 1 ES RES ESC 358 = lyrata"Boisse eee eee 336 — macrocephala_ (Muss.-Puschk.) Pale tea er a PRN oe 334 — pterocaula (Trautv.) Czer. ....... 338 = saligna\@? Kochsts2232..2855% 336 — Tournefortii Jaub. and Spach....337 — Umesaoi Kitamura. ........... 336 Cheirolepis Boiss. ...........0..0. 2. 356 — drabifolia (Sibth. and Smith) Boiss.... obec AS OCR ois asco toot 358 — Kotschyi Boiss and Heldr. ...... 359 == "PETSiCa BOISS ieee 358 Chiyseis (Casss 2 ee 323 — glauca (Willd.) Cass. .......... 0... 326 =="odorata Casson tee 331 Ciliata Petrak, subsect. ..........0... ... 128 =—*2) Gilidta’ Petrake ee ee 128 — 3. Fimbriata Petrak. ............. ... 128 — |. Serulata Petrak... 100, 109, 128 Cirsiicarduus lutetianus Arénes...... 39 Cirsium Mill. emend. Scop............ 51 — subgen. Cephalonoplos Nakai...209 SESE CET aa a Ne 149 638 — sect. Acarna DC. ....... cece ee 218 — sect. Anacantha Ijin. ............... 223 — sect. Apalocentron (Cass.) Kitam... Berna deorc cob ee eh eae 160 — sect. Breea (Less.) Boiss..... ... 209 SSUSUDSE Clee sersteee caer Sete. 164 — abkhasicum (Petrak) Grossh. .... 86 — Acarna Moench.............::ee 218 SHACAUIE AUCLY. eis: -cehscer sees Rea 204 SS bd bey tae teelesonund 201, 203 = sacaule’ (Is) Scop hieesceeseeecoee 205 — — VAT. ACAUIE. 000... ieeeceettetee es 205 — — var. genuina Birq. and Cav... ... Boe ri necnaace aoe tees 205 — — B. sibiricum Ldb. ............ ... 201 — — y. Gmelini DC... oe 201 — adjaricum Somm. and Lev........ 79 — — ssp. nutans Petrak................. 79 — aduncum Fisch. and Mey. ex DC Seetpehentewort Bote oa. ole ete Buse. 97 — — XC. ciliatum (Murr.) Moench.. wan SeaR Rea meteraracadaetten fC a 0 Rin aan 134 =—Jaffine Mausch: scvsc.-twcevene ess 166 — aggregatum Ldb. .......... eee 69 — — var. macrophyllum Alb. ....... 70 — alatum (S. G. Gmel.) Bobr. ... 192 — Albertii Rgl. and Schmalh. ..... 145 — Albertii Wink). .............00008 oe 145 — Albowianum Somm. and Lev....189 — Allionii Sch. and Spenn. ..... ... 205 — anatolicum Petrak. .................... 102 — — var. microcephalum Grossh....... seals ce RRC so te sk Mare oy soca te 102 — — var. ramosum Grossh.......... 102 = apiculatum) DG) ..c8 eee 190 — — ssp. glaberrimum Petrak.... 190 — arachnoideum Boiss. ............... ++. 96 — arachnoideum (MB.) MB.... ... 110 — — var. incanum Lipsky........... 111 — argillosum V. Petr. ex Charadze. ... ssncah Sian SRN NES, cdeetlarmbewue, Dhronek 123 — angunense DC. ...eeeeseeseesrees eee 210 — armenum DC. ...........cccc0sceeeeee eee 200 — ALVENSE AUCL. woececccccccccccccsesees aoe 210 — arvense (L.) Scop............ eee 213 =Varvenser Viethareessseese: eee 21h — — SSP. AIVENSE. .........sccceseeeeee ee 213 — — ssp. Hookerianum Petrak. .. 212 — — ssp. incanum Iljin. .......... ... 212 — — ssp. rubricaule Novopoktr... 213 — — ssp. setosum Ijin. ........... ... 210 — — ssp. vestitum Petrak............ 211 — — var. commune Beck........ ... 214 — — var. commune f. incanum Beck. 20 desl ocd RORY Sete, seve od 211 — — var. horridum Koch. ....... ... 214 — — var. horridum O. and B. Fedtsch. pines go ct Re RRSP See 212 — — var. incanum O. and B. Fedtsch. sae A SME, oo toc codeeh 211 — — var. mite Koch. .................. 214 — — var. obtusilobum Beck... ... 214 — — var. setosum Koch........... ... 214 — — var. setosum Kom. .......... ... 210 — — f. horridum Vietn. ..... 213, 214 == — f. nite Vieniencc see One Le — —c. setosum Schmalh. ....... ... 210 — — Q. horridum Wimm. and Grab... Cirsium esculentum @. sibiricum C. A. M. b) Caulescens C. A. M..... 201 — — B. caucasicum C. A.M. ... 203 — — — a) rhodanthum C. A. M... ... seveanctasw dls NORRIE Sa Leas ee 204 -— euxinum Charadze. ................... 110 — fallax Fisch. and Mey................ 7) — ficifolium Fisch. ............00006 302 — fimbriatum Spreng. ..............0.05 83 — — ssp. Bornmiilleri Petrak. .... 112 — — ssp. tricholoma (Fisch. and Mey.) Petra, cise secustnandsucteentoteedsataee 113 — — var. diversifolium Somm. and Lev. SR, EAs, PR 85 — Fominii Petrak. ............. eee 87 — fraternum DC. ......... eee eee 98 — Frickii Fisch. and Mey........ ... 204 — furiens Griseb. and Schenk..... 132 — Gagnidzei Charadze........... 71, 600 — glabrifolium (Winkl.) O. and B. edits Chive ss. ee cis te 143 — x glabrifolium Petrak. ......... ... 143 — — var. spinosissimum O. and B. Redtschey.. es sec eke 143 — glutinosum Lam. ............:00. 0 163 — Gmelinii Tausch............e 201 — Grossheimii Petrak. .................. 149 — grumosum Fisch. and Mey...... 203 — helenioides (L.) Hill. ............ ... 166 — heterophylloides Pavl........... ... 166 — heterophyllum auct. ................ 166 — heterophyllum (L.) Hill............ 165 — — xC. palustre (1.) Scop. .............. bbe RG Ube FO oh. 166, 184 — heterophyllum ssp. angarense M. Poprniais.n dl Sees See. ae 166 — — var. dealbatum Schmalh. ... 168 — — a. helenioides DC. ............. 165 — — Q. indivisum DC. .............++ 165 — f. dealbatum Lipsky. ......... 168 — Pimeisum DC. vce A UR 165 horridum aucty....26 senna 116 — (Adams) Petrak..........0..... 0... 116 = SD) Ca ie oessrashcessy ese nesin aMn ee 115 — (MB.) MB. ....... ... WSs ITS 2S Stanképr. SR eer: MOMS thee Bye) — ssp. isophyllum Petrak ... ... 118 — ssp. macrocephalum Petrak. .. ... Bi vise Mod heaitd rab aeanays Mowe waa cet ce cactktamie 116 — ssp. prasinolepis Petrak..... 122 — ssp. tomentosum Petrak.......... ... we attattinsacsubebnaien See 115, 118 — var. tomentosum auct. ........ 115 — var. tomentosum Boiss. ...... 118 — B. eriocephalum Boiss. ...... 122 hydrophiloides Charadze. .... ... 186 hydrophylum Boiss. .............. ... 187 hydrophylum auct. ..........c 188 hypoleucum!' DCs eee oe 154 — x C. echinus (MB.) Hand.-Mazz. AEMENORE 2 WNALRTAE ort Gea ETB AS, BRM: 156 — x C. obvallatum (MB.) MB. f. subhypoleucum Petrak. ........ ... 156 — x C. obvallatum (MB.) MB. f. Woronowii Petrak. .................... 156 hypoleucum ssp. drymeium Petrak. egtteaes Meath. SOMA ee AEE EM 3 LSS) — var. ponticum Somm. and Lev. eRRU RPLEIVERE - MORON SR TAN, 154 — B. lazicum C. Koch. ....... ... 154 igniarium Spreng. ..........0008 0 222 imereticum Boiss: ss. 74 — incanum (S. G. Gmel.) Fisch. ex MB. oat ete ARSE 211 — — var. pinnatilobum Sosn. ex Grosshy-wsicccieeve. RO: 211 — interpositum Petrak. .......0..0.0.... 90 — isophyllum Grossh. ............... 118 = italicum DC. 22est..csk wens ee 89 — japonicum DC. ..... ee oe 157 — — ssp. Maackii (Maxim.) Nakai. .. devia ckibandatt rer ipnezscedassonzeaees Ree 157 — — var. amurense Kitam. ..... ... Si iStjucunatm Ijin oe 224 — kamtschaticum Kitam. .......... ... 181 — kamtschaticum Ldb. ex DC.... 179 — — Q. genuinum Herd. ............. 179 — — B. Weyrichii Maxim. ...... ... 181 — Kemulariae Charadze. ................ 75 — Ketzkhovelii Charadze......... ... 120 — Komarovii Schischk ... .... 177, 178 — Kotschyanum Boiss.. ................ 190 — x Kozlowskyi Petrak. ........... ... 199 639 — Kusnetzowianum Somm. and Lev.. spset sasteceds tM ies MEMEO RADE WE, UA 150 — — var. oblongifolium A\lb....... 150 — — var. polycephalum Alb... ... 150 —=MAGleund. AUCE. EIR... 0.2, MER 162 — laevigatum Tausch...............0.0. 210 — lamyroides Tamamsch.............. 138 — lanceolatum auct. ..........0. 135 =i.) MSCOP PO RO. DES oo 135 — — x C. chlorocomos Somm. and Lev. — — ssp. eulanceolatum Beger... 135 — — ssp. hypoleucum (DC.) Beger. .. became cagcaecviexana ie Mietand eRe Ae Ss) — — ssp. Savianum Arénes. ... ... 135 — — ssp. silvaticum Arénes. .. ... 135 — — var. nemorale Naegeli ex Koch. scisbasssussdeciteys MMs ks ER ct 135 — — var. vulgare Naegeli ex Koch... DAR, SENTING, MEIER ND, | OOM oy 135 — — B. hypoleucum DC.............. 135 — laniflorum Grossh. .............0.00.++ 111 — laniflorum (MB.) MB............ ... 111 — — B. Szowitsianum DC........... 105 642 — — y. subnudum DC............. ... 102 — lappaceum auct. 0... 125 — lappaceum (MB.) MB.... 101, 107 — ssp. anatolicum var. microcephalum Petrak. ............ 102 — — ssp. anatolicum var. ramosum Petra. <:.8. S950) PR, | IR as 102 — — ssp. eulappaceum Petrak var. Biebersteinii Petrak. ................. 101 — — var. Biebersteinii Grossh. .. 101 — — var. ferox auct. oo... 125 — — var. lineariloba Trautv. ..... 102 — — var. tomentosum auct........ 104 — — var. tomentosum (Boiss.) Petrak. segeaacibeansar dSvanaseledeatitirisadassasdesooeusers 105 — — f. subracemosum DC..... ... 106 — — f. subracemosum Ldb........ 105 — — Y. macrocephalum Ldb. ..... 106 — — Y. monocephalum Bge........ 125 — — y. tomentosum Boiss.. ....... . 102 — — €. microcephalum Boiss..... 102 — leucopsis Petrak. ............ee 194 — — var. caucasicum Somm. and Lev. cs égueae SRT EEED a. RIE ER, TINE os 193 — — var. indivisa Trautv. ........... 195 — — f. indivisum C. A. M....... 195 — — ¥. cinarescens C. A. M..... 195 — libanoticum Boiss. ............... + 190 — libanoticum DC............... 185, 190 == Wipskyi KOK ste ccccyecsesceencseee ee 621 640 — littorale Maxim. ..........cc 159 — — var. usuriense Rgl.............. 157 — — 6. nudum Rl... 159 — x Lojkae Somm. and Lev.... ... 149 — lokense Conr. and Fr. .............. 107 — longifolium Charadze. ................ 71 — Maackii Maxim. .........c eee 157 — macrobotrys (C. Koch) Boiss. . 106 — — var. longilobum Bordz. ...... 106 — macrocephalum C. A. M......... 122 — macrocephalum Grossh........ ... 116 — maculatum SCOP... ss 227 — Mamiyanum Koidz.................... 182 — megricum Charadze. ................. 104 =} mirum Alpin ve eee ees. eae. 225 — mongolicum Petrak. ................. 177 — munitum (MB.) MB..... ... 116, 117 — — Somm. and Lev... 115 — var. hypopolia Trautv......... 116 — — var. stenophyllum Somm. and Lev. saaanecaeoacastae ahh dhe smaseensae Wnament 1335) =—asimplexnG. vAeiME sean. ae 199 — — XC. obvallatum (MB.) MB. ..... adage stints ates ce A 199 — — ssp. armenum Petrak.. ........ 200 — — var. armenum Grossh. .... ... 200 — — f. armenum Boiss............... 200 — sinuatum (Trautv.) Boiss..... ... 137 — Sommieri Petrak............ eee 69 — sorocephalum DC.........0.::00000 a9, — sorocephalum Fisch. and Mey... 91 — Sosnowskyi Charadze................. 85 — strigosum Fisch. and Mey. ex DC. eee eae eee Oa Eee eae 93 — strigosum (MB.) MB....... .... 94, 96 — var. gymnocalycina Trautv... 93 — f. khorassanicum Boiss... .... 96 subinerme Fisch. and Mey...... 195 — x C. echinus (MB.) Hand.-Mazz. Peasaue aise lan saeevesbicesls) epee oy Se cuasaeaites 149 — — XC. obvallatum (MB.) MB.... 196 — sublaniflorum Sojak ............. ... 112 — X succinctum Ldb.................... . 109 svaneticum Sommi and Lev.... 193 — var. Albowianum (Somm. and Wey.) ChanadzZ@uererecre acer 189 — — var. ramosum Somm. and Lev.. Ee PE Nai ren ee Nc 3 cere eer 193 — sychnosanthum Petrak............. gS — syriacum (L.) Gaertn................ 216 — Szowitsii (C. Koch) Boiss... ... 132 — LALATICUM AUCL... 00... ecrrrneccceeeeeee . 170 — tauricum Sojak.. ............. ee . 114 — tindaicum Charadze. ................. 121 — tomentosum C. A. M.............. . 118 644 641 — tomentosum Grossh. ............006 6+ 115 oe RAMU tVed tedcetet wli22 — tricholoma auct................... . 84, 87 tricholoma Fisch. and Mey ....... 82 — var. aciculare Somm. and Lev... rte Canaan sooner ine Ech RM 84 — X trifurcum Petrak. .......0..... 108 — x tsebeldinum Woron.................. 77 — turkestanicum (Rgl.) Petrak.... 124 — ukranicum Be€SS..............c008 ce 129 — uliginosum Grossh. . .............. . 186 — uliginosum (MB.) MB............. 194 — — B. glabrum Ldb ............. .... 153 — Vlassovianum Fisch............. .... 173 — Vlassovianum Kitam............. ... 174 — — Q. genuinum Herd.......... .... 173 — — B. bracteatum Ldb......... .... 173 — — yY. laciniatum Herd.. ........... 173 — vulgare (Savi) airy-Chow........ 135 = — (Savi) )Petrak. 2 eee 135 — vulgare (Savi) Ten... 6. 135 — — ssp. Savianum Arénes. ......... 39 — ssp. silvaticum (Tausch) Arénes. teadhive yiremanaes akan a ethene Sak oe rn 135 — — SSP. VUIZATE... ...cceccereeeee s WBS — Wachtangii Woron.. ...............- 220) — Waldsteinii Rouy.. «00.0.0... 153 — x Wankelii Reichard............ ... 166 — Weyrichii Maxim. ............00.. 181 — x Woronowii Petrak ............. ... 156 — Wwedenskyi (Petrak) Sojak ...... 79 — X xenogenum Petrak............. ... 134 Cnicus: auctico ee eee 160 == 'Benthis cca ee . 209 Cnius Le cicicGenta are S25 87 — subgen. Breea Sch. Bip. ex Maxim. Hanne eR OEE LL rt 209 == Acarna- li te ees 218 — acaulis Willd.. ...................0000008 . 205 — arachnoideus MB... ..............06+ 110 — arvensis Hoff... ............c0000 w23 —— y. setosus Maxim. .............. 210 — benedictus L. ..............::cccsecsceeees 587 — bracteatus Gilib.. ................0006 . 161 = 'canus MBiikcninseen ee 170 = Wyle, Lee a eee 169 — carthamoides Willd. ................. 311 — centauroides Willd.. ..... .. 311, 312 == Cernuus WO ane ae AE ee 40 <= /ciliatus’ Willds 0). ee: 130 — cynaroides MB...........:.0c1cee 137 — darwasicus Winkl.................... 224 — echinocephalus Willd... ........... 220 = VE FISINQLES eee ee ee 163 642 — esculentus Si€VETS... «0.1.0.0. 202 — fimbriatus MB... «ccc 83 — glabrifolius Winkl... .........0.00 143 — — var. hirtus Winkl... ............ 143 — — var. spinosissimus Winkl ... 143 — Gmelini Spreng. .........ccccce es 202 — heleniodes (L.) Retz............... 5 MOS = Willd, RE ue een. 166 — heterophyllus (L.) Retz........... . 165 =f hornidys) MIB. Veeest. reese seeceteeeee 78 — igniarisu Bemth.. ..........cce 22 — japonicus €. Maackii Maxim.... 157 — — &. schantarensis (Trautv. and Mey) Miaxcitinierss.pescrstsoerenes 159 — jucundus WInKI. ......ccccee oes 224 — kamtschaticum var. alata Trautv... 180 — — Q. genuinus Maxim........... _ ay? = VWaniflonusANiBry eens ese ee 111 == lappaceus MBE weettntt see eee 101 SS iwnunitus MBP eee. e ee 116 — obvallatus MB. «.....-:.1c.0ciee 198 — OJ era GeUs) Nays. cc evsecis tes HG. se 161 == JOVOIISUAIS, NIVGINIGIS ceppocceecsccossecece! coe 183 — — var. B. MB. occ one 191 — pauciflorus Waldst. and Kit. .. 153 — pectinellus (A. Gray) Maxim... 182 — pendulus Maxim. ..........c0cce 208 == pratensis Watniiseerydtsce-scrrepece 161 — rivularis WAIId.. .......seee ee 162 — sairamensis Wink. ................+. 142 — salisburgensis Willd................. 162 — Semenovii Winkl............0-:0+ . 140 — serratuloides Roth.................-. 176 =tserrulatus MIB... ..-:..cc:1--2:--25-se8 128 —s'setosisy) Bess:: (ik. eet ea: 210 SS simple TOT ce eecerscet sec: >» Og — spinosissimus Sievets. ...........:.. 146 SSUStTFISOSUS NIB. ver-:. hates: ereaseee 95 a= Steveigunnctiewetre ees 96 == Isyriaeus (Ie) Wildes -seseeece: 216 — Trautvetteri Wink. .............0..... 123 — uliginosus MB... ......cccccce es 195 == uniflorus Wh p22. Ase be teres 318 Se ="Sievers en ee eee Sapte 311 — Vlassovianus (Fisch.) Maxim...... ... 173 — Weyrichii Maxim. .........:0000 181 Congesta Petrak, subsect................. 90 Cousinia subgen. Oligochaeta (C. Koch) Wink. 1,238 5 a eas Oh 346 — Massalskyi Winkl.. ............0..0 . 347 — Trautvetteri Rgl. .....cccceeee 42 645 Crupina Casswnse. saserdtes. 2a. 252 — crupinastrum (Moris) Vis........ 253 =+(Morisit Bor. \....02.04- ae eee 253 — oligantha Tschern................. ... 255 — pauciflora Kar. and Kir... 254, 255 — VUIZATIS AUCH. .......eeceeseereeeseee es 253 == jvulgaris' Cassg .<..5) Saisie 254 — Zuccarinii Bge. ex Nym.......... 253 Cyaneae Stef. and Georg., subsect. ..... eves chaps dare eaeeseressaal shasta 416 Gyanus Juss) See eee = 0370387 Cyanus (Juss.) Hayek, subgen...... 387 PATS CC beer sec sccceee eee eee 414 ==tsect. Cyanus\DO ye) es 387 — sect. Lepteranthus (Neck.) De.. 449 — sect. Zoegea (L.) DC. ............. 246 — arvensis Monech. ............0.:000+ 417 — jacea (L.) Gaertn. «0... 445 = jsegetum Hill. ......---enceeeeseeseeeee ss 417 — solstitialis (L.) Baumg............. 572 == \Wwulgaris Delanbhe-..2.-......eteer 417 Cynaray Epes ie ee 225 SS (CandunCwls) Ween cers eee 226 SStscolymu ple sbe ts peste. sche 226 Czerniakovskya Czer. subgen. .............. aseceanrtcne sche Eee CERO EUS ECE 418, 609 Demetria Boriss., sect....... ... 300, 607 Depressae Stef. and Georg., subsect. .. vs ee Mens a. cee RA, SRE a 414 HeWenais: @assaece sree ee 52, 139 Echenais (Cass.) Petrak, sect. ...... 139 — carlinoides Cass. .....:cce1cce 148 — — O. and B. Fedtsch............... 146 — — B. nutans DC..... ee oe 148 Echenais nutans Cass.............0....+- 148 — Sieversii Fisch. and Mey......... 146 = Swans edbr yeh 146 Eleutherochaetum Soskov, sect. ........... ate TEE Meee .. 315, 607 BE prirachiy sy Cay ko Chie tees eases ay? Epitrachys DC., sect. ................ 69, 89 — erythrolepis C. Koch. ............... 81 — macrobotrys C. Koch........... ... 106 — ossetica C. Koch. ........... cesses ee 85 — penicillata C. Koch.................. 101 = rigida’ CaO hie. ete wettest 96 — Szowitsti C. Koch. ............... . 133 Erecta Rouy, S€Ct. .........::cceeeeee eee 234 Eremopappus Takht. ............00. 367 — pulchellus (Ldb.) Takht....... ... 369 Eriolepiss@asshes svc... 52, 69, 89 — lanceolata (L.) Cass............ .... 135 Eriophora Petrak, subsect. ............ 126 Eriophora Petrak, subsect. 4. Caucasica Rettalkk.,. 2 cee 32.228. Th RON 100 — 1. Eriocephala Petrak.... 100, 126 — 2. Ferocia Petrak.............05 6 100 ae SH MLOT TUG PUUT AK -eveceercccc ee osnc eee aes pee Bi cee . 100, 114, 118, 124 — 5. Lappacea Petrak. ee LOO OM Erysithales DC., SE€Ct. ...........::0006 + 150 Eucirsium Rouy, subgen. ................ 52 Eucyanus Hayek, sect...... .... 391, 414 — a) Segetales Hayek......... 414, 416 — b) Phrygiae Hayek ............... ... 448 — c) Montanae Hayek. ..................0 tise salrshie Sas . 391, 395, 400, 407 —e) Mrichbeeriaine Hayek. ....... 459 Eujacea, sect. ser. Vulgares_ (Hayek) I ODROCZS eS ck toes caterer cot nana: 444 — a) Vulgares Hayek................. ... 444 Euphalolepis Dobrocz., sect. ........ 551 Eurhaponticum DC., sect . ........... 316 Fischerianae Czer.,; subsect................+. ccat Re cctascstoue Re Vs ea 395, 608 Hornicium,Cass.:...0¢-e08 «n-ne 308 Fornicium (Cass.) Soskov, sect.... 311 — rhaponticoides Cass................. 313 Gallactites ates ar eee: 39 Gerberarauct: eer cents ecee oe 592 GerbernaiCassss eee 589 — § 3, Uechtritzia Beauverd....... 597 — anandria Sch. Bip.................+ 593 — armena Beauverd.................004. 596 — jamesonii Bolus. ...............0.. 0. 590 — Knorringiana B. Fedtsch......... 594 — kokanica Rgl. and Schmalh. ... 597 — Kuntzeana A. Br. and Aschets....... rsdhie ok SINS cnnle Sanaa AR ane 592 — Linnaei Cass. ..........ccccccceceeeee cee 590 — ruficoma Franch. ..............:c006+- 592 — Schimper Sch. Bip. ................ 590 Gerbereae Benth., subtrib. ........... 589 Grossheimia Sosn. and Takht. .... 332 — helenioides (Boiss. and Hausskn.) Sosn. and Takht......... ee 333 — macrocephala (Muss.-Puschk.) Sosn. and) Takhtaei.... 820k)... Ree 333 — macrocephala var. minor. Sosn. and PakN tayacsectetd. ce PRR AAR BORD 333 — ossica (C. Koch) Sosn. and Takht.... 334 646 643 — ossica var. tuba (Somm. and Lev.) Sosn. and Takht.....................5. 334 Halocharis carthamoides MB. ex DC. Se ct eee a tec ee Silt Heterolophus Cass. ............ .. 370, 463 Heterolophus (Cass.) Dobrocz., subgen... msacerence tat Aco Pt Est Dee 463 — sibiricus (L.) Cass. .........068 6 469 Hippophaeslum S. F. Gray... 370, 573 — vulgare S. F. Gray.. ...........0.+- s575 Hyalea (DC.) Jaub. and Spach............ sarvecs cet y bo REF hoa 367, 370, 486 487 369 368 — leuzeoides Jaub. and Spach. ... — pulchella (Ldb.) C. Koch. ....... — tadshicorum (Tzvel.) Sojak. .... Hyalinella (Tzvel.) Tzvel., subgen....... ead Se Sua suseusen Ween es var acum mes a Be a 488 SSSCCHE ch. cccasterteeoeseanbospeverh 2. exteveeestts 489 Hypoleucae (Sosn.) Sosn., sect. ... 424 Iljinia Boriss., sect. ............ .. 296, 606 Infrarosulares Stef. and Georg., SuUbSECtiv ee tote 395, 407 Intermediae Stef. and Georg., subsect. BCE | Oh. A Oma 416 TA CCAM AUC crcese terres eter eovnenes oe 444 SS USS CON eet 370, 440 Jacea (Juss.) Hayek, subgen. ........ 440 a SECM eetecens ccttvstereersncasatieinccsiesees . 444 Jacea (Juss.) DC., sect... ...........0 444 —— © BUjaced BOSS! ceccsceccetcce ee « 444 — communis Delarb.... ..............+++- 445 ——NIGTANSe EGTA. erect ceccee eee: 445 — phrygia ssp. austriaca (Willd.) DOStal eee ee a , . 450 — pratensis Lam... .........:::ccceeee 445 — segetum (Hill.) Lam... ............. . 417 — vulgaris ssp. angustifolia (Schrank) DOStal en ieritccterccetetececess . 446 — — SSP. VUIQATIS.... cscs 445 Jurinea Korolkowii Rgl. and Schmalh. Rs i enna Reis A srt Te 349 — Paulseni O. Hoffm. ex Pauls.......... AE ee) DaRAC EES, Senet 287 = "“lenttis’ BRE eeeeisccs oe os 304 Kentrophyllum Neck... ..........0.0006 581 Kentrophyllum (Neck.) Fiori and Paol., SECU iicnssencceiirerecsrebavenieseeersvces . 583 — anatolicum Fisch. and Mey. ex DC. ee LN LN TA NAL BE el Snes 584 — glaucum Fisch. and Mey. ex DC... Le I Oe ae ED Earned er 584 644 — lanatum DC... voccccccccccccccccessceeees . 583 ——ttauricum CUA. Mit: nse. nes 583 Klaséay Cassty ns ike Arcachon: ee Klasea (Cass.) DC., sect. .......... ... 272 — centauroides (L.) Cass... ......... 273 — heterophylla (L.) Cass. ........ ... 280 — quinquefolia (MB.) Cass..... ... 294 Lamyra’' Cass... cc... ae 219 — angustifolia Cass............... Bhi 220 — casabone (L.) Tamamsch......... 219 — echinocephala (Willd.) Tamamsch. ust ate adds, Be Sete, cote ee 220 Lamyropappus Knorr. and Tamamsch... Lieedte cee Solna eee ee 221 — schakaptaricus (B. Fedtsch.) Knorr. and Tamamsch... ............:csee 221 Lamyropsis Charadze, sect.... 136, 603 Lanceolate Petrak, subsect........... . 134 Lanceolata Petrak, subsect. 2. alata PEM AK 2 ope aceesees Meee ee 135 Leibnitzia Cass... 0.0.0. 592 — anandria (L.) Turcz... ........... 593 — cryptogama CaAs6.... ...ccccscee 593 — Knorringiana (B. Fedtsch.) Pobed... een eh Apis eaten tan Aen eed 594 — Kuntzeana (A. Br. and Aschers.) RODEO nn reresicersraueeer etree 592 — phaenogama Cass.. ..........01 S28) — ruficoma (Franch.) Pobed........ 592 Lepteranthus Neck. ... .. 370, 440, 449 Lepteranthus (Neck.) DC., sect.... 449 Lerieae Less., subtrib ................ ... 589 Leucantha S. F. Gray........ .. 370, 570 — cyanifolia S. F. Gray.. ............ Sy — iberica (Trev.) A. and D. Léve... 574 — solstitialis (L.) A. and D. Léve... 572 Leucophyllae (Sosn.) Sosn., sect... 428 Weuzed DC vice eee cccssccse-necases eaecns oh 309 Leuzea sect. Fornicium (Cass.) DC... . Ea at ert pO MR a NN nee ata 311 — altaica Fisch. ex Schauet........ 311 Sep ATI Ks waevte ee a soeecs enero ne ned 313 — carthamoides DC. ..........:.c00000 6 311 — — ssp. orientalis Serg......... ... 312 — AAUTICA BRE... ..ccccccecccercescservercens . 318 — salina Rgl. ex O. Fedtsch. ...... 315 = = BSYOTKETN ED enc dronenoerecr codaoertoneceosae ic 313 — serratuloides Fisch. and Mey... 300 Leuzeopsis Boriss., sect. .. ... 298, 607 Lophiolepis Cass. ............... 52, 69, 89 = SUD BOM crc teats stecs less phasgd ee manne ynt 69 SMSC Cbs crasievs chtactone toot rers arenas 69, 128 — calocephala Cass .......:csccee vee 130 == idubiasC@assie a eee 135 Lopholoma Cass .........000005 «0+ 370, 493 Lopholoma (Cass.) Dobrocz., subgen. .. eISBN ead i et ewe OL 493 SS isectais Ak dee ae, 503 Malacocephalus altaicus Tausch... 313 Malacolepis Ldb., sect. ................ +. 40 Mariana Lactea Hill. ................... pnd) — Mariana (L.) Hill... 0... 227 Mastrucium pinnatifidum Cass... . 268 Mesocentron Cas6.... .........00 370, 570 Microcephala Petrak, subsect.. ...... 92 MicrolonchuS auct. .........cccccesccee oe 346 — albispinus Bge. 0.0.0... . 352 — minimus BOISG... ......c.cccceccceeees . 349 — oligochaetus BOiss.. ............0.... 347 Microlophopsis plumosa (Aitch. and Hemsl!)'Czercet eee 307 Microlophus Cass.... .........0.+. 370, 560 Microlophus (Cass.) Hayek, subgen.... sacra ilk resp g ny Aaa aac 560 == SEC... «2: Meee, 2 NT? eh 561 — alatus (Lam.) Cass. ...... ... 306, 562 — behen (L.) Takht......... . 562 — polypodiifolius (Boiss.) Agadsh... .. Par ern tna totter en 563 — Szovitsianus (Boiss.) Takht... Aiea basta nite atiaeueek oar ol ecmteamas 562, 563 Modestia Charadze and Tamamsch. ... cerita tee sos Ssanesngisuseanc saan Mec eeate 223 — darwasica (Winkl.) Charadze... 224 — jucunda Charadze and Tamamsch... AO tik 10h: Webb ats tae, Mae 224 — mira (lIljin) Charadze and Maram hisses eeccee vos. ceenccceacances 225 Montana Charadze, subsect... 150, 604 Montanae (Hayek) Stef. and Georg., SUBSE Chergeteican Mesite nt tescasreet aoe: 391 Mutisiées Baill., trib... ................0- 589 Mutisiaceae Bessey, fam.. ........... . 589 Sessa, tribes scsi cc ddaceh scapasacereee . 588 Mutisieae Cass., trib.. ..............0005 . 588 Mutisieae Less., subtrib... ............. 589 Mutisinae O. Hoffm., subtrib....... 589 Napuliferae Hayek, gr.. ...........00 . 391 Nassauvieae, Benth., subtrib........ 589 Notobasis Cass. ...........ccccceceeere es 215 — syriaca (L.) Cass... 0... eens 216 Odontolophopsis Tzvel., subgen... 485 647 Odontolophus Cass... ............ 370, 481 Odontolophus (Cass.) Hayek, subgen... Ree oer le MALIA htc Mec 481 — cyanoides CASS... ....:ccee . 483 Olgaeallhpinwe 2. ee a. 43 — baldshuanica (Winkl.) Iljin.. ..... 50 — eriocephala (Winkl.) Iljin.......... 44 — lanipes (Winkl.) Iljin.. ........0..... 47 — longifolia (Winkl.) Ijin... ............ 47 — nidulans (Rupr.) [jin. ...0...... 48 — nivea (Winkl.) Ijin... .........04.. 45 =" pectinata) Ijineys.. 2S eee ee 50 — Roborovskyi Ijin... ......... ee 46 — spinifera Ijin... oo... eee 49 — Wvedenskyi Ijin... . 46 Oligochaeta C. Koch... ................ . 346 — divaricata (Fisch. and Mey.) C. Koch....... Be er oy Cy ce Pere eee 347 — leucosmerinx Rech. f. and KGie... 353 — minima (Boiss.) Brig... ............ 349 — — f. lyratisecta Czer... 0.0.0.0... 350 =—itomentosa’ @Zen.2.4/...au ee 348 Onopordum L. ... ......... ee 228 MSE Clie ets, ee keh, RR 231 — acanthifolium Gilib.. ........... ... 231 — — ssp. araneoso-tomentosum Rech. BAM J RR A RN Bch auto 231 =—"acanthiwmm. Res.. Peteas beter: 231 — — xX O. tauricum Willd... ....... 232 — anisacanthum BOISS.. ........66 239 — armenum Grossh ....... eee vee 234 == "Bra cteatuimneiacectcss-aee. tonne 229 — candidum Nabelek.. ................. . 238 — carmanicum Bornm. ...............-: 239 -— cinereum Grossh. ..........:ccc 232 — deltoides Ait. .....ccccccccsceeeee es 302 — elatum Sibth. and Smith. ........ 233 — Frickii Tamamsch. ........ .. 236, 605 — heteracanthum C. A. M........... 239 — — var. ortholepis Bornm........ 237 — — B. anisacanthum Boiss... ... 239 — lanigerum BOISS...........0.0 5 PBv =—'leptolépis DG... 28... 1i23)7 — — x O. acanthium L. ............... 238 = "Ole ae) Ree Ree aut, > 23h — polycephalum Boiss... .............. 239 — Prjachinii Tamamsch... ... 233, 605 — seravschanicum Tamamsch... .......... zest tein) ARIE, ASME 238, 605 — tauricum Willd... 0.0.00... 233 ae VEEN DC eicc Hee. Beal ans . 233 — — PB. tauricum DC........0000... W238 — viscosum HOrm.......0..cccccccceee . 233 Onoserideae Benth., subtrib....... ... 589 Onotrophe Cass.. ........::eeee . 52, 149 — (Cass.) DC., sect... ... 90, 149, 172 — sect. Apalocentron Cass... 150, 160 — sect. Microcentron Cass. ......... 150 WIC AUIISH CASS i... f8te ROR 205 — palustris CasS.........:ccceeceeseee . 183 Orientales (Hayek) Tzvel., sect.... 500 Oroboidea Ijin, sect. ...............006 306 Orthocentron Cass... ...... . 52, 150, 184 Orthocentrum DC., sect... ............ 150 Orthocentrum (Cass.) Charadze, subs... Ee peeled cal AMOR Ne shat onaaceancaseeceses see 184 Perdicieae Link, ordo.. ................ . 588 Perdicium anadria R. Bvr.... .......... 593 — tomentosum Thunb ............... .. 593 Phaecopappus MDE). niveccsssse.ctire sees 359 — sect. Cheirolepis (Boiss.) Boiss. § 1. Eucheirolepidei Boiss. ............. 356 — sect. Euphaeopappus Boiss. § 3. Macrocephali Boiss ............. ..- 359 — sect. Euphaeopappus § 4. Micro- cephali BOISS .......cccecceciee oe 477 — sect. Psephelloides Jaub. and Spach.. LUN congue Pees ANN) eel fa 485 — sect. Rhaponticoides Jaub. and Spachiccdisccisss.vsrne Rh. BRS 359 — amblyolepis (Ldb.) Boiss.. ...... 479 B. daghestanicus (Trautv.) Ten Shey se sic ech ee ee Reenass 479 — armenus Jaub. and Spach.. ..... 361 — Aucheri 8. integrifolius (Boiss.) Bonnin te. 2G) aaa . 362 — — y. Szovitsii (Boiss.) Bornm... 363 — cheirolophus Boiss... ...........++ 358 — daghestanicus Lipsky............... 479 — — var. laceratus Lipsky.. ........ 480 — daralaghezicus Fom. ................ 364 — Freynii Hausskn. and Sink... .. 300 — inuloides Fisch. ex Nym. ........ 580 — Kotschyi B. persicus (Boiss.) Boiss... SEU Nae MEM I cee ite. eos 358 — leuzeoides BOISS...........0:0005 0 362 — — B. integrifolius Boiss... ....... 362 — — y. Szovitsii (Boiss.) Boiss... 363 — macracanthus Boiss. and Buhse .... SES Peedi xt aE Mahle RR SY he 363 — macrocephalus (Muss.-Puschk.) BOiSS 5.5 334 S="OSSICUSiIC. NIKO A te.cee teeter 334 — Ruprechtii BOiISS. ...........::0e 480 — — B. samurensis Lipsky. ........ 479 — scleroblepharus Freyn.............. 363 648 646 — spectabilis (Fisch. and Mey.) Boiss. MRE ee rec sn busnestteges Mees BAG) — Stevenii (MB.) Boiss................ 564 — SZovitsii BOISS. .........2:..0ec.e00 2 363 — trinervius (Steph.) Boiss. ........ 483 Phalacrachena Iljin...................... 579 — calva (Ldb.) Ijin. «0.0... 581 — inuloides (Fisch. ex Schmalh.) Iljin. TE SR, eee ath eRE seanhs hey 580 Phalolepis Cass. .......... 340, 370, 538 Phalolepis (Cass.) Dobrocz., subgen... Bee) esate br bah 5 abe teh BRR 538 Phalolepis (Cass.) DC., sect. ... ... 444 Phrygia S.F. Gray. .... .. 370, 440, 449 Phyllocephala (Petrak) Charadze, subsect) S4.25 G2... ae 99 Picnomon adans. .................::0 218 = Sfacarna’ (L.)sCass Pee nese: 218 Piptoceras Cass. ...........00. + 370, 560 — behen (L.) Cass............ . 306, 562 Piptochaete Boiss., sect. ................ 269 Plagiobasis#auct:, 2.2::eese- ee 250 Plagiobasis Schrenk. ..................... 255 — centauroides Schrenk. .............. 256 — dshungaricus IMljin................... 256 — sogdiana BE. ...........scsceseeee es 252 Platycophali Rouy, sect. .......... 10, 20 Platylophus Cass. .......0.00. +. 370, 440 Platyraphium Cass. ......:.:c1:cceee es 219